Proceedings of the International Symposium

on Air Pollution and Climate Change Effects

on Forest Ecosystems

February 5-9, 1996 Riverside, California

United States

Department

of Agriculture

Forest Service

Pacific Southwest

Research Station

General Technical Report

PSW-GTR-166

3000

2000

1000

2

4

6

8

10

4

6

8

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12

14

2

South-North(km)

West-East(km)

Study Site

Silverwood

Lake Gregory Lake

Arrowhead

Holcomb Greek

Big Bear Lake

Baldwin

Lake (dry)

Bear 

River

1212 m

Los Angeles

Santa Ana

SantaRiver

Ana

Creek

Lake

1212 m 1212 m

San Bernardino 1212 m

Mountains

California

Kilometers

Study Area

Lake

National Forest boundary

1212 m Contour

(a)

(b) Barton Flats Area

San Bernardino Mountains

Barton Flats Area
Abstract

 Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan L., technical coordinators.

1998. Proceedings of the international symposium on air pollution and climate

change effects on forest ecosystems; 1996 February 5-9; Riverside, CA. Gen. Tech.

Rep. PSW-GTR-166. Albany, CA: Pacific Southwest Research Station, Forest Service,

U.S. Department of Agriculture; 332 p.

 The purpose of the International Symposium on Air Pollution and Climate Change Effects

on Forest Ecosystems held February 5-9, 1996 in Riverside, California, was to summarize

results of several years of scientific collaboration between the United States and European

specialists about air pollution and climate change effects in Central European and North

American forests. More than 40 technical presentations given by scientists and forest

managers from the United States and Central Europe were divided into 5 sessions

discussing mechanisms of air pollution and climate change effects on plants; air pollution

status of forests; results from several gradient and case studies on the effects of air

pollution/climate change on forests; information on forest health in several Central

European countries; and information regarding photosynthetic active radiation

measurements and their use in forest health monitoring.

Retrieval Terms: effects on forests, monitoring, nitrogen deposition, ozone, sulfur dioxide

Technical Coordinators

Andrzej Bytnerowicz is Ecologist, Michael J. Arbaugh is Statistician, and Susan L.

Schilling is Computer Programmer with the Atmospheric Deposition Effects in Western

Forest Ecosystems Research Unit at the Pacific Southwest Research Station, USDA Forest

Service, 4955 Canyon Crest Dr., Riverside, CA 92507.

Publisher:

Albany, California

Mailing address:

PO Box 245, Berkeley CA

94701-0245

(510) 559-6300

http://www.psw.fs.fed.us

September 1998

Pacific Southwest Research Station

Forest Service

U.S. Department of Agriculture
Proceedings of the International Symposium

on Air Pollution and Climate Change Effects

on Forest Ecosystems

February 5-9, 1996 Riverside, California

Andrzej Bytnerowicz Michael J. Arbaugh Susan L. Schilling

Technical Coordinators

Pacific Southwest

Research Station

USDA Forest Service

General Technical

Report

PSW-GTR-166

September 1998

i

Contents

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv

SESSION 1: Air Pollution/Climate Change Effects on

Plants: General Topics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

William J. Manning and Stefan Godzik, Chairs

Biochemical Reactions of Ozone in Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

J. Brian Mudd

Oxidant Induced Alteration of Carbohydrate Production

and Allocation in Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Robert L. Heath

The Use of Plants as Bioindicators of Ozone . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

William J. Manning

State of Polish Mountain Forests: Past, Present, and Future . . . . . . . . . . . . . . . . 27

Krystyna Grodzinska and Grazyna Szarek-Lukaszewska

Ozone Injury Responses of Ponderosa and Jeffrey Pine in the Sierra Nevada

and San Bernardino Mountains in California . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

Paul Miller, Raleigh Guthrey, Susan Schilling, and John Carroll

SESSION II: Air Pollution Status of Forest Sites . . . . . . . . . . . . . . . . . . . . . 43

Paul Miller and Juri Martin, Chairs

Ozone Air Pollution in the Ukrainian Carpathian Mountains and Kiev Region. . 45

Oleg Blum, Andrzej Bytnerowicz, William Manning, and Ludmila Popovicheva

Ozone Monitoring in the Krakow Province, Southern Poland . . . . . . . . . . . . . . . 51

Barbara Godzik

Ozone in the Upper Silesia Region—Concentration and Effects on Plants . . . . . 59

Stefan Godzik

Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site in

Early Summer of 1994 and 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

Evgeny Donev, Karl Zeller, and Christo Bojinov

Ozone and Modeled Stomatal Conductance at a High Elevation

Subalpine Site in Southeastern Wyoming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

Robert C. Musselman, Karl F. Zeller, and Nedialko T. Nikolov

Bulgarian Rila Mountain Forest Ecosystems Study Site: Site

Description and SO4

2-, NO3-

 Deposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Karl Zeller, Christo Bojinov, Evgeny Donev, and Nedialko Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. ii

Fire in Siberian Boreal Forests—Implications for Global Climate

and Air Quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Susan G. Conard and Eduard P. Davidenko

Nutrient Fluxes in Forests of the Eastern Sierra Nevada: Comparisons

with Humid Forest Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

Dale W. Johnson, Richard B. Susfalk, Randy A. Dalgren, Virginia Boucher, and

Andrzej Bytnerowicz

Concentrations and Deposition of Nitrogenous Air Pollutants

in a Ponderosa/Jeffrey Pine Canopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

Andrzej Bytnerowicz, Mark E. Fenn, and Michael J. Arbaugh

Predicting Nitrogen Flux Along a Vertical Canopy Gradient in a Mixed Conifer

Forest Stand of the San Bernardino Mountains in California . . . . . . . . . . . . . . 115

Michael J. Arbaugh, Andrzej Bytnerowicz, and Mark E. Fenn

Indicators of Nitrogen Status in California Forests . . . . . . . . . . . . . . . . . . . . . . . . 123

Mark E. Fenn and Mark A. Poth

Nitrogen Deposition Effects on Coastal Sage Vegetation of

Southern California . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131

Edith B. Allen, Pamela E. Padgett, Andrzej Bytnerowicz, and Richard Minnich

SESSION III: Air Pollution/Climate Change Effects on Forests: Results

of Gradient and Case Studies . . . . . . . . . . . . . . . . . . . . . . . . . . 141

Reginald Noble and Alicja Breymeyer, Chairs

Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East and

Along a Pollution Gradient in Poland: General Assumptions . . . . . . . . . . . . . . 143

Alicja Breymeyer

Geographical Differentiation of the Floristic Composition and Structure

of the Herb Layer of Forest Permanent Plots in East Germany,

Poland, and Belarus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

Ewa Roo-Zielinska and Jerzy Solon

Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions of

Forest Permanent Plots in Poland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Ewa Roo-Zielinska and Jerzy Solon

Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the

Content of Heavy Metals in the Pedosphere in Poland . . . . . . . . . . . . . . . . . . . 169

Marek Degorski

Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical

Properties of Upper Soil Layers in Polish Forests . . . . . . . . . . . . . . . . . . . . . . . 179

Alicja Breymeyer, Marek Degorski, and David Reed

Geographic Information System (GIS) Analysis of Ecosystem Response to Industrial

Pollution in the Niepolomice Forest in Southern Poland . . . . . . . . . . . . . . . . . 187

January Weiner, Stefan Fredro-Boniecki, David Reed, Ann Maclean,

Marshall Strong, and Michael Hyslop

Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine

Trees Affected by Air Pollutants in Polish Forests . . . . . . . . . . . . . . . . . . . . . . 197

Wojciech Dmuchowski, Ewa U. Kurczynska, and Wieslaw Wloch

Ectomycorrhizae of Young and Mature Scots Pine Trees in

Industrial Regions in Poland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205

Barbara Kieliszewska-Rokicka, Maria Rudawska, and Tomasz Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. iii

Impact of Urban Environmental Pollution on Growth, Leaf Damage and Chemical

Constituents of Warsaw Urban Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215

Waldemar Chmielewski, Wojciech Dmuchowski, and Stanislaw Suplat

Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa

and Jeffrey Pines at a Southern California Site . . . . . . . . . . . . . . . . . . . . . . . . . . 221

Patrick J. Temple and Paul R. Miller

Effect of Ozone Exposure on Seasonal Gas Exchange of

Five Western Conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229

Nancy E. Grulke, Paul R. Miller, and Theodor D. Leininger

Mature Ponderosa Pine Nutrient Use and Allocation Responses

to Air Pollution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239

Mark A. Poth and Mark E. Fenn

SESSION IV: Forest Health Monitoring: National Perspectives . . . . . . . 249

Paul Miller and Oleg Blum, Chairs

Assessment of Forest Quality in Southwestern Poland with

the Use of Remotely Sensed Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251

Zbigniew Bochenek, Andrzej Ciolkosz, and Maria Iracka

Response of Scots Pine Stand Vitality to Changes in Environmental

Factors in Poland, 1991-1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

Jerzy Wawrzoniak

Air Pollution and Forest Health Studies along a

South-North Transect in Poland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259

Stefan Godzik, Jerzy Szdzuj, Tomasz Staszewski, and Wlodzimierz Lukasik

Forest Health Status in Russia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

Vladislav A. Alexeyev

Air Pollution and Climate Change Effects on Health of the

Ukrainian Forests: Monitoring and Evaluation . . . . . . . . . . . . . . . . . . . . . . . . . . 271

Igor F. Buksha, Valentina L. Meshkova, Oleg M. Radchenko, and Alexander S. Sidorov

Forest Condition in Latvia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281

Madis Sipols

Forest Health Monitoring and Forestry Implications in the Czech Republic. . . . 287

Martin Cerny and Pavel Moravcik

Dynamics of Forest Health Status in Slovakia from 1987 to 1994 . . . . . . . . . . . . 293

Julius Oszlanyi

Forest Health Status in Hungary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299

András Szepesi

Forest Health Monitoring Indicators and Their Interpretability:

A Lithuanian Case Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313

Romualdas Juknys and Algirdas Augustaitis

SESSION V: Photosynthetically Active Radiation (PAR) Measurements

and Their Use in Forest Health Monitoring . . . . . . . . . . . . . 321

Craig Palmer, Chair

Photosynthetically Active Radiation Measurements in Pure Pine and Mixed

Pine Forests in Poland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323

Jaroslaw Smialkowski

Remote Sensing Applied to Resource Management . . . . . . . . . . . . . . . . . . . . . . . 327

Henry M. Lachowski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. iv

Preface Industrial air pollution has been identified as one of the primary causes of

severe damage to forests of central Europe in the past 30 to 40 years. The

mountain forest ecosystems have been affected considerably, resulting in

extensive areas of severely deteriorated forest stands (e.g., the Krusne Hory of

the Czech Republic or the Izerske and Sudety Mountains along the Polish and

Czech border). In addition, the increase of motor vehicles in central Europe has

caused a higher amount of nitrogen oxide and hydrocarbon emissions. Under

particular weather conditions (high solar radiation, high temperatures,

temperature inversions), these compounds may undergo complex chemical

transformations resulting in the formation of photochemical smog and a buildup of potentially phytotoxic ozone concentrations. Ozone has already been

responsible for injury to ponderosa pine stands in the San Bernardino Mountains

in southern California and to forest vegetation in the eastern United States and

western Europe. Elevated concentrations of nitrogenous components in

photochemical smog and the emissions of ammonia from agricultural activities

have caused eutrophication of natural ecosystems, including forests, both in

Europe and North America. In addition, changing climatic factors, especially the

amount of precipitation, temperature and solar radiation (including ultraviolet-B),

modify responses of plants to air pollution, and must be considered when these

effects are evaluated.

To successfully prevent additional deterioration of forest health both in

central Europe and in the United States, well-coordinated research activities are

needed in the areas of air pollution and forest health monitoring, investigations

of the effects of air pollution and climate change on forest functioning, and

selection and implementation of proper response strategies. These studies may

result in stricter air pollution standards in the forested areas and more effective

control strategies. Reduction of air pollution and the simultaneous improvement

in understanding forest functioning and the application of research results should

help prevent future forest decline and rebuild the already damaged forests.

Cleaner air also results in the preservation of valuable biological resources and

healthier living conditions of people.

Since 1991, U.S. agencies such as the USDA Forest Service (FS), USDA

International Cooperation and Development (ICD), and U.S. Environmental

Protection Agency (EPA), and the European institutions such as the Polish

Academy of Sciences have sponsored a collaborative scientific program entitled

“The Effects of Atmospheric Deposition and Climate Change on Forest

Ecosystems in Central and Eastern Europe and the United States.” This program

has resulted in several cooperative studies between U.S. and Polish scientists,

introduced the European cooperators to the forest health monitoring procedures

used in the U.S., and provided training in using plants as bioindicators of air

pollution phytotoxicity. Other cooperative studies, sponsored mainly by the FS,

ICD, and European institutions, have been conducted in Bulgaria, Poland, the

Czech Republic, and Ukraine. Thanks to the financial support provided by the

U.S. State Department, FS, and EPA, and governments of the European countries,

the European participants have received advanced training in the U.S. Forest

Health Monitoring methodologies, and selected countries have implemented

this system. In addition, funds received from the USDA FS Pacific Global Change

Research Program and the Southern California Edison Company made possible

collaborative research between Polish, U.S., and Canadian scientists on the effects

of elevated levels of nitric acid vapor (an important component of photochemical

smog) on California tree species.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

v

Partial presentation of the developing plans for this cooperative research

and some of the preliminary results were presented at the international

conference “Climate and Atmospheric Deposition Studies in Forests” organized

by the Polish Academy of Sciences and the USDA Forest Service in October 1992

in Nieborow, Poland. In addition, at the Nieborow meeting, Ann Bartuska of the

FS suggested that the Pacific Southwest Research Station, USDA FS organize an

international symposium in Riverside, California in 1995/1996. The purpose of

such a meeting would be to present and discuss the continuing progress of the

U.S.-European research cooperation.

Thanks to the financial support of the USDA ICD, the USDA FS, and a grant

from the Southern California Edison, the International Symposium on Air

Pollution and Climate Change Effects of Forest Ecosystem was held in the

recently renovated historic Mission Inn in downtown Riverside on February 5-9,

1996. The primary goal of the meeting was to discuss the effects of air pollution

and climate change and modification of these effects by other abiotic and biotic

factors on forest ecosystems. Seventy scientists, forest managers, and decision makers

from Belarus, the Czech Republic, Estonia, Hungary, Latvia, Lithuania, Poland,

Russia, Slovakia, Ukraine and the United States participated in the meeting.

The participants of the symposium were greeted by the Mayor of the City of

Riverside, Ronald Loveridge, who is also a member of the South Coast Air

Quality Management District, and James Space, Director of the Pacific Southwest

Research Station, USDA Forest Service. In his opening remarks James Space

stated, “The issues of global climate change and human-made air pollution are

closely interwined and have been the subject of major inter-governmental

research programs, both locally and worldwide. The implications are vital to the

long-term sustainability of our forest ecosystems.” William T. Sommers, Director

of the Forest Fire and Atmospheric Sciences Research Staff, USDA Forest Service,

outlined the domestic and international activities of the FS related to air pollution

and climate change effects on forests and the U.S. forest health monitoring

activities. Whetten Reed, Director of the USDA International Cooperation and

Development, discussed international cooperative research sponsored by the

USDA in central and eastern Europe.

 These introductory speeches were followed by five scientific sessions in

which more than 40 papers by European and U.S. forest scientists and managers

were presented. Session 1 included the mechanisms of the effects of oxidant air

pollutants on plants, the use of plants as bioindicators of air pollutants, and the

effects of air pollution and climatic factors on mountain forests of southern

Poland and the San Bernardino Mountains and Sierra Nevada of California. At

Session 2 papers were presented on air pollution status in European and U.S.

forest sites, effects of fire on the Siberian boreal forests, and deposition of

nitrogen air pollutants to forests and its effects on the health and ecology of

natural ecosystems. At Session 3 the results of case studies addressing air

pollution and climate change on forests, both in Europe and the United States,

were discussed. Session 4 included information on the status of forests in the

individual countries, based on the results from the Forest Health Monitoring

networks. Session 5 dealt with measurements of photosynthetic active radiation

(PAR) and its use for forest health monitoring efforts; and a video, “Integrating

Remote Sensing with GIS,” produced by the USDA FS, was shown. In general,

the meeting provided an update on forest health and air pollution status in

forests, mechanisms of air pollution and climate change on forest vegetation, and

an evaluation of the possible ecological effects of air pollution and climatic

factors in forests of the represented countries.

After 2 days of sessions, three field trips were offered to the symposium

participants: (1) in the San Bernardino National Forest and Joshua Tree National
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. vi

Park in California, the air pollution monitoring efforts, effects of air pollution on

natural resources of forest and desert ecosystems, and programs that involve

school children in nature preservation were presented; (2) at the Harry Reid

Center for Environmental Studies, University of Nevada, Las Vegas, software used

for forest health monitoring activities in the United States and some of the central

European countries was demonstrated, and the participants received hands-on

training of the presented methodologies; and (3) at the Pacific Southwest Research

Station, USDA FS in Riverside visitors were acquainted with scientific activities

related to the effects of air pollution and climate change on forests, various

aspects of research and management related to fire presence in forests,

meteorology and fire severity forecasting, and wildland recreation and the urban

culture research, and visited the Botanic Garden of the University of California,

Riverside and natural history collections of the Riverside Municipal Museum.

Future research cooperation between the U.S. researchers and their European

colleagues may include climatic gradient studies in Poland, Belarus, and Russia

or cooperation between the Czech Republic, Poland, Slovakia, Ukraine and the

United States on ozone distribution studies and phytotoxicity in forests of the

Carpathian Mountains. In addition, a similar meeting may be organized in a few

of years in one of the European countries to discuss and evaluate progress of the

ongoing cooperative work.

The Organizing Committee for the Symposium on February 5-9, 1996

consisted of Alicja Breymeyer, Polish Academy of Sciences; Andrzej Bytnerowicz,

USDA Forest Service; William Manning, University of Massachusetts; and

Reginald Noble, Bowling Green State University. The Local Arrangements

Committee consisted of Michael Arbaugh, Linda Burkholder, Andrzej

Bytnerowicz, Paul Miller, Mark Poth and Susan Schilling, Pacific Southwest

Research Station, USDA Forest Service.

Andrzej Bytnerowicz,

Technical Coordinator
SESSION I

Air Pollution/Climate Change

Effects on Plants: General Topics

William J. Manning and Stefan Godzik, Chairs

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 3

Biochemical Reactions of Ozone in Plants1

Abstract

Plants react biochemically to ozone in three phases: with constitutive chemicals in the apoplastic fluid and cell

membranes; by forming messenger molecules by the affected constitutive materials (ethylene); and by responding to

the messenger molecules with pathogenic RNAs and proteins. For instance, plant reactions with ozone result in

constitutive molecules such as the ozonolysis of ethylene in the gas spaces of the leaf, and the reaction with ascorbic

acid in the apoplastic fluid. Formation of messenger molecules include the stimulation of ethylene production.

Responses to the messenger molecules include the formation of the pathogen related proteins and their mRNAs.

Reactions of ozone with biological molecules also frequently result in classical ozonolysis of double bonds, with the

production of various aldehydes and peroxides, such as ethylene, isoprene, fatty acids, tryptophan, and some

phenylpropenoic acids. Some reactions of ozone with biological molecules do not fit the classical ozonolysis

mechanism, such as the oxidation of methionine and some phenylpropenoic acids.

Introduction

Although ozone has toxic effects on both plants and animals, the mechanisms by

which the toxic effects are elicited are inadequately understood. However, plants

react biochemically to ozone in three phases: with constitutive cells and materials;

by generating messenger molecules; and by responding to the messenger

molecules.

Reactions with constitutive cells and materials result in substances in the

gas phase, (e.g., ethylene) that are dissolved in the apoplastic fluid, (e.g.,

ascorbic acid) and that react with the lipids and proteins of the plasma

membrane of cells surrounding the sub-stomatal cavity. The concentration of

ozone in the leaf spaces inside the leaf is zero (Laisk and others 1969). This

indicates that ozone passing through the stomata is immediately consumed by

oxidizable substances lining the leaf spaces. Because lesions characteristic of

ozone damage occur in the palisade parenchyma, palisade cells are part of the

sink for ozone.

Reactions of messenger molecules cause stimulation of the synthesis of

molecules such as ethylene, while the response of messenger molecules results in

the formation of messenger RNA and the proteins for which they provide the

genetic code. However, because these three stages of response to ozone are simply

an analysis of the problem, a practical method of protecting plants needs to be

developed.

This paper discusses the biochemical reactions of plants to ozone, focusing on

the reactions of constitutive cell materials, the synthesis of messenger molecules,

the responses of the messenger molecules, and various methods to protect plants

from ozone damage.

Discussion

Ozone Reaction with Constitutive Materials

Ethylene

Ethylene has been known as a “ripening hormone.” The arrival of the ripening

is coincident with a “climacteric” in ethylene production. In addition to this

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor Emeritus, Department of

Botany, University of California, Riverside, CA 92521, U.S.A. email:

Mudd@citrus.ucr.edu

J. Brian Mudd2
4 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

function in normal development, ethylene is known to be produced under

various forms of stress. The biosynthesis of ethylene has posed a problem that

has consumed many years of research, but it is now clearly understood as the

following function:

S-adenosyl-methionine ----> ACC (1-aminocyclopropane-1-carboxylate)

ACC -----> C2H2

The two critical enzymes are ACC synthase (which is the rate-limiting enzyme),

and ethylene synthetase.

The reaction of ozone with ethylene in the gas phase involves a classical

ozonolysis (fig. 1). The degradation products include formaldehyde,

hydroxymethylhydroperoxide (HMHP), and hydrogen peroxide. Mehlhorn and

Wellburn (1990) and Hewitt and Kok (1991) have suggested that ozone toxicity is

a consequence of the reaction of ozone with ethylene. Mehlhorn and others (1990)

attribute radicals detected after exposure of plants to ozone to the reaction of ozone

with ethylene. Are these reactions taking place in the atmosphere? Hellpointer and

Gab (1989) measured the peroxide content in rainwater collected on different days

(table 1). The major peroxide usually found was hydrogen peroxide. Small amounts

of HMHP were also detected. These measurements show that the reaction of ozone

with ethylene in the atmosphere is a clear possibility. In the gas phase reaction it

was found that hydroxyl radicals could be detected by the conversion of

cyclohexane to cyclohexanol and cyclohexanone (Atkinson and others 1992). The

yield of radicals was 0.12 per mole of ethylene reacted.

Table 1 Table 1 — Peroxides in rainwater (Hellpointer and Gab 1989). Table 1 1

H H2O2 

 2 HMHP 3 HEHP 4 CH3

OOH 5

5.9 7 0.2 0.1 0.2

6.7 63.1 nd nd 0.3

6.4 110.6 nd nd 0.4

Isoprene

Isoprene is the basic member of the group of plant natural products known as

isoprenoids. Important classes of these compounds are formed from multiples of

the basic five carbon unit: monoterpenes (C10), sesquiterpenes (C15), and

diterpenes (C20). These compounds are important articles of commerce because of

Figure 1 — Reaction of ozone

with ethylene.

Session I Biochemical Reactions of Ozone in Plants Mudd

H

H

C C

H

H

+O3

H H

H H

O

O O

H

H

C O +

CH2 HOCH2

OO OOH

H O2

.

.

H

H

C O + H O2 2

1 Data are expressed as μmoles/liter.

2 H2O2: hydrogen peroxide.

3 HMHP: hydroxymethyl hydroperoxide.

4 HEHP: hydroxyethyl hydroperoxide.

5 CH3OOH: methyl hydroperoxide.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 5

characteristic odors. Many terpenoids are volatile and give rise to the characteristic

odors associated with some plants.

Another volatile compound produced in sometimes copious amounts by

plants is isoprene, which can react with ozone (fig 2). Theoretically one molecule

of isoprene can give rise to two molecules of HMHP. The HMHP can degrade to

formaldehyde and hydrogen peroxide. Results of different laboratory studies

indicate in one case that the major product is hydrogen peroxide, and in another

case that the major product is HMHP. These compounds are in equilibrium:

HMHP <-----> formaldehyde + hydrogen peroxide.

Hydroxyl radicals are also formed in the reaction of ozone with isoprene. The yield

is 0.27 radicals per isoprene reacted. (Atkinson and others 1992).

Fatty acids

Fatty acids are most frequently found in plant tissues esterified with the trihydric

alcohol glycerol. Triglycerides are the most important storage fat of seeds. More

importantly, when glycerol is esterified with two fatty acids and one polar head

group such as phosphorylcholine, the compound formed is amphipathic, which

has a lipid soluble moiety and a water soluble moiety. The property of

amphipathicity is essential for the formation of the lipid bilayer of the plant

membranes. Any oxidation of the fatty acids in the plant membranes will destroy

the integrity of the membrane.

The reaction of ozone with fatty acids has a long history — the reaction was

used to determine the position of double bonds for many years. Using oleic acid as

an example, the reaction is exactly analogous to the reaction of ozone with ethylene

(fig.3). The double bond is broken with the formation of keto groups on the two

nine carbon fragments of oleic acid. The hydroxyhydroperoxide is an intermediate,

similar to the case of ethylene. The very significant difference in the ozonolysis of

Figure 3 — Reaction of ozone

with oleic acid.

Figure 2 — Reaction of ozone

with isoprene.

O3

CH CH CH CH 2

H

H

C O + H O2 2

HOCH2

OOH

(HMHP)

2

Session I Biochemical Reactions of Ozone in Plants Mudd

(CH ) CH CH(CH ) CH C 2 7 2 7 3

O

O

oleate

O3

+

+

9 - oxo - nonanoate

O

O

H

O

H

C 2 7 (CH ) COOH

nonanaldehyde

OHC(CH ) CH 2 7 3

O

O C 2 7 (CH ) CHO H O2 2
6 USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998.

fatty acids is that they are most often esterified to form phospholipids, which are

a universal feature of biological membranes, forming a lipid bilayer. After

ozononlysis the product of phospholipid oxidation has only a nine-carbon chain

(1-acyl-2-(9-oxononanyl) sn-glycerophosphocholine (Santrock and others 1992)

(fig. 4). This compound has the capability to lyse biological membranes.

Even though these reactions can be measured in solutions of phospholipid, it

is doubtful that ambient ozone has access to the double bonds in the phospholipid

bilayer of the biological membrane.

Amino Acids and Proteins

All parts of plant cells contain proteins. They may be structural in nature, or they

may be catalytic. They may be soluble or membrane-bound. The plasma membrane

is close to 50 percent lipid and 50 percent protein. The proteins of the membrane

are responsible for various catalytic activities, including pumps for sodium,

potassium, and calcium. The proteins of the plasma membrane are probably also

responsible for the reception of chemical stimuli (natural and unnatural). The

reaction of ozone with membrane proteins can be visualized as having a

devastating effect.

Reaction of ozone with a few amino acids and proteins has been well

documented (Mudd and others 1969, Previero and others 1963) The reactive amino

acids include tryptophan, histidine, and methionine (fig. 5). The product of

tryptophan ozonolysis is N-formylkynurenine (Kuroda and others 1975, Previero

and others 1963), which is consistent with a classical ozonolysis breaking the

double bond. The product of ozonolysis of histidine, when it is in peptide linkage,

is aspartic acid which also implies a classical ozonolysis of the double bond of the

imidazole ring (Berlett and others 1996). The product of oxidation of methionine

is methionine sulfoxide (Johnson and Travis 1979, Mudd and others 1969) showing

a pathway of ozone oxidation that is clearly not a classical ozonolysis. Some other

amino acids that are oxidized by ozone include cysteine, tyrosine, and

phenylalanine. Proteins that are inactivated by ozone include lysozyme (Kuroda

and others 1975), glyceraldehde-3-phosphate dehydrogenase (Knight and Mudd

1984), and glutamine synthase (Berlett and others 1996).

Ascorbic Acid

Ascorbic acid (vitamin c ) is a widely distributed constituent of plant tissue. It has

long been known that fruits can prevent and correct the results of vitamin c

deficiency in humans. Ascorbic acid is an outstanding antioxidant. It is fundamental

important in the detoxification of superoxide produced during the reactions of

photosynthesis. Ascorbic acid is also an important cofactor in hydroxylation

reactions such as the formation of hydroxyproline. The presence of ascorbic acid

in apoplastic fluid can be viewed as a protection against ozone, but it is incomplete

since the effects of ozone on plant cells is obvious.

Figure 4 — 1-palmitoyl-2 (9-

oxononanyl)-sn3-glycerophosphorylcholine.

1-palmitoyl-2-(9-oxononanyl)-sn3-glycerophosphorylcholine

O

O

O

-

H OC(CH ) CH 2C 2 14 3

H 2 7 C OC(CH ) CHO

OPOCH CH N(CH ) + 2 2 3 3

H C2 O

(PN PC) 1

Session I Biochemical Reactions of Ozone in Plants Mudd
USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998. 7

Foliar spray of ascorbic acid can prevent ozone damage to plants (Freebairn

1957). Ascorbic acid in the apoplastic fluid of plant leaves has also been shown

to be oxidized by ozone (Luwe and others 1993). The reaction of ozone with

ascorbic acid is stoichiometric, and the reaction is more rapid at pHs above the

first pK (pH 4.7) (fig. 6) (Giamalva and others 1985). The reactivity of ascorbic

acid may also be caused by the cleavage of the double bond in the ring

(Chameides 1989), and the oxidation product may be an epoxide formed at the

double bond (Kanofsky and Sima 1991). There are precedents for both reactions.

An examination of the product of oxidation of ascorbic acid by ozone using

nuclear magnetic resonance, mass spectrometry, and enzymic re-reduction

using DHA reductase (fig. 7) has shown beyond doubt that the product is

dehydroascorbic acid. Since dehydroascorbate reductase is common in plants,

the oxidation of ascorbic acid by ozone in the apoplastic fluid is not a suicideprotection mechanism but rather a renewable form of defense. This defense

does depend on the ability of the plant to take up DHA, re-reduce it in the

cytoplasm and then export AA to the apoplastic fluid.

Phenylpropenoids

The phenylpropenoids are unique plant natural products. They have been

recognized and studied for many years because of their abundance and their

characteristic reactions. The phenylpropenoids are synthesized by way of shikimic

acid and phenylalanine. Phenylalanine is the direct precursor of the simplest

phenylpropenoic acid: cinnamic acid, catalyzed by phenylalanine-ammonia lyase

(PAL). PAL has received a great deal of attention because it responds to many

stimuli, including irradiation, pathogen infection, and ozone. Phenylpropenoic

acids are common in plants. They comprise cinnamic acid coumaric acid, caffeic

Figure 5 — Reaction of ozone

with amino acids.

Figure 6 — Reaction of ozone

with ascorbic acid.

Session I Biochemical Reactions of Ozone in Plants Mudd

N-formylkynurenine

1. TRYPTOPHAN

2. HISTIDINE

3. METHIONINE

aspartic acid

methionine sulfoxide

–

N

H

NH

C O

CH CH 2 O3

C O

COCH CH 2

NCHO

NH

NH

C O

CH CH 2

O3

+ N N

H H2

OOCCH CH 2

NH

C O

O3

NH

C O

CH SCH CH CH 3 2 2

NH

C O

CH SCH CH CH 3 2 2

O

OH

O

O

O

–

ascorbic acid

CHOH.CH OH 2 CHOH.CH OH 2

dehydro ascorbic acid

O

O

O

O

O3
8 USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998.

acid, ferulic acid and sinapic acid. These compounds are important because they

are precursors of lignin and flavanoids. They may also have a role in protection of

plants from the damaging effects of ultraviolet-B radiation.

The phenylpropenoic acids are susceptible to oxidation by ozone.3 In some

cases the major pathway of oxidation is ozonolytic cleavage of the double bond in

the propenoic acid side chain. In other cases the oxidation seems to be directed to

the phenolic ring. Pryor (1974) has suggested a mechanism for the attack of ozone

on the phenolic compounds that involves radical attack on the phenolate ion.

However, the reaction of ozone with the propenoic acid moiety is clearly important.

Messenger Molecule Synthesis

Ethylene

The production of ethylene after the exposure to ozone is one of the fastest

responses known. The speed of the response is consistent with the role of ethylene

as a messenger molecule in the succession of events after ozone exposure

(Langbartels and others 1991). The messenger RNA for aminocyclopropane

carboxylic acid synthase has also been shown to rapidly increase after exposure to

ozone (Schlagnhaufer and others 1995). Gunderson and Taylor (1988) have shown

that ethylene causes stomatal responses similar to those when plants are exposed

to ethylene. Other messenger molecules could be examined such as jasmonic acid

and salicylic acid. In the case of salicylic acid the response is too slow to account for

responses to ozone (Yalpani and others 1994).

Responses to the Messenger Molecules

Clear responses to messenger molecules are the formation of chitinase and bglycosidase (Schraudner and others 1992). The formation of these proteins may

be a beneficial response to the challenge of ozone, or merely a response to the

stress by production of proteins analogous to “pathogen related proteins.” It is

probable that the late production of these proteins are an indication that

damage has been done.

Protection from Ozone

In the early 1950’s the protection of plants from ozone was accomplished by

application of ascorbic acid (Freebairn 1957). But the protection was short-lived

because of the ready oxidation of ascorbic acid in the atmosphere. The protection

afforded by ascorbic acid points to a critical distinction among protectant chemicals:

3 Unpublished data on file at the Department of Botany, University of California, Riverside.

Figure 7 — Re-reduction of

ozone-oxidized ascorbic acid by

DHA reductase.

Session I Biochemical Reactions of Ozone in Plants Mudd

Absorbance 260 nm

0:0 0:05 0:15 0:25 0:40 1:00 1:30 2:00

Time

Oxidation Product

91 percent of the theoretical recovery

1.4

1.2

1

0.8

0.6

0.4

0.2

0
USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998. 9

those that are simply sacrificed and lower the effective dose of ozone to the plant,

such as foliar applications of ascorbic acid, and those that alter the plant

physiologically and biochemically in such a way as to protect the plant from

ozone. There are no convincing examples of the second class of protectants at

present. Many compounds have been tested for protection, and several have

been found effective. The most interesting and most effective of these

compounds is N-[2 2-oxo-1-imidazolidynil)ethyl]-N’-phenylurea (EDU). EDU

was first synthesized and tested for protection of plants against ozone by chemists

at DuPont Chemical Company (Carnahan and others 1978). EDU has been found

to be effective both as a foliar spray and as a soil drench (Kostka-Rick and Manning

1993). It has been stated that EDU does not react with ozone (Pitcher and others

1993), and it is therefore a candidate for effectiveness by changing the physiology

and biochemistry of the plant. Lee and Chen (1982) have reported that EDU has

cytokinin-like activity, which could be related to its activity as an antiozonant. Lee

and Bennett (1982) found that the application of EDU increased the activities of

superoxide dismutase, catalase, and peroxidase, but these findings have been

contradicted by Pitcher and others (1993). The mode of action of EDU is therefore

still controversial and worthy of much greater research effort.

Another method for plants to resist the effects of ozone is to develop resistant

varieties. Resistant varieties of several agricultural crops include tobacco

(Heggestad 1993), alfalfa (Graumann 1972), and maize (Cameron and others 1970).

These plants can be studied to understand the reasons for resistance, and breeding

programs can be initiated to introduce the desirable trait into all commercially

available lines. One method to detect differences in resistant and susceptible plants

is to analyze the DNA by restriction fragment length polymorphism (RFLP). Neale

and Harry (1995) have discussed the uses of these technologies with particular

attention to coniferous trees. The identification of DNA fragments associated with

resistance could be useful to breeders, especially since the DNA testing for

resistant crosses can be made on seedlings rather than mature plants.

Conclusion

The reaction of ozone with many types of compounds found in plants have been

studied in vitro. It is difficult to connect these known chemical reactions to

reactions which are responsible for the succession of events in the plant which

eventually result in characteristic lesions of ozone damage. How are we going to

make the connection between chemical reactions and visual symptoms? One

approach is to compare the DNA of genetically related plants that are either ozoneresistant or ozone susceptible. Methods of analysis include the use of DNA probes

for known enzymes that may be involved in ozone resistance. Another method is

the comparison of DNA fragments derived by action of restriction endonucleases.

Techniques of molecular biology offer the most promise for understanding the

differences between ozone-resistant and ozone susceptible plants.

References

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ozone with a series of terpenes. ozone with a series of terpenes. Journal of Geophysical Research 97:6065. ozone with a series of terpenes. 

Berlett, B.S.; Levine, R.L.; Stadtman, E.R. 1996. A comparison of the effects of ozone on the modification of amino acid

residues in glutamine synthetase and bovine serum albumin. residues in glutamine synthetase and bovine serum albumin. Journal of Biological Chemistry 271:4177. residues in glutamine synthetase and bovine serum albumin. 

Cameron, J.W.; Johnson Jr, H.; Otto, H.W. 1970. Differential susceptibility of sweet corn hybrids to field injury by air

pollution. pollution. Horticultural Science 5:217. pollution. 

Carnahan, J.E.; Jenner, E.L.; Wat, E.K.W. 1978. Prevention of ozone injury to plants by a new protectant chemical. . Prevention of ozone injury to plants by a new protectant chemical.

Phytopathology 68:1225.

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Chameides, W.L.; 1989. The chemistry of ozone deposition to plant leaves: Role of ascorbic acid. The chemistry of ozone deposition to plant leaves: Role of ascorbic acid. Environmental Science The chemistry of ozone deposition to plant leaves: Role of ascorbic acid. 

Technology 23:595.

Freebairn, H.T. 1957. Reversal of inhibitory effects of ozone on oxygen uptake by mitochondria. . Reversal of inhibitory effects of ozone on oxygen uptake by mitochondria. Science 126:303. . Reversal of inhibitory effects of ozone on oxygen uptake by mitochondria. 

Giamalva, D.; Church, D.F.; Pryor, W.A. 1985. A comparison of the rates of ozonation of biological antioxidants and

oleate and linoleate esters. oleate and linoleate esters. Biochemical Biophysical Research Communication 133:773. oleate and linoleate esters. 

Graumann, H.O. 1972. Notice of release of alfalfa clones resistant to ozone. . Notice of release of alfalfa clones resistant to ozone. USDA Bulletin. . Notice of release of alfalfa clones resistant to ozone. 

Gunderson, C.A.; Taylor Jr, G.E. 1988. Kinetics of inhibition of foliar gas exchange by exogenous ethylene: an

ultrasensitive response. ultrasensitive response. New Phytology 110:517. ultrasensitive response. 

Heggestad, H.E. 1991. Origin of Bel-W3, Bel-C and Bel-B tobacco varieties and their use as indicators of ozone. Origin of Bel-W3, Bel-C and Bel-B tobacco varieties and their use as indicators of ozone.

Environmental Pollution 74:264.

Hellpointer, E.; Gab, S. 1989. Detection of methyl, hydroxymethyl and hydroxyethy hydroperoxides in air and

precipitation. precipitation. Nature 337:631. precipitation. 

Hewitt, C.N.; Kok, G.L. 1991. Formation and occurrence of organic hydroperoxides in the troposphere: Laboratory and

field observations. field observations. Journal of Atmospheric Chemistry 12:181. field observations. 

Johnson, D.; Travis, J. 1979. The oxidative inactivation of human a-1-proteinase inhibitor. Further evidence for

methionine at the reactive center. methionine at the reactive center. Journal of Biological Chemistry 254:4022. methionine at the reactive center.

Kanofsky, J.R.; Sima, P. 1991. Singlet oxygen production from the reactions of ozone with biological molecules. Singlet oxygen production from the reactions of ozone with biological molecules. Journal Singlet oxygen production from the reactions of ozone with biological molecules. 

of Biological Chemistry 266:9039.

Knight, K.L.; Mudd, J.B. 1984. The reaction of ozone with glyceraldehyde-3-phosphate dehydrogenase. The reaction of ozone with glyceraldehyde-3-phosphate dehydrogenase. Arch Biochemical The reaction of ozone with glyceraldehyde-3-phosphate dehydrogenase. 

Biophysics 229:259.

Kostka-Rick, R.; Manning, W.J. 1993. Dose-response studies with the antiozonant EDU, applied as a soil drench to two

growth substrates, on greenhouse grown varieties of Phaseolus vulgaris L. Environmental Pollution 82:63. growth substrates, on greenhouse grown varieties of Phaseolus vulgaris L

Kuroda, M.; Sakiyama, F.; Narita, K. 1975. Oxidation of tryptophan in lysozyme by ozone in aqueous solution. Journal of Oxidation of tryptophan in lysozyme by ozone in aqueous solution

Biochemistry 78:641.

Laisk, A.; Kull, O.; Moldau, H. 1989. Ozone concentration in leaf intercellular spaces is close to zero. Plant Physiology Ozone concentration in leaf intercellular spaces is close to zero

90:1163.

Langebartels, C.; Kerner, K.; Leonardi, S.; Schraudner, M.; Trost, M.; Heller, W.; Sandermann Jr, H. 1991. Biochemical Biochemical

plant response to ozone I Differential induction of polyamine and ethylene biosynthesis in tobacco. Plant Physiology plant response to ozone I Differential induction of polyamine and ethylene biosynthesis in tobacco

95:882.

Lee, E.H.; Bennett, J.H. 1982. Superoxide dismutase. A possible protective enzyme against ozone injury in snap beans

(Phaseolus vulgaris Phaseolus vulgaris L.). Plant Physiology 69:1444.

Lee, E.H.; Chen, C.M. 1982. Studies of the mechanisms of ozone tolerance: Cytokinin-like activity of EDU, a compound

protecting against ozone injury. Physiol Plant 56:486. protecting against ozone injury

Luwe, M.W.F.; Takahama, U.; Heber, U. 1993. Role of ascorbate in detoxifying ozone in the apoplast of spinach (Spinacia

oleracea L.) leaves. Plant Physiology 101:969. L.) leaves

Mehlhorn, H.; Tabner, B.J.; Wellburn, A.R. 1990. Electron spin evidence for the formation of free radicals in plants

exposed to ozone. Physiol. Plant 79:377. exposed to ozone

Mehlhorn, H.; Wellburn, A.R. 1987. Stress ethylene formation determines plant sensitivity to ozone. Nature 327:417. Stress ethylene formation determines plant sensitivity to ozone

Mudd, J.B.; Leavitt, R.; Ongun, A.; McManus, T.T. 1969. Reaction of ozone with amino acids and proteins. Atmospheric Reaction of ozone with amino acids and proteins

Environment 3:669.

Neale, D.B.; Harry, D.E. 1994. Genetic mapping in forest trees: RFLPs, RAPDs, and beyond. AgBiotechnology 6:107. Genetic mapping in forest trees: RFLPs, RAPDs, and beyond

Pitcher, L.H.; Brennan, E.; Zilinskas, B.A. 1992. The antiozonant EDU does not act via superoxide dismutase induction in

bean. Plant Physiology 99:1388. bean

Previero, A.; Scoffone, E.; Pajetta, P.; Benassi, C.A. 1963. Indagini sulla struttura delle proteine. Nota X. Compartmento

degli amminoacidi di fronte all’ozono. Gazz ChimItal 93:841. degli amminoacidi di fronte all’ozono

Pryor, W.A. 1994. Mechanisms of radical formation from the reactions of ozone with target molecules in the lung. Free Mechanisms of radical formation from the reactions of ozone with target molecules in the lung

Rad Biol Med 17:451.

Santrock, J.; Gorski, R.A.; O’Gara, J.F. 1992. Products and mechanism of the reaction of ozone with phospholipids in

unilamellar phospholipid vesicles. Chemical Research Toxicology 5:134. unilamellar phospholipid vesicles

Schlagnhaufer, C.D.; Glick, R.E.; Arteca, R.N.; Pell, E.J. 1995. Molecular cloning of an ozone induced 1- Molecular cloning of an ozone induced 1-

aminocyclopropane-1-carboxylate synthase cDNA and its relationship with loss of rbcS in potato (Solanum

tuberosum L) plants. Plant Molecular Biology 28:93. tuberosum L) plants

Schraudner, M.; Ernst, D; Langebartels, C.; Sandermann Jr, H. 1992. Biochemical plant responses ozone. III. Activation

of the defense-related proteins b 1,3-glucanase and chitinase in tobacco leaves. Plant Physiology 99:1321. of the defense-related proteins b 1,3-glucanase and chitinase in tobacco leaves

Yalpani, N.; Enyedi, A.J.; Raskin, I. 1994. Ultraviolet light and ozone stimulate accumulation of salicylic acid, Ultraviolet light and ozone stimulate accumulation of salicylic acid,

papthogenesis related proteins and virus resistance in tobacco. Planta 193:372. papthogenesis related proteins and virus resistance in tobacco

Session I Biochemical Reactions of Ozone in Plants Mudd
USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998. 11

Abstract

Urban air basin produced oxidants, notably ozone, induce a decline in productivity in plants. This loss of productivity

is manifested by slower growth, hindered development, lower reproduction rates, impaired ability to resist disease,

and other stresses. While many metabolic events have been linked to oxidant exposure, three major shifts have been

well-studied: increased production and more rapid turnover of antioxidant systems; production of symptoms similar

to a mechanical wounding of the tissue, especially ethylene production; and decline in photosynthesis. Although these

processes may be linked metabolically at a fundamental level, the mechanisms leading to a decline in photosynthesis

have been shown to directly lower plant productivity. There are two distinct changes in physiology which can directly

alter the photosynthetic rate by leaves: a closure of the stomata limiting CO2

 concentration, and a decline in the

ability to fix CO2

 within the chloroplast. In many studies it is difficult to discriminate which is more critical and

which triggers various effects because in the final analysis, both limit carbon assimilation. The mechanisms of

stomatal closure may be linked to the loss of membrane permeability and transport because of oxidation of

membrane channels and transport proteins and/or oxidation to an increased sensitivity of the stomata to closure 

signals, such as internal Ca2+ levels or abscisic acid. On the other hand, impairment of the processes of photosynthesis

is probably not caused by changes in the light gathering photosystems, but rather by a loss of CO2

 fixing ability

induced by a decline in Ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) or by an alteration in normal

metabolite flow via changes in ionic balance. It is not yet clear which of these mechanism is more critical to each 

individual plant species under varied environmental conditions. However, the loss of photosynthetic products is only 

the first step in the loss of productivity. Much evidence indicates that translocation of the fixed carbon is altered by

oxidants. Because translocation is very sensitive to energy and carbohydrate status as well as membrane function of

the leaf, phloem loading would also be at risk by oxidant exposure. A decline in carbohydrate levels to roots and

growing shoot tips would have profound effects upon the plant’s ability to grow and respond normally to the

integrated effects of the remainder of the environment.

Introduction

Many studies that describe the attack of ozone upon the tissues and physiological

processes of green plants have used agriculturally important crops (Heagle 1989;

Heath 1980, 1994a, [In press]; Heck and others 1988; Karpinski and others 1993).

While native plants in the ecosystem may respond differently to ozone than crop

plants, certain fundamentals seem to be constant between species. Thus, some

generalities of physiology can be stated with certain confidence. To be sure, we

have much more to learn, but the concepts and ideas from basic research adds to

our “more practical” knowledge base. Understanding how ecosystems can be

protected from pollutants requires the foundations of basic research.

Definitions of Forest Decline

“Forest decline” can be defined as the negative changes within forest ecosystems

induced by a degeneration in air quality. Two examples of this decline include the

process of sinking to a “weaker” or inferior condition, or a diversion from the

“normal” development process. In both cases we are forced to define what is meant

by the terms “weaker” and “normal.” Studies of “normal” ecosystems are few;

thus, normal is not inadequate when defined experimentally. Currently, a “normal”

ecosystem is that with very low levels of air pollutants. Finding such an ecosystem

is not a simple task (Sandermann and others [In press]). A “weaker” ecosystem is

one which is not resistant to attack by biotic or abiotic stresses. The loss of trees in

1 An abbreviated version of this manuscript was presented at the International Symposium on Air Pollution and

Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Plant Physiology and Biophysics, Department of Botany and

Plant Sciences, University of California, Riverside, CA 92521-0124 U.S.A.

Oxidant Induced Alteration of Carbohydrate

Production and Allocation in Plants1

Robert L. Heath2
12 USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998.

Session I Oxidant Induced Alteration of Carbohydrate Production and Allocation in Plants Heath

the San Bernardino Mountains in southern California is a good example because

ozone causes an early loss of foliage, seemingly giving the tree less resources to

fight a later attack by bark beetle which ultimately kills the tree (Miller 1992).

The observational definitions of plant “changes” caused by air quality include:

[1] visible injury, [2] loss of productivity, [3] inability to withstand other stresses,

[4] accelerated senescence, and [5] changes in metabolic pathways. Visible injury

is often used as the primary observation of forest decline and includes loss of

chlorophyll, necrosis, “water-logging” in deciduous leaves, and production of

anthocyanin-containing “spots” (Jacobson and Hill 1970). The loss of productivity

has been most easily measured by a decline in timber production in a forest. The

inability to withstand other stresses has been found in forests attacked by bark

beetles, and an inability to compensate for a lowering of nutrient and water

availability are examples of other natural abiotic stress. Accelerated senescence is

most easily observed as early loss of needles and crown thinning but can include

an altered leaf or root development. Changes in metabolic pathways are the most

difficult to observe in the field, yet these observations can most clearly indicate a

plant’s ability to respond ultimately to the ozone attack.

Plant Physiology and Ozone

Various hypotheses have been proposed to explain the interaction of plant

physiological processes and ozone attack (table 1). Although the hypotheses

overlap at the fundamental biochemical level, each hypothesis is an independent

process, and individual research groups tend to focus on only one at a time. The

antioxidant protection hypothesis involves both the amount of each antioxidant

present and the ability to produce antioxidants which serve to eliminate ozone so

that ozone cannot move into any cellular site where it will induce damage. Ozone

Table 1 — Various hypotheses that explain the modification of plant physiological processes induced by ozone. Table 1

General General Specific Detailed

Antioxidant protection

Superoxide:

Superoxide dismutase

Hydrogen peroxide

Peroxidases

Hydroxyl radical

Ascorbate, glutathione

Tocopherol

Wounding response

Wounding proteins or pathogen

response proteins

Chitinase

β -glucanase

Ethylene production

Loss of photosynthetic capacity

Inappropriate stomatal response

Photosynthetic processes

Photo-inhibition of photosystems

Loss of carboxylation

Slowing of translocation

Membrane dysfunction

Loss of ion channels (K+

 , Ca2+)

Loss of permeability

Channels

Membrane structure

Loss of signal transduction receptors
USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998. 13

itself is not expected to be stable within the cell or its wall because of its high

reactivity. Three major transformation species are currently believed to be formed:

superoxide (O2

-

), hydrogen peroxide (H2O2), and hydroxyl radical (HO.) (Grimes

and others 1983, Möller 1989, Mudd [In press]). Other possibilities, such as ozonide

radical (O3

.

) and peroxyl radical (HOO.) have been described (Heath 1986), but

their reactions are currently poorly understood. Data seem to indicate that many

oxidative species from ozone can be detoxified easily by reactions with ascorbic

acid or glutathione (Benes and others 1995, Bors and others 1989, Chameides 1989,

Gupta and others 1991, Guzy and Heath 1993, Kangasjärvi and others 1994, Luweand

others 1993, Polle and others 1995, Willekons and others 1994).

If interception by antioxidants is unsuccessful, then ozone (or its oxidative

products) will alter the membrane function through oxidation of critical sulfhydryls

of ion channels or pumps and induce an increase in general membrane permeability

via inhibition of channel closure or alterations in the membrane structure (Heath

1988, 1994a). Another, but unproved, mechanism is the false triggering of the

signal transduction receptors by a direct or indirect chemical modification. Both

effects would lead to a non-natural balance of ions across all membranes. The

most critical of these ions is Ca2+; its plasmamembrane efflux pump is inhibited

and its general permeability is increased by ozone (Castillo and Heath 1990).

Many metabolic processes are in turn activated as the Ca2+ concentration within

the cell rises.

One major response of ozone exposure mimics a general wounding, either

mechanical or pathogen induced. Since one of the initial triggers of wounding

responses is a rise of Ca2+ level within the cell, wounding fits well with what is

known about ozone attack. This rise in Ca2+ level, in turn, triggers a series of

metabolic cascades, ultimately generating ethylene and the production of

pathogen-response (PR) proteins (Fengmeier and others 1994, Kärenlampi and

others 1994, Langebartels and others 1991, Schraudner and others 1992, Sharma

and Davis 1994). It is not clear that ozone directly produces these responses; some

oxidative product of ozone may be the key chemical. For example, the prevention

of ethylene release has been shown to prevent the induction of visible injury

(Mehlhorn and Wellburn 1987, Mehlhorn and others 1991), and the visible injury

ultimately has been considered a result of the chemical interaction of ozone and the

double bond of ethylene producing a toxic product (Gunderson and Taylor 1991,

Taylor and others 1988, Tingey and others 1976).

A clear, but mechanistically confusing, plant response to ozone is the loss of

photosynthetic capacity. Stomata generally partially close during ozone exposure

but, under some conditions, can open further (Heath 1994b). Under many

conditions the level of the primary enzyme of CO2 fixation (ribulose 1,5-

bisphosphate carboxylase/oxygenase, Rubisco) declines (Dann and Pell 1989, Nie

and others 1993, Pell and others 1994). Yet one clear visible injury pattern, the loss

of chlorophyll, signals a problem within the photosystems of the chloroplast

(Heath 1989). Photoinhibition can be induced if CO2 fixation becomes too slow for

a given rate of photon capture. Many observed events suggest that while carbon

assimilation within the leaf declines, translocation of carbon is inhibited even more

so such that growing points of the plant are inhibited and root/shoot ratios are

altered (Dugger and Ting 1970, Tjoelker and others 1995).

Examination of the changes in photosynthetic capacity suggests that a multiple

series of declines are at the heart of a loss in productivity and resource accumulation

of the plant. In general, stomata partially close during ozone exposure. The

apertures of stomata are governed by a dynamic balance of water loss and internal

CO2 concentration (which is fixed by a balance of gas flow through the stomata and

carbon assimilation) (Farquhar and Sharkey 1982). We do not know if a membrane

imbalance first leads to a loss of osmotically accumulated water from the guard cell

or if the imbalance is caused by an inhibition of CO2 fixation. In fact, there are many

possible interactive mechanisms: stomata closure can occur with an influx of Ca2+

into the guard cell in which its higher concentration changes the sensitivity of the

Session I Oxidant Induced Alteration of Carbohydrate Production and Allocation in Plants Heath
14 USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998.

Figure 1 — Changes in the

production of m-RNA for the

small subunit of Rubisco. Tomato

plants, grown in a growth

chamber for 3 weeks, were

fumigated with 0.33 ppm ozone

(produced within O2

 stream) at

an air temperature of 28 degrees

C and a relative humidity of 25

percent for 3 hours. No visible

injury could be seen even after

several days, but a slight (ca. 40

percent) stomatal closure could

be observed towards the end of

the fumigation period (fig. 3). The

message RNA was probed

(Cohen and Bray 1990) and

quantified on the slot blot

membrane by the phosphor

Imager System (Molecular

Dynamics). 3

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

guard cell to abscisic acid (ABA) transported to the leaf from the roots or produced

within the leaf due to a lowered cell water potential (Atkinson and others 1990).

These interactive mechanisms make understanding of the full process difficult.

A poor balance between light gathering (photosystems) and CO2 fixation by

Rubisco seems to lead to photoinhibition (Farage and others 1991). Changes in

chlorophyll fluorescence patterns and in kinetics of protein production (e.g., the

D1 protein of photosystem II) (Barber and Anderson 1992, Osmond 1981) suggest

that this is a real mechanism with the possibility that it is not expressed in all plants

(Nie and others 1993, Pino and others 1995). The loss of Rubisco is difficult to

measure as Rubisco is present in very high concentration. Its level declines in

senescing leaves much faster after ozone exposure, suggesting a role in early

senescence. An easier method to follow Rubisco changes is by measuring the

concentration of its m-RNA leading to the production of the protein rbcS (the

message for its small subunit is nuclear transcribed). According to studies by Pell

and others (1994) and Reddy and others (1993), a rapid, but not complete, loss of

the protein message occurs within an hour or so of exposure but recovers within

a day after the exposure ceases (fig. 1).

The full scheme for the interaction between membrane dysfunction and Ca2+

changes and the other events observed during ozone exposure is not easily

decoded. However, the current data indicates that metabolic pathways are

regulated and mutually dependent. When ozone alters one metabolic event many

others far removed from that initial site of interaction are changed.

We have used a dual label isotope porometer (Johnson and others 1979) to

measure the stomata conductance of water vapor (using inward flowing 3H2O as

an analog for normal outward flow of water vapor from the leaf). Assimilation is

measured simultaneously by 14CO2 fixation. The gas stream passing over the leaf

has both isotopes (fig. 2). Knowing the stomata conductance for H2O vapor allows

the calculation of the conductance for O3 and with the known external level of

ozone, a dose of ozone within the leaf can be calculated (Heath 1994b). The

inhibition of stomata conductance is greater than the inhibition of assimilation, but

under the low light intensity of these experiments, the stomata can close a great

deal without inhibiting assimilation (Heath 1994b). We are currently using the

dual label porometer in the single label mode (using only 14CO2

 at high specific

activity) to label photosynthetically- fixed carbon in a leaf in order to follow its

movement through translocation. Preliminary results suggest ozone exposure

dramatically slows carbon movement.

Another method to continuously measure stomata conductance involves the

use of leaf temperature as a probe of evaporative water loss through transpiration.

Time (hr) after start of fumigation

150

100

50

0

1 2 3 3.5 3.5 3.5 3.2 3.2 3.2 5.2

Level compared to control (pct)

Session I Oxidant Induced Alteration of Carbohydrate Production and Allocation in Plants Heath
USDA Forest Service Gen. Tech. Rep. PSW-GTR-166. 1998. 15

Figure 2 — Dual isotope label

porometer (Johnson and others

1979). This machine allows the

measurement of water conductivity (measured inversely by

the flow of tritiated water from

the gas stream into the leaf) and

carbon assimilation (measured by

the flow of 14C carbon dioxide

into the leaf). The area labeled is

a circle of diameter of 0.55 cm.

By using two thermocouples wired in series but with legs reversed (the same metal

joined at a common junction) and one thermocouple touching the leaf and the other

in the air just below the leaf, the differential temperature (dT = Tleaf - Tair) can be

continuously recorded. Calibrations allows dT to be used as a measure of

transpiration rate. By using the air temperature and relative humidity, the stomata

conductance can be calculated, and the ozone dose can be “measured” continuously

during exposure. A plot for tomato exposed to ozone was measured by using this

technique (fig. 3). The change in conductance is measured against the actual

delivered dose of ozone to six leaves. Each leaf begins with a slightly different

conductance (as they are different leaves on the two plants and are at different

developmental ages, which affects their conductance); thus, the total time of

exposure at which stomata begin to close varies. However, converting time into

accumulated dose for each gives clearer results; the dose for the beginning of

closure is about the same as 8 nmole cm-2-leaf area. The stomata close to about 50-

60 percent of the initial level at a dose of about 30-35 nmole cm-2-leaf area. Higher

doses do not seem to close the stomata any further.

Conclusion

Figure 3 — Changes of stomatal conductance during fumigation of tomato plants with ozone. The data were collected and dose was

calculated by the amount of ozone in the air multiplied by the stomatal conductance for ozone each 210 seconds. This is an average

of thermocouple measurements for six leaves (2 on 2 plants, 1 on 4 plants). Several plants were destructively removed during the course of

the experiment for m-RNA sampling.

14CO2

+3H2O

uptake of CO2 gc

gw

rm=rc- 1.56 rw

uptake of H2O

g=1/r

VENT Leaf

Light Trap

Trigger

Valve

Gun

Chamber

Capilliary Tube

tank Hydrator

Session I Oxidant Induced Alteration of Carbohydrate Production and Allocation in Plants Heath

Fumigated Dose of Ozone (nmol/cm )

0.10

0.00

0.10

0.20

0.30

0.40

0 20 40 50

change in net conductance (cm/sec)

Fumigated Dose of Ozone (nmol/cm )

120

100

80

60

40

0 20 40 50

2 

Change in Conductance over Control (pct of initial)

2 

pct
16 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Results of the various studies discussed in this paper have made sizable

advancement in the understanding of ozone induced plant changes. Several

testable hypotheses regarding the mechanism of ozone induced injury have been

formulated (Bors and others 1989, Castillo and Heath 1990, Eckardt and Pell 1994,

Langebartels and others 1991, Pino and others 1995) and cultivars and mutants of

varied species have been generated which differ in their sensitivities to ozone

(Guzy and Heath 1993, Sharma and Davis 1994). The understanding of the basic

processes of plants has made this progress possible. Our comprehension of how

plants respond to the environment continues to grow. In the future, better

technology to allow us to measure physiological events, and a more complete

understanding of genetics and the use of mutants should make it possible to

understand how ozone exposure puts a plant at risk and how we can aid the plant

in its attempt to protect and repair itself in the face of that risk.

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Session I Oxidant Induced Alteration of Carbohydrate Production and Allocation in Plants Heath

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 19

Abstract

A variety of vascular plant species exhibit typical foliar injury symptoms when exposed to ambient ozone, making

them useful as bioindicators of relative air quality for a particular location or region. They are quite useful in areas

where mechanical ozone monitors are not available. Bioindicators are often introduced plant species known as

sentinels. They are known to be sensitive to ozone and will respond rapidly if they are given special care to ensure

ozone uptake and injury. Sentinels are usually genetically-uniform, rapid growing herbaceous annuals. Their rapid

and well-characterized response to ozone exposure has made them quite useful. Bel-W3 tobacco (Nicotiana tabacum

L.) is a sentinel plant bioindicator for ozone that is used all over the world. Detector bioindicators are plant species

that are found growing naturally in an area and known to be sensitive to ozone only when conditions are appropriate

for ozone uptake and plant injury. Detectors are often slow-growing, determinate perennial plants, shrubs, or trees

that respond slowly to ozone, with symptoms occurring quite late in the growing season. Populations of detectors are

not genetically uniform and only part of a population may show ozone injury symptoms. Black cherry (Prunus

serotina L.) is a common detector bioindicator for ozone in North America. A comparison of surveys of sentinel and

detector bioindicators in the same area often show different results. From an ecological perspective, visible injury on

a detector bioindicator is more significant than visible injury on a sentinel bioindicator. When using plants as

bioindicators, careful consideration needs to be given to the nature, requirements, and utility of sentinels and

detectors in relation to the relevance and utility of the results obtained.

Introduction

For more than 50 years, certain plant species have been used as bioindicators

because they are sensitive to ozone under ambient conditions (Middleton and

others 1950, Noble and Wright 1958). Sensitive individuals exhibit typical foliar

injury symptoms when exposed to ambient ozone under conditions appropriate

for ozone uptake. These symptoms are considered to be diagnostic or typical as

they have been verified in exposure/response studies under experimental

conditions (Krupa and Manning 1988). These plants are considered to be reliable

biological indicators or bioindicators for ambient ozone. The subjective

determination of the intensity or extent of foliar injury of bioindicators is used as

an index of relative air quality for ozone for a particular location or region. Ozone

has become an air pollution problem in most industrialized nations, resulting in an

increased interest in using bioindicator plants on a world-wide basis. According

to Guderian and others (1985), injury on Bel-W3 tobacco (Nicotiana tabacum L.) is

usually the first indication a county or region has developed an ozone problem.

The history and use of plants as ozone bioindicators has been extensively reviewed

elsewhere (Arndt and others 1987, Burton 1986, deBauer 1972, Feder and Manning

1979, Guderian and others 1985, Heck 1966, Hernandez and deBauer 1989, Manning

1991, Manning and Feder 1980, Posthumus 1982, Stuebing and Jager 1982, Tonneijck

and Posthumus 1987, Weintstein and Laurence 1989).

This paper discusses the use of introduced species or sentinels and naturally

occurring plants or detectors as bioindicators of ozone exposure.

Sentinels and Detectors

From an ecological perspective, Spellerberg (1991) recognizes two types of plant

bioindicators useful in studies designed to detect gaseous air pollutants (table 1).

Sentinels are non-indigenous plant species, consisting of well-defined selections,

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Phytopathologist, Department of Microbiology, University of Massachusetts, Amherst, MA 01003-5730.

The Use of Plants as Bioindicators of Ozone1

William J. Manning2
20 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

cultivars or clones, that exhibit diagnostic reliable foliar symptoms when exposed

to ambient ozone. Sentinels are grown in charcoal-filtered air and then introduced

into areas, usually for short time periods. They are quite useful in identifying areas

where ozone concentrations and exposures are suspected to be high enough and

of a long enough duration to cause foliar injury (Ashmore and others 1980,

Bytnerowicz and others 1993, deBauer 1972, Heck and Heagle 1970, Hernandez

and deBauer 1989, Jacobson and Feder 1974, Kelleher and Feder 1978, Lipa and

Votapkova 1995, Noble and Wright 1958, Oshima 1974, Posthumus 1982). Examples

of sentinel bioindicators for ozone include Bel-W3 tobacco (Heck and others 1964,

Heck and Heagle 1970, Heggestad 1991) and morning glory (Ipomoea purpurea)

(Manning 1977, Nouchi and Aoki 1979). Detector plants are indigenous to an area

that may exhibit typical foliar injury symptoms to ozone exposure in situ. They are

useful in assessing the long-term or cumulative effects of ozone. Examples include

the sensitive individuals in populations of black cherry (Prunus serotina) (Davis

and others 1982), Ponderosa (Pinus ponderosa) and Jeffrey (P. Jeffreyi) pines (Stolte

and others 1992), and milkweed (Asclepias syriaca) (Duchelle and Skelly 1981).

Table 1 — Types of indicator plant species used to assess relative air quality for ozone (Spellerberg 1991). Table 1

Sentinels Well-defined plants known to be sensitive to ozone are introduced into an

area to serve as early warning devices or checks on the efficiency of

abatement practices.

Examples—Bel-W3 tobacco (Nicotiana tabacum), Morning glory

(Ipomoea purpurea)

Detectors Plants that naturally occur in the area of interest that may exhibit typical

foliar responses to ozone.

Examples—Black cherry (Prunus serotina), Milkweed (Asclepias syriaca)

Sentinels are the most commonly used bioindicators of ambient ozone. They

are usually well-defined, genetically-uniform, herbaceous annuals that grow

rapidly. Their response time to ozone is rapid, serving as an early warning of ozone

presence. Rapid response, however, may only occur in the early stages of their

growth cycles, necessitating frequent re-introduction of new plants. They must be

grown in charcoal-filtered air until they are old enough to move to the field. To

minimize edaphic factors, they are usually grown in pots of a uniform artificial

growing medium. They require water, fertilizer, shading, and often pesticide

applications, on a regular basis. Protection from animals and vandals may also be

required. If sentinels do not receive the special care they require, they will not

respond well to ozone in a typical or relevant manner.

Detectors are native plants selected in situ and usually are not given any special

cultural care. Usually they are determinate perennial plants, trees, or shrubs that

respond slowly and often fairly late in the growing season. Only the sensitive

individuals in a population of a detector bioindicator will respond to ozone and

only when they experience appropriate edaphic and tropospheric conditions

coupled with ozone exposures and concentrations sufficient to cause foliar injury.

The distribution of ozone-sensitive genotypes (where there is little phenotypic

variation) in a population of a detector bioindicator is usually not well-known.

This adds uncertainty in interpreting results from detectors. When large

numbers of individuals are present, confidence coefficients increase (Stolte

and others 1992).

Session I The Use of Plants as Bioindicators of Ozone Manning
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 21

Session I The Use of Plants as Bioindicators of Ozone Manning

Bioindicator Plant Response to Ozone

Sentinels and detectors evaluated in the same area often show different results:

sentinels may respond but detectors may not. Results from ozone plant bioindicator

studies must be interpreted on the basis of a number of interacting biological,

cultural, and physical factors (table 2). Foliar ozone injury symptoms are only an

indication of previous exposure to ozone, when the concentration was high

enough for a certain time period, and environmental conditions were conducive

to ozone uptake and cellular injury. The visible response is a relative index of air

quality for ozone during a previous exposure period. It is not possible to use this

response to quantitatively assess air quality for ozone. This is because we do not

yet truly understand the relationship between the occurrence and magnitude of

plant response and the environmental conditions and concentrations and durations

of ambient ozone present when the plant injury response was initiated. If data from

a co-located or nearby mechanical ozone monitor is available, however, periods

when monitored ozone concentrations have potential biological significance or

meaning can be indicated, as reflected by frequency, duration, and intensity of

plant responses.

Table 2 Table 2 — Biological, cultural, and physical factors affecting responses to ozone (Manning 1991). Table 2 

Biological and Cultural Biological and Cultural

Factors Responses

Biological

Genetic diversity Homogeneous plants give uniform responses; species,

clones, cultivars, and provenances react differently to O3

Stage of plant Plant developmental stages and leaf age affect

development responses to O3

Cultural

General cultural Optimal or usual practices result in “typical”

practices responses to O3

Growth media Soilless media allow uniformity and reproductibility,

but are less relevant than natural soils.

Nutrients Optimal levels usually result in optimal injury from O3.

Pesticides Variable effects, ranging from none to protection, from

injury to reduced tolerance to joint effects with O3

.

Physical Physical

Factors Responses

Air movement Must be sufficient to alter boundary layer resistance

to allow O3 uptake.

Light

Intensity, Ideal value varies for each plant; less or more than

photoperiod, ideal value for each plant reduces O3

 sensitivity

and quality

Temperature Injury increases in a range from 3 to 30 °

C.

Water

Relative humidity Controls stomatal opening exchange; uptake of O3

 and

injury should increase as relative humidity increases.

Soil moisture tolerance Water stress increases O3

 due to stomatal closure
22 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Previous research has determined some well-defined bioindicators for ambient

ozone (table 3). Each of these bioindicators will exhibit typical ozone injury

symptoms (table 4). Tobacco cultivar Bel-W3 is the best-defined, best-described,

and most widely-used bioindicator for ambient ozone (Heggestad 1991). It has

become the standard sentinel bioindicator for ozone and has been used all over the

world to detect ambient ozone in areas ranging from local sites to whole countries

and continents. Heck and Heagle (1970) used Bel-W3 tobacco to determine

incidence of phytotoxic concentrations of ozone around Cincinnati, Ohio. Bel-W3

tobacco plants were used by Manning to demonstrate ozone in Warsaw, Krakow,

and forested areas in southern Poland (Bytnerowicz and others 1993). Jacobson

and Feder (1974) reported the results of a network of Bel-W3 tobacco plantings in

the northeastern United States. Long-range transport of ozone from metropolitan

New York to Nantucket Island was demonstrated by response of Bel-W3 tobacco

plants on Nantucket (Kelleher and Feder 1978). Ashmore and others (1980)

mapped phytotoxic ozone in the British Isles by using networks of Bel-W3 tobacco

plants. Schoolchildren in many European countries use Bel-W3 tobacco to assess

relative air quality for ozone (Lipa and Votapkova 1995).

Table 3 Table 3 — Selected bioindicators of ozone. Table 3 1

 Plant Latin Name References

Sentinels

Blue grass Poa annua Noble and Wright 1958

Bean Phaseolus vulgaris Oshima 1974, Sanders and others 1992

Clover Trifolium repens Heagle and others 1992

T. subterraneum Sanders and others 1992

Morning glory Ipomoea purpurea Manning 1977, Nouchi and Aoki 1979

Spinach Spinacea oleraceae Posthumus 1982

Tobacco Nicotiana tabacum Heck and Heagle 1970,

Bel-W3 (sensitive) Heaggestad 1991

Bel-B (tolerant)

Detectors

Blackberry Rubus spp. Chappelka and others 1986, Manning 1991

Black cherry Prunus serotina Chappelka and others 1992, Davis and

 others 1982

Green ash Fraxinus pennsylvatica Davis and Wilhour 1976

Milkweed Asdepias sytiaca Duchelle and Skelly 1981

Quaking aspen Populas tremuloides Karnosky 1976, Keller 1988

Sassafras Sassafras albidum Chappelka 1992

Tulip poplar Liriodendron tulipifera Davis and Chappelka 1986, 1992; Davis

 and Wilhour 1976

White ash Fraxinus americana Chappelka 1992, Davis and Wilhour 1976

Several plant species have cultivars or clones that differ in their response to

ambient ozone. The best defined incidence of this are the tobacco cultivars Bel-W3

(ozone-sensitive) and Bel-B (ozone-tolerant). Bel-W3 tobacco responds to ambient

ozone at lower concentrations than does Bel-B. When grown together in an area,

their comparative responses to ozone can provide a better description of relative

air quality for ozone. If both cultivars do not respond, then the air is relatively clean

and ozone concentrations are probably low. If Bel-W3 responds, but Bel-B does

not, then ozone concentrations are intermediate. If both cultivars respond, then

ozone concentrations are higher (Manning and Feder 1980).

1 Not all varieties or individuals in a species will be ozone-sensitive.

Session I The Use of Plants as Bioindicators of Ozone Manning
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 23

Detectors are being used to assess relative long-term air quality in forested and

wilderness areas. Study areas are selected and certain species are examined on an

annual basis for incidence and severity of foliar ozone injury. In wilderness areas

of New Hampshire and Vermont, Manning and others (1991) have surveyed black

cherry, white ash, and milkweed. Chappelka and others (1986, 1992) have used

black cherry, sassafras, white ash, yellow poplar, and blackberry in the southeastern

United States. The USDA Forest Service’s Forest Health Monitoring Program uses

detector bioindicators in its forest surveys in the U.S. (Conkling and Byers 1993).

Detector bioindicators are used in forested regions of the Carpathian Mountains

in Poland, Ukraine, Czech Republic, Slovakia, and Rumania (Manning and others,

unpublished paper).

EDU (ethylenediurea) is a chemical known to protect plants from ozone injury

(Carnahan and others 1978). It can be used to verify the response of an ozone

bioindicator in the field. This is especially useful if an ozone monitor is not readily

available or there are questions about the nature of the response of the bioindicator.

This approach was used in Poland (Bytnerowicz and others 1993). Bel-W3 tobacco

plants were outplanted in a number of locations. Half of the plants were treated

with EDU before outplanting. The treated plants did not develop any symptoms

of foliar injury while the non-treated plants developed varying degrees of typical

ozone injury symptoms, verifying that ozone was the cause.

Table 4 Table 4 — Common symptoms of foliar ozone injury (Krupa and Manning 1988). Table 4 

Methodology for Assessing

Bioindicator Plants

Great care must be taken in assessing the response of bioindicator plants to

ambient ozone. In most cases, assessment of bioindicators involves the subjective

determination of the intensity or extent of acute ozone injury symptoms (table 4).

Leaf injury evaluations should be made at regular intervals, often weekly, and by

the same person. The use of a set of color photographs illustrating degrees or

categories of severity of injury can help to standardize the evaluation process

(Heck 1966; Heck and others 1966, 1969; Oshima 1976). If plants are left in the field

for more than 1 week, then new injury on both older and new leaves needs to be

estimated and recorded.

The observer should look at each leaf and visually integrate the injured areas

of each leaf and then determine the percentage of the total leaf area that has been

injured. Depending on the type of plant, extent of injury, and purpose of the study,

the evaluation system can be quite simple and uncomplicated (table 5). With only

six indices, this is a simple system to use. Where more precise information is

required, an expanded system is more appropriate (table 6).

Acute injury

Flecking Small necrotic areas due to death of palisade cells, metallic or Flecking

brown, fading to grey or white.

Stippling Tiny punctate spots where a few palisade cells are dead or Stippling

injured, may be white, black-red, or red-purple.

 Chronic injury

Pigmentation Leaves turn red-brown to brown as phenolic tan pigments Pigmentation

(Bronzing) (Bronzing) accumulate. (Bronzing)

Chlorosis May result from pigmentation or may occur alone as chlorophyll Chlorosis

breaks down.

Premature Early loss of leaves or fruit. Premature

senescence

Session I The Use of Plants as Bioindicators of Ozone Manning
24 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Data can be depicted graphically. Plotting weekly injury scores, or number of

leaves injured, on a weekly basis against elapsed time gives a cumulative injury

curve over time. If data are available from a nearby mechanical ozone monitor,

ozone injury can be plotted against cumulative ozone or related to episodes in

which thresholds are exceeded (Manning and Feder 1980).

As an alternative to using subjective determination of the intensity or extent

of foliar injury as a measure of ozone effects, other investigators have developed

bioindicator systems that use shoot biomass response from sequential harvests as

a measure of ozone effects. Oshima and others (1976) developed a standardized

pot culture system with alfalfa (Medicago sativa L.) along an ozone gradient in

southern California. Heagle and others (1994) have developed an ozone

bioindicator system by using sensitive (NCS) and resistant (NCR) clones of ladino

clover (Trifolium repens L.) NCS and NCR plants are grown in standardized pot

culture. Shoot biomass is removed at 28-day intervals and dry weights are obtained.

Ozone impact is determined by obtaining dry weights of harvested shoot biomass

and calculating the ratios of NCS to NCR. A ratio of less than one indicates that

ambient ozone has had an adverse effect on foliar biomass of NCS.

Summary

Bioindicator plants for ozone can be extremely useful in assessing relative air

quality, especially in areas where ozone monitors are not available. If they are not

used properly, however, poor quality or misleading results will be obtained.

Common factors that produce poor results are poor plant culture for sentinels,

failure to adhere to regular evaluation schedules, careless and inaccurate evaluation

of symptoms, and the use of more than one person to evaluate plant responses.

Table 5 Table 5 — System for evaluation of ozone injury for bean ( Table 5 Phaseolus vulgaris L.) (Manning and Feder 1980).

 Rating Systems

 Injury rating Injury severity index Percent leaf injury

None 0 0

Slight 1 1-25

Moderate 2 26-50

Moderate-severe 3 51-75

Severe 4 76-99

Complete 5 100

(Percent severity or class incidence) (Percent severity or class incidence)

0 0

1 0-3

2 3-6

 3 6-12

 4 12-25

5 25-50

6 50-75

7 75-88

8 88-94

 9 94-97

10 97-100

11 100

Table 6 — The Horsfall-Barratt Scale for assessing foliar injury (Horsfall and Barratt 1945, Horsfall and Cowling 1978). Table 6

Session I The Use of Plants as Bioindicators of Ozone Manning
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 25

Care should be taken in interpreting responses of sentinel bioindicators. As

they are well-watered and fertilized, they respond much more frequently than do

detectors. Their response indicates the occurrence of periods of ozone exposure

when detectors might respond if they were also growing under conditions of

sufficient soil moisture and fertility. From an ecological perspective, response of

detector bioindicators to ozone is more important than a response by a sentinel

bioindicator. A response by a sentinel indicates what could happen under ideal

conditions, while a response by a detector indicates what did happen under more

realistic conditions.

There is an unfortunate tendency to confuse a bioindicator with a biomonitor.

A bioindicator indicates that the system has been affected. A biomonitor should

also indicate how much of the causal factor was present and caused the observed

effect. Currently, bioindicators of ambient ozone cannot be used as biomonitors.

It is not possible to make quantitative inferences about air quality for ozone on the

basis of plant symptom expression alone.

With a clear understanding of the strengths and weaknesses of the use of plants

as bioindicators of ozone exposure, they can be quite useful in assessing relative

air quality, especially in programs like Forest Health Monitoring, in remote areas,

and in emerging nations, where technology is lacking.

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Session I The Use of Plants as Bioindicators of Ozone Manning
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 27

Abstract

Mountains occupy only 3 percent of Poland. They are the northern part of the European arc of the Carpathian and

Sudety Mountains, extending about 700 km along the southern Polish border. They are of medium height (about

1,500 m., maximum 2,600 m. a.s.l.), and diversified in terms of climate, geology, soils, vegetation, and anthropogenic

impacts. The forest vegetation of the Sudety and Carpathian Mountains forms three elevational zones. Forests occur

in the foothills (as high as 600 m. a.s.l.), the lower mountain forest zone (as high as 1,250 m. a.s.l.) and the upper

montane forest zone (as high as 1,500 m. a.s.l.). The original lower mountain forests consist mainly of the fir (Abies

alba Mill.) and beech (Fagus sylvatica L.), while upper montane forests consist of spruce (Picea abies [L.] Karst.).

Centuries of economic activity have changed the species composition of mountain forests. The share of the spruce has

increased considerably, and the percentage of the fir and beech has decreased significantly. In the Sudety and

western part of the Carpathian Mountains prevalence of spruce is the highest (83 percent), while in the eastern part

of the Carpathians, it does not exceed 10 percent. The percentage of beech in the Sudety and western part of

Carpathian forests is below 20 percent, but about 40 percent in the eastern part of the Carpathians. Severe weather

conditions, frequently poor habitats and improper management and considerable air pollution occurring in the past

50 years, followed by infestations of primary and secondary insects and spread of parasite fungi, have led to

considerable destruction of mountain forests. According to forecasts, the area of totally destroyed and severely

damaged mountain forests will increase considerably through the year 2010. Deterioration of forest health will

proceed from west to east. Protection of forest health against deterioration requires reduction of industrial emissions,

changing existing forests into less sensitive habitats more compatible with their carrying capacity, and recultivation

of contaminated forests.

Introduction

Generally Poland is a lowland country. Mountains occupy only 3 percent of the

country. Even though the area occupied by mountains is small, their landscape

makes them the most beautiful areas of Poland, and they are valuable in terms of

wildlife, with numerous endemic and rare plant and animal species, and specific

biocenoses (Denisiuk 1995, Szafer and Zarzycki 1972, Zarzycki and others 1991).

Humans have affected the wildlife of the Polish mountains for centuries.

Forms of human intervention have changed over the years, evolving from direct

activity (cutting forests, raking duff, grazing) to air pollution, a much more

dangerous indirect impact that is difficult to control. Human impacts, exacerbated

by specific climatic, orographic and soil conditions, have seriously worsened the

condition of mountain ecosystems, including forests, in at least part of the ranges.

General Characteristics of Polish Mountains

Polish mountains form the northern part of the European arc of the Sudety and

Carpathian Mountains (fig. 1) extending about 700 km along the southern boundary

of Poland (fig. 2). They are of medium height, with average elevation not exceeding

1500 m. a.s.l. (fig. 3). They are diversified in terms of climate, geology, soil and

vegetation, and also vary with anthropogenic impacts (Fabiszewski and Jenik 1994,

Paschalis 1995, Starkel 1991, Szafer and Zarzycki 1972, Zarzycki and others 1991).

The Carpathian climate is more continental (colder and drier) than that in the

Sudety Mountains. Continentalism increases along the west-east transect. The

Sudety Mountains are composed mainly of granite, gneiss and basalt, and the

Carpathians are comprised of sandstone and shales (Carpathian flysch) with small

State of Polish Mountain Forests: Past, Present, and Future1

Krystyna Grodzinska,2 Grazyna Szarek-Lukaszewska2

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Ecology and Associate

Professor of Ecology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland.
28 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

0 20

40

40

20 0 20 40 60 60

areas occupied by granite and limestone rock (Starkel 1991). Thus, the Sudety have

a poor substratum and strongly acidic soils, while the Carpathians have less acidic,

more nutrient-rich substratum and soil.

The vegetation of the Sudety and Carpathians is diverse by elevation (fig. 4).

The foothill zone, as high as 600-700 m. a.s.l., is occupied by deciduous and mixed

forests (deciduous and coniferous). The lower montane zone (as high as 1,250 m.

a.s.l.) is represented by mixed forests, usually beech and fir, and the upper

montane belt (as high as 1,500 m. a.s.l.) consists of spruce. Above the montane

forest zones in the higher mountain ranges, there is a dwarf mountain pine zone

(up to 1,750 m. a.s.l.), and alpine (up to 2,300 m. a.s.l.) and subalpine zones (> 2,300

m. a.s.l.) (fig. 4).

Figure 1 — Central European

Mountains Province (Szafer and

Zarzycki 1972).

Figure 2 — Polish Mountains:

the Sudety and Carpathians.

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska

16° 20° 24°

54° 54°

52° 52°

16 20° 24° °

50°

CARPATHIAN

MOUNTAINS

MOUNTAINS SUDETY

50°
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 29

Figure 4 — Vegetation zones

in Polish mountains (Denisiuk

1995, Fabiszewski and Janik

1994).

Figure 3 — The Carpathian

Mountains (Warszynska 1995).

Polish Mountains Forests

The primeval lower montane forests are comprised mainly of the fir (Abies alba

Mill.) and beech (Fagus sylvatica L.), while the upper montane forests have spruce

(Picea abies [L.] Karst). The first two species are abundant in the Carpathians, and

the third species is common in the Sudety. Generally, poor acidophilic forest

communities occur in the Sudety, while the Carpathians had more eutrophic

communities (Matuszkiewicz 1984).

The current species composition of the forests significantly differs from the

primeval one because of centuries-long, often improper management. Currently,

spruce stands occupy considerably larger mountain areas, and often the seed

source is from non-native plantations. The percentage of the fir and beech in forests

in Polish mountains has significantly decreased. In Sudety the current percentage

of the spruce in forests is more than 83 percent (Capecki 1989) (table 1). In the

Carpathians the average percentage of this species does not exceed 22 percent, and

beech and fir are about 25 percent each (Fabijanowski and Jaworski 1995). The

percentages of these three forest-forming species are different in various parts of

the Carpathian arc. In the western part (Silesian Beskid Mountains, Zywiecki

Beskid Mountains) the spruce dominates (> 70 percent), while in the eastern part

(Beskid Niski and Bieszczady Mountains beech (27 and 41 percent) and fir (31 and

20 percent) dominate (Fabijanowski and Jaworski 1995) (table 1).

The average age of Carpathian forest trees is about 50 years, and their volume

per hectare is as high as 200 m3

, with average annual growth 2.6 m3/ha/year. In selected

regions of the Carpathians there are considerably older tree stands (> 80 years), with

BESKID ZYWIECKI BESKID SADECKI BESKID NISKI BIESZCZADY

TATRY

m.a.s.l.

2500

2000

1500

1000

500

0

50 100 150 200 250 300 km

W E

19° 20° 21° 22°

.

0

 -.-.-.-

 .-.-.-.-

 -.-.-.-

 -.-.-.-.

-.-.-.-.-.-.-.-.-

 -.-.-.-.-.-.-.-

 -.-.-.-.-.-.-.-

 -.-.-.-.-.-.

 -.-.-.-.-.-

 .-.-.-.-

 -.-.-.-.-.-.-.

 .-.-.-.-.-.-.-

 -.-.-.-.-.-.-

 -.-.-.-.-.-.-

-.-.-.-.-.-.- -.-.-.-.-.-.-.-.-.-.

-.-.-.-.-.-.-.-.-.-.

-.-.-.-.-.-.-.-.-.-.- .-.-.-.-.-.-.-.-.-.-.-

.-.-.-.-.-.-.-.-.-.-.-.

.-.-.-.-.-.-.-.-.-.-.-.-

The Sudety The Carpathians

Western

Eastern

W E

m.a.s.t.

2500

2300

2100

1900

1700

1500

1300

1100

0900

0700

0500

0300

subnival zone

alpine zone

dwarf pine zone

upper montane zone

lower montane zone

foothills

m.a.s.t.

2500

2300

2100

1900

1700

1500

1300

1100

0900

0700

0500

0300

 -.-.-.-.-.-.-.-.-.-.-

 -.-.-.-.-.-.-.-.-.-.-.

 -.-.-.-.-.-.-.-.-.-.-.-

 | O | O | O | O |

| O | O | O | O | O

 O | O | O | O | O | O

O | O | O | O | O | O | O

 | O | O | O | O | O | O

O | O | O | O | O | O | O

 | O | O | O | O | O | O

O | O | O | O | O | O | 

| O | O | O | O | O |

 | O | O | O | O | 

 | O | O | O | O | O | O | O

O | O | O | O | O | O | O

O | O | O | O | O | O | O

 | O | O | O | O | O | O | O |

 | O | O | O | O | 

| O | O | O | 

 | O | O | O 

O | O | 

O | O | 

| O | O

 | O

O | O | O | O | O | O | O |

 | O | O | O | O | O | O

O | O | O | O | O 

O | O | O | 

| O | O |

x x x x x

x x x x x x x

x x x x x x x

x x x x x x 

x x x x x x x

x x x x x x x x

x x x x x x x x x x x x x x x x x x x x x x x

x x x x x x x x x x x x

x x x x x x x x x x x x x

x x x x x

vvvvvv

vvvv vvvvvv

vvvvv

vvv vvvvvvvvvvvvvv

vv

vvvvvvvvvvvvv

vvvvvvvvvvvvv

UUUUUUUUU

UUUUUUUU

UUUUUUU

UUUUU

UUUU

UU

U

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska
30 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

higher volume and valuable native populations (e.g., Beskid spruce) and ecotypes

(Istebnianski spruce), but they occupy small areas.

Mountain forests, both in the Sudety and Carpathians, are affected by severe

climatic conditions, including strong winds, low temperatures, frequent mists, and high

snowfall. Tree stands, in habitats unsuitable for them and thus weakened, are more

easily broken by wind and snow.

In the last 50 years, the severe climatic conditions have been accompanied by

gaseous (SO2, NOx) and particulate (heavy metals) air pollutants. The Sudety, located

closest to large industrial centers in the Czech Republic, Germany, and Poland, are

among the most threatened and damaged Polish mountains. In the Sudety the average

annual SO2 concentration reaches 25 μg m-3, fluctuating during the year between 16 and

45 μg m-3; average annual NOx reaches 32 μg m-3 (Godzik and Szdzuj 1994). The average

24-h ozone concentrations reaches in the spring month of May 100 μg m-3 (Zwozdziak

and others 1994). Annual sulfur deposition is estimated at about 30 kg/ha-1, and nitrogen

deposition at 8-9 kg ha-1. Atmospheric precipitation is acidic, below pH 4 (Stachurski and

others 1994, Zwozdziak and others 1995). Critical ambient concentrations of sulfur and

ammonia compounds are considerably exceeded in the Sudety forests. In the

westernmost parts of the Carpathians (Silesian Beskid Mountains), air pollutant

concentrations and deposition of these pollutants are nearly the same as those in the

Sudety (Godzik and Szdzuj 1994). Air pollution is considerably lower eastwards (Beskid

Niski and Bieszczady Mountains) because no large emission sources are found in these

areas (Dmuchowski and Wawrzyniak 1994) (figs. 5, 6).

The decrease in the air pollution gradient from west to east (from the Sudety

Mountains to the Carpathians) is confirmed by sensitive plant indicators (Dmuchowski

and Wawrzyniak 1994, Zolnierz and others 1995). Concentrations of and toxic elements

(cadmium and aluminum) in the needles of the spruce and pine (Pinus silvestris L.) are

considerably higher in the Sudety than in the Carpathians (figs. 7, 8).

Mountain forests that have altered species composition, improperly managed and

affected by industrial emissions are attacked by primary insect pests, such as Zeiraphera

griseana, Cephalcia falleni, and Lymantria monacha, and then by secondary pests, such as Ips

typographus and Pityogenes chalcographus as well as by parasite fungi (among others

Armillaria obscura and Heterobasidion annosum). Insects of both groups inflict the greatest

damage in the Sudety and western Carpathians (Capecki 1989, Godzik 1995, National

Inspectorate for Environmental Protection 1993).

Table 1 — Main tree species in forest stands in the Sudety and Carpathian Mountains. Table 1

Forest Stand Picea abies Abies alba Fagus sylvatica

pct

Sudety 83.4 - 5.0

Carpathians 21.7 25.0 25.3

Beskid Slaski 73.6 3.5 17.2

Beskid Zywiecki 74.8 8.8 12.1

Beskid Makowski, Wyspowy 12.9 43.9 17.2

Gorce, Beskid Sadecki 36.9 28.1 25.9

Tatry 77.6 2.8 2.2

Beskid Niski 3.7 31.0 27.0

Bieszczady 9.3 20.0 40.9

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 31

Figure 5 — SO2 deposition index, average value for 1989-1991,

summer period (mg/m2

/24 h) (Dmuchowski and Wawrzoniak 1994).

Figure 6 — NOx

 deposition index, average value for 1989-1991,

summer period (mg/m2

/24 h) (Dmuchowski and Wawrzoniak 1994).

Figure 7 — Total sulfur concentration in spruce needles (Zolnierz

and others 1995).

Figure 8 — Total sulfur concentration (percent in d.wt) in Scots pine

needles (previous year growth) (Dmuchowski and Wawrzoniak 1994).

< 4

4 - 8

8 - 12

>12

μg g-1

2500

2000

1500

1000

500

0

1 2 3 4

S

C

Needle Age

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska

< 0.2

0.2 - 0.4

0.4 - 0.6

>0.6

< 0.120

0.120 - 0.140

0.141 - 0.160

0.161 - 0.180

> 0.180
32 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Fgure 9 — Forest damage in

Poland. Damage degree:

1 = low,

2 = moderate,

3 = heavy,

4 = very heavy,

5 = deforestation (Paschalis 1995).

The Present Health of

Mountain Forests in Poland

According to the general map of forest health by Paschalis (1995), based on data from the

Institute of Forest Research, there were no healthy forests in Poland in 1995. The Sudety

forests and westernmost Carpathian ranges are in a state of total disaster or very

damaged (fig. 9). The forests in the central and eastern Carpathians have been classified

as strongly damaged. Compared to 1990, the state of the forests has deteriorated

considerably (Paschalis 1995).

Factors involved in damage to the mountain forests, which is highest in the western

part and lowest in the eastern part of the Sudety-Carpathian arc include climatic,

geological, and soil conditions; the intensity of anthropogenic impact; the level of air

pollution from industrial emissions; the occurrence of insect pests and parasite fungi

(fig. 10).

In addition to the horizontal east-west gradient of contamination of the mountain

forests, a vertical gradient has been found. Upper montane forests, higher than 1,200 m.

a.s.l., are exposed to greater amounts of wet deposited pollutants than lower ones (<

1,000 m a.s.l.). These pollutants are brought there together with orographic mists and

clouds that occur more frequently at higher elevations than lower elevations (Zwozdziak

and others 1994).

Locally, beautiful stands of forests can be still found in Poland. They are usually

under protection from economic activity in national parks. The country’s seven mountain

national parks are located at regular intervals along the Sudeten-Carpathian arc. Forests

occupy between 67 percent and 91 percent of their areas.

Future of Mountain Forests in Poland

According to forecasts of the Institute of Forest Research based on a large-area inventory

of damage to tree stands, the area of completely and very damaged forests will increase

considerably during the next 15 years (through the year 2010) (Paschalis 1995).

Deterioration of forest health will proceed from west to east (fig. 11).

To counteract the deterioration of mountain forest health, managers and scientists

should reduce both local and transboundary industrial emissions; change existing

forests into less sensitive genotypes, more compatible with local habitats; and

recultivate contaminated forests.

1

2

3

4

5

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 33

Figure 10 — Factors involving

various degree of damage to

mountain forests in Poland.

Figure 11 — Forest damage in

Poland, prognosis 2010. Damage

degree:

1 = moderate,

2 = heavy and very heavy,

3 =deforestation (Paschalis 1995).

POLISH MOUNTAINS

CARPATHIAN MTS.

SUDETY MTS.

50°

24° 20° 16°

50°

W E

continentality

index

soil fertility

human impact

 natural vegetation

 actual forest stand composition

spruce

beech

air pollution

noxious insects

pathogenic fungi

forest damage

1

2

3

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska
34 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The future of mountain forests is a primary concern of Polish ecologists. We are

reminded of the consequences of total forest destruction on the environment by an

admonition more than a dozen years ago by Keating who stated, “The fate of mountain

ecosystems affects half the world’s people” (The Earth Summit Agenda for Change 1983).

Acknowledgments

We are grateful to the hosts of the conference “Air Pollution and Climate Change Effects

on Forest Ecosystems” for financial support that enabled us to participate in the

conference and present this paper on mountain forests in Poland. We thank Laurie Dunn

for technical editing of this manuscript.

References

Capecki, Zenon. 1989. Rejony zdrowotnosci lasow sudeckich. Prace IBL 688:1-93. Rejony zdrowotnosci lasow sudeckich

Denisiuk, Zygmunt. 1995. Ochrona przyrody i krajobrazu (Nature and Landscape Conservation) In: Warszynska, Ochrona przyrody i krajobrazu (Nature and Landscape Conservation)

Jadwiga, ed. Karpaty Polskie -Przyroda, czlowiek i jego dzialalnosc (The Polish Carpathians—nature, man and his

activities). Krakow: Uniwerytet Jagiellonski;131-143.

Dmuchowski, Wojciech; Wawrzoniak,Jerzy. 1994. Spatial distribution of and nitrogen content in needles of Scots Pine

(Pinus sylvestris Pinus sylvestris L.) as related to air pollution and tree stands vitality in Poland. In: Solon, Jerzy; Roo-Zielinska, Ewa; L.) as related to air pollution and tree stands vitality in Poland

Bytnerowicz, Andrzej, eds. Climate and atmospheric deposition studies in forests. Conference Papers 19. Warszawa:

JGSO PAS; 177-186.

Fabijanowski, Jerzy; Jaworski, Andrzej. 1995. Gospodarstwo lesne (Silviculture). In: Warszynska, Jadwiga, ed. Karpaty Gospodarstwo lesne (Silviculture)

Polskie - Przyroda, czlowiek i jego dzialalnosc (The Polish Carpathians—nature, man and his activities). Krakow:

Uniwersytet Jagiellonski; 253-263.

Fabiszewski, Jerzy; Janik, Jan. 1994. Wartosci przyrodnicze i zagrozenia Karkonoskiego Parku Narodowego. Kosmos Wartosci przyrodnicze i zagrozenia Karkonoskiego Parku Narodowego

43(1): 101-115.

Godzik, Stefan; Szdzuj, Jerzy. 1994. A comparison of air, soil, and spruce needle chemistry in Izerskie and Beskidy

Mountains. In: Solon, Jerzy; Roo-Zielinska, Ewa; Bytnerowicz, Andrzej, eds. Climate and atmospheric deposition Mountains

studies in forests. Conference Papers 19. Warszawa: IGSO PAS; 187-195.

Grodzki, W. 1995. Zanieczyszczenia przemyslowe a gradacje szkodnikow owadzich w lasach gorskich. Sylwan 89(5): 13-19. Zanieczyszczenia przemyslowe a gradacje szkodnikow owadzich w lasach gorskich

Matuszkiewicz, Wladyslaw. 1984. Die Karte der Potentiellen Naturlichen Vegetation von Polen. Braun-Blanquetia 1: 1-99. Die Karte der Potentiellen Naturlichen Vegetation von Polen

Panstwowa Inspekcja Ochrony Srodowiska. 1993. Ocena stanu lasow Polski na podstawie badan monitoringowych. Ocena stanu lasow Polski na podstawie badan monitoringowych

Warszawa: Biblioteka Monitoringowa Srodowiska; 71 p.

Paschalis, Piotr. 1995. Stan lasow w Polsce i prognoza zmian. In: Kozlowski, Stefan, ed. Prognoza ostrzegawcza zmian Stan lasow w Polsce i prognoza zmian

srodowiskowych warunkow zycia czlowieka w Polsce na poczatku XXI wieku, Ekspertyza. Zeszyty Naukowe Komitetu

PAN “Czlowiek i Srodowisko” 10; Instytut Ekologii PAN Oficyna Wydawnicza; 191-218.

Stachurski, Antoni; Zimka, Jerzy R., Kwiecien, M. 1994. Niektore aspekty krazenia pierwiastkow w ekosystemach

lesnych na terenie Jakuszyc (II). In: Fischer, Zofia, ed. Karkonoskie Badania Ekologiczne. II Konferencja, Dziekanow lesnych na terenie Jakuszyc (II)

Lesny, 17-19 stycznia 1994. Dziekanow Lesny: Oficyna Wydawnicza Instytutu Ekologii PAN; 207-232.

Starkel, Leszek, ed. 1991. Geografia Polski. Srodowisko Przyrodnicze. Geografia Polski. Srodowisko Przyrodnicze. Warszawa: PWN; 669 p. Geografia Polski. Srodowisko Przyrodnicze.

Szafer, Wladyslaw; Zarzycki, Kazimierz, eds. 1972. Szata roslinna Polski Szata roslinna Polski. Warszawa: PWN; Vol. I:609 p; Vol. II: 347 p. Szata roslinna Polski

Warszynska, Jadwiga, ed. 1995. Karpaty Polskie - Przyroda, czlowiek i jego dzialalnosc (The Polish Carpathians - nature, Karpaty Polskie - Przyroda, czlowiek i jego dzialalnosc (The Polish Carpathians - nature,

man and his activities). Krakow: Uniwersytet Jagiellonski; 367 p. man and his activities

Zarzycki, Kazimierz; Landolt, Elias; Wojcicki, Jan J., eds. 1991. Contributions to the knowledge of flora and vegetation of Contributions to the knowledge of flora and vegetation of

Poland. Veroff. Geobot, Inst, ETH, Stiftung Rubel, Zurich 106: 1-304. Poland

Zwozdziak, Jerzy; Kmiec, G.; Zwozdziak, A.; Kacperczyk, K. 1995. Presja zanieczyszczen przemyslowych w ostatnim

wieloleciu a stan obecny. In: Fischer, Zofia, ed. Problemy Ekologiczne Wysokogorskiej Czesci Karkonoszy. Dziekanow wieloleciu a stan obecny

Lesny: Oficyna Wydawnicza Instytutu Ekologii PAN; 79-96.

Zwozdziak, Jerzy; Zwozdziak, A.; Kmiec, G.; Kacperczyk, K. 1994. Analiza stezen ozonu i skladu frakcyjnego aerozolu

atmosferycznego w Sudetach Zachodnich. In: Fisher, Zofia, ed. Karkonoskie Badania Ekologiczne. II Konferencja, atmosferycznego w Sudetach Zachodnich

Dziekanow Lesny, 17-19 stycznia 1994. Dziekanow Lesny: Oficyna Wydawnicza Instytutu Ekologii PAN; 63-75.

Zolnierz, Ludwik; Fabiszewski, Jerzy; Matula, Jan; Sobierajski, Z.; Wojtun, Bronislaw. 1995. Obraz skazenia fitocenoz

karkonoskich na podstawie badan skladu naturalnego wybranych gatunkow roslin. In: Fischer, Zofia, ed. Problemy karkonoskich na podstawie badan skladu naturalnego wybranych gatunkow roslin

Ekologiczne Wysokogorskiej Czesci Karkonoszy. Dziekanow Lesny: Oficyna Wydawnicza Instytutu Ekologii PAN;

169-185.

Session I State of Polish Mountain Forests: Past, Present, and Future Grodzinska, Szarek-Lukaszewska
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 35

Paul Miller,2 Raleigh Guthrey,2 Susan Schilling,2 John Carroll3

Abstract

Ozone injury was monitored on foliage of ponderosa (Pinus ponderosa Dougl. ex Laws.) and Jeffrey (Pinus

jeffreyi Grev. & Balf.) pines at 11 locations in the Sierra Nevada and 1 site in the San Bernardino Mountains of

southern California. Ozone injury on all age cohorts of needles on about 1,600 trees was surveyed annually from

1991 to 1994. A new method for describing ozone injury to whole tree crowns, the ozone injury index (OII), was field

tested and improved. The OII ranged from 0 (no injury) to 100 (the maximum possible injury). The OII multi-year

average was only 5 at Lassen Volcanic Park in rural northern California and gradually increased in a southward

direction west of the Sierra Nevada crest to moderate amounts (28-41) in the Sequoia National Forest and Sequoia

National Park. The OII multi-year average measured at the San Bernardino Mountain site was 46. An assessment

of annual changes at the 12 individual sites indicated both increases and decreases in OII from 1991 to 1994. The

two most responsive indicators of the annual increments of accumulated injury, contributing 40 percent each, were

chlorotic mottle and needle fascicle retention (within remaining needle whorls). Data for these components were

tested with quadratic and Weibull functions against several expressions of ozone exposure (including ozone

exposure indices Sum 0, Sum 60, W126, and number of hours exceeding 80 ppb during the summer exposure

periods). Sum 0 was a suitable exposure index having Weibull correlation coefficients of 0.57 with percent chlorotic

mottle and 0.74 with percent fascicle retention. These results provide estimates of ozone injury responses across a

range of annual accumulated ozone exposures and environmental conditions during four summers.

Introduction

In 1990-91 two companion projects were begun at analogous locations in the Sierra

Nevada of California: the Sierra Cooperative Ozone Impact Assessment Study

(SCOIAS) and the Forest Ozone Response Study (FOREST). Funded by a contract with

the California State Air Resources Board, the principal activity of SCOIAS was to

monitor ozone and meteorological variables at six Sierra Nevada sites managed by the

University of California, Davis. FOREST was developed as a companion project to

SCOIAS through an agreement of intent between the USDA Forest Service, Pacific

Southwest Region , and the California State Air Resources Board.

The FOREST agreement led to the establishment of forest vegetation plots within

3 miles distance and 500 ft elevation of SCOIAS monitoring stations for the purpose of

annual assessments of ozone injury to ponderosa and Jeffrey pine populations. The Air

Resources Management Staff of the USDA Forest Service’s Pacific Southwest Region

provided training of field crews and actual data gathering activities in six National

Forests. Other participants included Yosemite, Sequoia-Kings Canyon, and Lassen

Volcanic National Parks, and the Forest Service’s Pacific Southwest Research Station.

The participants used accepted procedures for instrument calibration and maintenance

for ozone measurements and tree injury assessment (Miller and others 1996b). The

Pacific Southwest Research Station provided data management, data analysis and

reporting services for FOREST (Guthrey and others 1993, 1994).

This report describes the use of the ozone injury index (OII) to monitor ozone

injury to ponderosa pine (Pinus ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus

jeffreyi Grev. and Balf.) in the Sierra Nevada and San Bernardino Mountains in 1991-

1994; and it provides results of tests that analyzed the relationship of annual

accumulated ozone exposure expressed as Sum 0, Sum 60 and W126 to the intensity of

chlorotic mottle and the retention of needle fascicles in all remaining needle whorls.

Ozone Injury Responses of Ponderosa and

Jeffrey Pine in the Sierra Nevada and San Bernardino

Mountains in California1

1 An abbreviated version of this paper

was presented at the International

Symposium: Air Pollution and Climate

Change Effects on Forest Ecosystems,

February 5-9, 1996, Riverside,

California.

2 Plant Pathologist and Computer Programmers, respectively, Pacific Southwest Research Station, USDA Forest

Service, 4955 Canyon Crest Drive, Riverside, CA 92507.

3 Meteorologist, Department of Land,

Air and Water Research, University of

California, Davis, CA 95616.
36 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Methods

Ozone monitoring stations established as part of SCOIAS were located at six

Sierra Nevada sites including National Forests, a State Park, and private land

(Van Ooy and Carroll 1995). Ozone data were also available from the National

Park Service at Lassen Volcanic, Yosemite, and Sequoia-Kings Canyon National

Parks and from the Forest Service at Barton Flats in the San Bernardino

Mountains. At each site, three forest plots with at least 40-50 tagged ponderosa

or Jeffrey pines were established within 5 km and 150 m elevation of the

monitoring stations (fig. 1). Trees were sampled annually by pruning five

branches each from about 1,700 trees distributed in 36 field plots. The ozone

injury index (OII) was computed for each tree based on four weighted variables:

number of annual needle whorls retained (40 percent), amount of chlorotic

mottle (the ozone injury symptom) on needles of each whorl (40 percent), length

of needles in each whorl retained (10 percent), and percent live crown (10

percent). The OII ranged from 0 (least severe injury) to 100 (most severe injury)

(Miller and others 1996b).

Two of the attributes included in the computation of the OII, amount of

chlorotic mottle (percent of the total needle surface) on needles in each whorl and

the percent of the original complement of needle fascicles retained in each whorl,

were considered the best indicators of the annual incremental increases of ozone

injury to ponderosa and Jeffrey pine foliage. A schedule was established for

computation of accumulated ozone exposure for each age cohort of needles

Figure 1 — Map locations of

ozone monitoring sites and

associated plots for monitoring

ozone injury to ponderosa or

Jeffrey pines.

LV = Lassen Volcanic,

WC = White Cloud,

SP = Sly Park,

LC = 5 mile Learning Center,

JD = Jerseydale,

YM = Yosemite Mather,

YW = Yosemite Wawona,

SL = Shaver Lake,

SF = Sequoia Giant Forest,

SG = Sequoia Grant Grove,

MH = Mountain Home,

BF = Barton Flats.

LV

120

40

35

BF

WC

SP

LC

JD YM

YW

SL

SF SG

MH

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 37

(table 1). Summer-season exposure was evaluated for each cohort, from

August 1 to the injury evaluation date. August 1 was determined as the estimated

date when current year needles had grown to maximum length and had

measurable stomatal conductance (Temple and Miller, [this volume]). Since the

SCOIAS sites were established in 1991, a full summer season of ozone data was

available for only a few sites in 1991. A record of dates of injury evaluation was

available for trees at each set of three plots near each monitoring station.

Evaluation of all three plots ranged from 1 to 2 weeks. The beginning of the

evaluation period was selected as the termination of the exposure period that

had caused observed injury in that season. The exposure was initially calculated

from August 1 to the first day of the foliar injury evaluation in the origin year of

the needle whorl. Accumulation of ozone exposure for the cohort’s second year

was continued after evaluation and until October 31. Accumulation was resumed

from June 1 each year until the next date of foliar symptom evaluation. This

procedure was carried out for each annual needle age cohort (1991-94) until the

beginning of the last period of the symptom evaluation in 1994, which had only

the 1994 season of ozone exposure. The ozone exposure indices which were

computed for each annual or consecutive annual time interval were Sum 0, Sum

60 (ppb), W126 (Lefohn 1992), and number of hours > 80 ppb. The sum of the

number of days in each seasonal exposure was included as a separate (dummy)

variable. This variable simulates the extreme case in which daily ozone exposure

is the same every day.

For each annual injury data set the amount of chlorotic mottle present

was recorded as 0 (0 percent), 1 (1-6 percent), 2 (7-25 percent), 3 (26-50

percent), 4 (51-75 percent), and 5 (76-100 percent). For the regressions these

ranges were represented by their midpoints (e.g., 0 = 0 percent, 1 = 3.5 percent, 2

= 16 percent, 3 = 38 percent, 4 = 63 percent, and 5 = 88 percent). Similarly, the 0 (0

percent), 1 (1-33 percent), 2 (34-66 percent), and 3 (67-100 percent) recorded

values for percent of needle fascicles retained in each whorl were represented as

0 = 0 percent, 1 = 17 percent, 2 = 50 percent, and 3 = 83.5 percent.

The Weibull function was selected as the most appropriate model for relating

the ozone exposure indices to foliar injury measurements (Rawlings and others

1988). The Weibull function was used successfully in the analysis of the ozone

exposure of annual crop plants (U.S. Environmental Protection Agency 1986). It

expresses the relationship between plant injury and ozone exposure as:

Y = αexp[-(x/σ)c

]

in which α = response at 0 O3

 exposure; x = accumulated O3 exposure (in ppbhrs); σ = accumulated ozone exposure at which α is reduced by 63 percent;

and c = a dimensionless shape parameter.

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll

Table 1 Table 1 — The schedule for computation of accumulated ozone exposure and evaluation of needle injury (E) Table 1 

at 12 ozone-monitoring, tree plot locations in California, 1991-94.

Origin year of Origin year of Periods of the summer each year when ozone exposure was

accumulated needle whorl before and after injury evaluation (-E) needle whorl before and after injury evaluation (-E)

1991 1992 1993 1994

1991 8/1-E-10/31 6/1-E-10/31 6/1-E-10/31 6/1-E-10/31

1992 8/1-E-10/31 6/1-E-10/31 6/1-E-10/31

1993 8/1-E-10/31 6/1-E-10/31

1994 8/1-E-10/31
38 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Results

Ozone Monitoring from 1991-1994

Although average ozone concentrations varied at stations within a given subregion, e.g., the central Sierra Nevada (Carroll and Dixon 1993, Van Ooy and

Carroll 1995), average concentrations from north (Lassen Volcanic National

Park) to south (San Bernardino Mountains) increased. Thus, we found an

adequate exposure gradient to examine the relationship between ozone exposure

and the injury response of pines.

OII Values Along a North to South Transect — 1992-1994

The least injury is present in the northern to central Sierra Nevada (fig. 2a) and

the most in the south central Sierra Nevada and the San Bernardino mountains

(fig. 2b). The observed injury amounts are generally proportional to seasonal

ozone exposures reported by Van Ooy and Carroll (1995) and by Miller and

others (1996a). The combined ozone and OII data sets are considered only

marginally useful for providing adequate detail of the spatial distribution of

injury because the locations of ozone monitors and tree plots could not be

randomly selected. The change in yearly OII at any single site is considered a

satisfactory indicator of exposure response because it was a repeated

measurement of the same trees. Between 1991 and 1994 at the northern locations

the OII rose slightly at four of six locations and was essentially unchanged at

each of the two remaining locations (fig 2a). At the six southern locations OII

declined slightly (less injury at all places where the record was complete) except

at Barton Flats where it increased slightly (fig. 2b).

Figure 2a — Annual changes in

the ozone injury index (OII) from

1991 to 1994 at six northern or

central California sites: Lassen

Volcanic , White Cloud, Sly Park,

Learning Center, and Camp

Mather and Wawona in Yosemite

National Park.

Figure 2b — Annual changes in

the ozone injury index (OII) from

1991 to 1994 at five south-central

Sierra Nevada sites and one

southern California site:

Jerseydale, Shaver Lake, Grant

Grove and Giant Forest in

Sequoia-Kings Canyon National

Park, Mountain Home State Park,

and Barton Flats in the San

Bernardino National Forest. Data

are incomplete for Jerseydale

from 1992 to 1994 because of

unusual amounts of bark beetlecaused mortality.

OII Units

Lassen Wh. Cloud Sly Park Learn Cn Mather Wawona

1991

1992

1993

1994

60

50

40

30

20

10

0

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll OII Units

Jerseydale Shav. Lk. Grnt. Grv. Giant For. Mtn. Home Bart. Flats

1991

1992

1993

1994

60

50

40

30

20

10

0
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 39

Correlation of Percent Chlorotic Mottle with Several

Ozone Exposure Indices and Cumulative Days of Exposure

The year by year change of percent chlorotic mottle is illustrated as a function of

accumulating ozone exposure received by current-year, 1-year-old, 2-year-old,

and 3-year-old needles. The data were tested with a number of fitted curves,

including quadratic and a Weibull function. The latter was selected for this

application because it is a monotonically decreasing (or increasing) fit that more

likely applies to the expected biological response in this case. The r2 for both

quadratic and Weibull fits were computed. The corrected regression coefficients

for percent chlorotic mottle and each of the exposure indices were:

Index Quadratic Weibull

Sum 0 .583 .574

Sum 60 .592 ——

W126 .596 ——

Hrs > 80 .553 ——

Expos. Days .433 .424

The r2

 for Sum 0 was not the highest value for quadratic fits but it was the only

form of accumulated ozone exposure that converged to the required parameters

needed for the Weibull fit. These results point to Sum 0 as an acceptable exposure

index to apply in this situation. Therefore, the Weibull equation for percent

chlorotic mottle (PCM) and Sum 0 was:

 PCM = 100-100*EXP(-(Sum 0/1579000) 2.170)

During the 1991 to 1993 seasons the Sum 0 accumulated exposure ranged

between 0 and 600,000 ppb-hrs (fig. 3). It was not feasible to observe a needle

whorl for longer than three seasons (e.g., 1991 needles could not be used in 1994)

because many needle fascicles would have abscised and those few remaining

would have less chlorotic mottle. These results indicate that Sum 60, W126, and

hrs > 80 are not feasible to use because these data did not converge to the

required parameters needed for the Weibull curve fit. Accumulated number of

exposure days had the lowest coefficient.

Figure 3 — Weibull curve fit of

percent chlorotic mottle

accumulated by 1991 to 1993

whorls in relation to Sum 0 ozone.

Percent chlorotic mottle

Ozone dose: SUM 0

0 1 105 2 105 3 105 4 105 5 105 6 105

r2 =.574

15

10

5

0

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll
40 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Correlation of Percent Fascicle Retention with Several

Ozone Exposure Indices and Cumulative Days of Exposure

As chlorotic mottle develops on needle fascicles, the natural abscission of needle

fascicles within each annual whorl increases thereby leaving whorls with fewer

needle fascicles until eventually all needle fascicles abscise and the entire whorl

is gone. The corrected r2 values show Sum 0 to be the Weibull fits:

Index Quadratic Weibull

Sum 0 .759 .742

Sum 60 .675 .613

W126 .653 .609

Hrs > 80 .443 .411

Expos. Days .692 .669

Because the expected biological response corresponds to the monotonically

decreasing form of the Weibull function, it was selected for relating percent

fascicle retention (PFR) to the selected exposure indices, e.g., in the case of Sum 0:

 PFR = 83.5*EXP(-(Sum 0/732600) 3.057)

For the Weibull fit of Sum 0 data versus percent fascicle retention (fig. 4) the

maximum amount for Sum 0 exposure is 800,000 ppb-hr as compared to 600,000

ppb-hr for percent chlorotic mottle because it was feasible to use a longer time

span (1991-94) for percent fascicle retention than for percent chlorotic mottle

(1991-93).

Three Dimensional Display of the Sum 0 Exposure Index, and the

Frequency of Sampled Trees in Original Classes of Chlorotic Mottle

Because the Weibull function correlations of Sum 0 versus percent chlorotic

mottle involve transformed data, such as the midpoint value of each class, 1 = 1-

6 percent (3.5 percent) and 2 = 7-25 percent (16 percent) etc., we examined the

distribution of the untransformed data. We related the frequency of trees to the

original chlorotic mottle classes as a function of Sum 0 exposure index (fig. 5).

The threshold above which category 1 (1 to 6 percent) chlorotic mottle appears is

about 150,000 ppb. The equivalent threshold values for category 2 (7 to 25

percent) chlorotic mottle could be more useful as a category to monitor because

needle fascicle abscission begins at this level.

Figure 4 — Weibull curve fit of

percent needle fascicle retention

accumulated by 1991 to 1994

whorls in relation to Sum 0

ozone.

Percent fascicle retention

Ozone dose: SUM 0

0 1 105 2 105 3 105 4 105 5 105 6 105 7 105 8 105

r2 =.759

80

70

60

50

40

30

20

10

0

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 41

Discussion

The time resolution for examining pine needle response to ozone can vary from

daily examination of physiological variables (Grulke and Lee, [In press]; Temple

and Miller, [this volume]) to annual visual ratings of injury. For the purpose of

evaluating annual increments of injury increase and comparing yearly changes,

the visual rating method used in this study, the OII (Miller and others 1996b),

appears to provide a reliable estimate. It could be improved by the inclusion of

an estimate of the seasonal pattern of stomatal conductance and an estimate of

ozone flux to pine foliage, but this improvement generally requires concurrent

field measurements of stomatal conductance. Attempts to apply models of ozone

flux that are parameterized primarily by atmospheric conditions (temperature

and relative humidity) have realized very limited success as predictors

(Fredericksen and others 1996) when compared to flux estimates based on actual

conductance measurements.

Because of the relative simplicity of the OII, we recommend it as a suitable

protocol for monitoring yearly injury. In our study we have used a large number

of trees (1,700) to evaluate the visual change of individual age cohorts of needles

on each tree exposed to ozone in their initial year and in successive years.

Certainly the climate and soil-water availability conditions from 1991 to 1994

were variable and caused important yearly differences in ozone uptake and

incremental changes of needle injury amounts. However, annual differences

were detected by the chlorotic mottle element of the OII (Temple and Miller, [this

volume]), and to a lesser degree by the OII itself. In further support of the visual

estimation procedure of the OII, it was possible to include enough trees to

represent the range of genetic resistance or susceptibility of ponderosa and

Jeffrey pines as well as trees in different crown position classes (e.g. dominant,

codominant, etc). Large sample numbers are usually difficult to achieve with

physiologically-based response variables (Grulke and Lee [In press]). By using

the OII or its most sensitive components, we can examine further the different

versions of a possible air quality standard that may be appropriate for the task of

protecting ozone-sensitive tree species.

Figure 5 — Distribution of 1991

to 1994 annual whorls from all

trees into original categories for

chlorotic mottle as a function of

Sum 0 ozone exposure index

during the same period.

Chlorotic mottle

Ozone dose: SUM 0

800

600

400

N

200

0

0

250,000

500,000

750,000

1

2

3

4

5

6

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll
42 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Acknowledgments

This study was sponsored in part by an interagency agreement with the U.S.

Environmental Protection Agency, AREAL, Research Triangle Park, NC.,

Deborah Mangis, agreement monitor. We thank Dan Duriscoe, Judy Rocchio,

Diane Ewell and other National Park Service (NPS) staff from several other Parks

who received training and collected annual data at the vegetation plots. The

Parks ozone data was obtained from the Air Quality Division, NPS. We thank

Kenneth Stolte, Forest Health Monitoring Program, USDA Forest Service; Trent

Procter, Pacific Southwest Region, Forest Service, Air Resources Management

staff for training and organization, and the tree injury survey crews from the

Eldorado, Sequoia, Sierra, Stanislaus and Tahoe National Forests. Jules Riley,

Stanislaus, did quality assurance measurements; Brent Takemoto of the

California State Air Resources Board, evaluated data. David Randall, Pacific

Southwest Research Station, Forest Service, provided statistical advice. We thank

Laurie Dunn for technical editing of this manuscript.

References

Carroll, J. J.; Dixon, A. J. 1993. Sierra cooperative ozone impact assessment study: year 3. Final Report: Interagency Sierra cooperative ozone impact assessment study: year 3

Agreement # A132-188. Volume 1. Sacramento, CA: California Air Resources Board; 74 p.

Fredericksen, T. S.; Skelly, J. M.; Snyder, K. R.; Steiner, K. C.; Kolb, T. E. 1996. Predicting ozone uptake from

meteorological and environmental variables. Journal of Air and Waste Management Association 46: 464-469. meteorological and environmental variables

Grulke, Nancy E.; Lee, E. Henry. [In press] Correlation of morphological, physiological and nearest neighbor

characteristics with visible ozone damage in ponderosa pine. Canadian Journal Forestry Research. characteristics with visible ozone damage in ponderosa pine

Guthrey, D. Raleigh; Schilling, Susan L.; Miller, Paul R. 1993. Initial progress report of an interagency forest monitoring

project: forest ozone response study (FOREST). Riverside, CA: Pacific Southwest Research Station, USDA Forest project: forest ozone response study (FOREST)

Service; 37 p.

Guthrey, D. Raleigh; Schilling, Susan L.; Miller, Paul R. 1994. Second progress report of an interagency forest monitoring

project: forest ozone response study (FOREST). Riverside, CA: Pacific Southwest Research Station; 13 p. project: forest ozone response study (FOREST)

Lefohn, A. S. 1992. The characterization of ambient ozone exposures. In: Lefohn, A. S., ed. Surface level ozone exposures The characterization of ambient ozone exposures

and their effects on vegetation. Chelsea, MI: Lewis Publishers, Inc; 31-92.

Miller, Paul R.; Chow, Judith R.; Watson, John G.; Bytnerowicz, Andrzej; Fenn, Mark; Poth, Mark; Taylor, George.

1996a. Final Report, Contract A032-180, Assessment of acidic deposition and ozone effects on conifer forests in the

San Bernardino mountains, Sacramento CA: California State Air Resources Board; 650 p. San Bernardino mountains

Miller, Paul R.; Stolte, Kenneth W.; Duriscoe, Daniel; Pronos, John, technical coordinators. 1996b. Evaluating ozone air

pollution effects on pines in the western United States. pollution effects on pines in the western United States. Gen. Tech. Rep. PSW-GTR-155. Albany, CA: Pacific Southwest pollution effects on pines in the western United States.

Research Station, USDA Forest Service; 79 p.

Temple, Patrick J.; Miller, Paul R. 1998. Seasonal influences on ozone uptake and foliar injury to ponderosa and Jeffrey

pines at a southern California site, In: Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan, technical pines at a southern California site

coordinators. Proceedings of the international symposium on air pollution and climate change effects on forest

ecosystems. 1996 February 5-9; Riverside, CA. Gen. Tech. Rep. PSW-GTR-166. Albany, CA: Pacific Southwest

Research Station, USDA Forest Service; [this volume].

U.S. Environmental Protection Agency. 1986. Air quality criteria for ozone and other photochemical oxidants. Research Air quality criteria for ozone and other photochemical oxidants

Triangle Park, NC: Office of Health and Environmental Assessment, Environmental Criteria Assessment Office; EPA

Report no. EPA-600/8-84-020aF-ef. Five volumes. Available from NTIS. Springfield, VA; PB87-142949.

Van Ooy, D. J.; Carroll, J. J. 1995. The spatial variation of ozone climatology on the western slope of the Sierra Nevada. The spatial variation of ozone climatology on the western slope of the Sierra Nevada

Atmospheric Environment 29: 1319-1330.

Session I Ozone Injury Responses of Ponderosa and Jeffery Pine Miller, Guthrey, Schilling, Carroll
SESSION II

Air Pollution Status of Forest Sites

Paul R. Miller and Juri Martin, Chairs

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 45

Ozone Air Pollution in the Ukrainian Carpathian Mountains

and Kiev Region1

Abstract

Ambient concentrations of ozone (O3 ) were measured at five highland forest locations in the Ukrainian Carpathians

and in two lowland locations in the Kiev region during August to September 1995 by using O3

 passive samplers. The

ozone passive samplers were calibrated against a Thermo Environmental Model 49 ozone monitor located at the

Central Botanical Garden in Kiev. The 2-week long average concentrations in August to September at the Carpathian

Mountains ranged from 27.4 to 51.8 ppb, and at the lowland forest location (Lutezh, near Kiev) ranged from 29.3 to

32.2 ppb. The 2-week long average concentrations in Kiev (Botanical Garden) ranged from 22.0 to 31.5 ppb. The

highest diurnal average concentration in August 1995 in the Botanical Garden was 42.1 ppb while the highest 1-hr

average concentration reached 84.4 ppb. Ozone-sensitive Bel-W3 tobacco (Nicotiana tabacum L.) plants at the

Botanical Garden in Kiev were injured from exposure to ambient concentration of ozone. Ozone injury symptoms were

found on native plants (e.g., Sambucus racemosa and Humulus lupulus) elsewhere in Kiev and at some of the study

sites in the Carpathians.

Introduction

In general, elevated concentrations of sulfur and nitrogen oxides have been blamed

for air pollution-related damage to forest vegetation in central and eastern Europe.

However, increasing concentrations of ozone may also play an important role in

the observed suite of symptoms of forest decline in that part of Europe. Ozone

alone may affect plant health, while the effects of mixtures of ozone with other

pollutants, especially with sulfur dioxide, may be synergistic (more than additive)

in nature (Guderian 1985).

Recently, ozone concentrations have significantly increased across Europe

mainly because of increased production of ozone precursors from combustion of

gasoline and other fuels (Derwent and Jenkin 1991). Long range atmospheric

transport may also be responsible for elevated concentrations of ozone in various

forested areas of the region (Dovland 1987).

In Ukraine the annual atmospheric emissions of gaseous pollutants are: NOx

(nitrogen oxides) — 1.1 million tons; CO (carbon monoxide) — 8 million tons;

Cx

Hy (hydrocarbons) — 1.4 million tons (National Report 1994). As a result of

photochemical transformation of these emissions, the level of surface background

concentrations of ozone could theoretically increase two-fold compared to the preindustrial pollution period (Nikolay Gurevich, personal communication).

Average 24-hour background O3 concentrations in the unpolluted lower

troposphere ranged from 10 to 40 ppb (Logan 1985). It has been estimated that in

the countries of the former USSR the background O3 concentrations ranged from

15 to 40 ppb (Israel 1984). Concentrations of ozone in European countries of higher

latitudes, such as Finland (Laurila and Lattila 1993) or Lithuania (Girgzdiene 1991),

are relatively low and do not pose a threat to vegetation. However, higher ozone

concentrations, potentially phytotoxic, occur in the lower latitudes of western and

central Europe (Emberson and others 1996, Semenov and Kouhta 1996).

Still relatively little is known about concentrations of ozone and potential toxic

effects of ozone in forests in the former Communist countries of central and eastern

Europe, particularly in Ukraine. Levels of ozone in central and western European

countries of comparable climatic conditions such as Poland (Bytnerowicz and others

1993, Godzik 1996), Czech Republic (Bytnerowicz and others 1995), Austria

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Lichenologist and Biochemist, respectively, Central Botanical Garden, National Academy of Sciences,

Timiryazevskaya St. 1, 252014 Kiev,

Ukraine.

3 Forest Ecologist, Pacific Southwest Research Station, USDA Forest Service,

4955 Canyon Crest Drive, Riverside,

CA 92507, USA.

4 Phytopathologist, Department of Microbiology, University of Massachusetts, Fernald Hall BOX 32420,

Amherst, MA 01003-2420, USA.

Oleg Blum,2 Andrzej Bytnerowicz,3 William Manning,4 and Ludmila Popovicheva2
46 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

(Dovland 1987), Switzerland (Ballaman 1993), or France (Proyou and others

1991) often exceed permissible levels for that pollutant. For example, in forested

areas of central and southern Poland 1-hour mean ozone concentrations can be as

high as 105 ppb (Bytnerowicz and others 1993). During long periods of hot weather

and high solar radiation, the highest 1-hour mean ozone concentrations monitored

in Katowice (the Upper Silesia Region of Poland) have reached 115 ppb (Godzik

and others 1994).

Information about the air pollution status of the Ukrainian Carpathians is

essential for a better understanding of environmental stresses that affect valuable

forest ecosystems of central Europe. Thus, level of ozone, one of the main

components of photochemical smog and a strong phytotoxic agent, is a primary

area of study.

Until recently, surface background concentrations of ozone in natural

landscapes of Ukraine were not measured, with the exception of O3 monitoring in

the territory of Karadag State Reserve on a coast of the Black Sea of the Crimea. At

that site, the 24-hour average O3 concentrations in the spring-summer period of

about 15 ppb and 1-hour average O3 maximum concentrations of about 105 ppb

have been recorded.5

This paper discusses results of a pilot study that used ozone-sensitive Bel-W3

tobacco and other plants to measure ozone concentrations in Ukraine.

Materials and Methods

Research Sites

In summer 1995 research sites were established in five forest locations in the

Ukrainian Carpathians (fig. 1) in the forest complex of the northern Ukraine

(Staropetrovskaya Forest Research Station, vil. Lutezh, near Kiev) and at two

locations, one the northern and one in the southern part, in the Central Botanical

Garden of the National Academy of Sciences in Kiev (50°24’47” N, 30°34’14” E;

190 m a.s.l.).

Forest sites were located at altitude 750-1,000 m a.s.l. along the Carpathian

range from the west to south-west: 1 — Uzhoksky Pass (Sjanki village, 850 m a.s.l.);

2 — Synevir (National Park “Synevir,” 1,000 m a.s.l.); 3 — Shiroky Lug (Carpathian

Biosphere Reserve, Kuzij massif, peak Sokolyne Berdo, 750 m a.s.l.); 4 — Yablunitsa

village, Yablunetski Pass (Carpathian National Park, 968 m a.s.l.); 5 — Kryvopilja

village (Carpathian National Park, 975 m a.s.l.).

5 Unpublished data on file, Gas Institute, National Academy of Sciences of

Ukraine

Figure 1 — Ozone monitoring

sites in the Carpathian Mountains

in Ukraine:

1 — Uzhoksky Pass,

2 — Synevir,

3 — Shiroky Lug,

4 — Yablunitsa,

5 — Kryvopilja.

Session I1 Ozone Air Pollution in the Ukrainian Carpathian Blum, Bytnerowicz, Manning, Popovicheva

IvanoFrankivsk

4 5

3

2

1

Romania

Hungary

Czech

Republic

Slovakia

Poland

M o u n t a i n s

0 15 30 45 Km

Rakhiv

Mukacheve

Uzhhorod U k r a i n e

C a r p a t h i a n
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 47

Ozone Monitoring

Average ambient concentrations of O3 were determined with the Ogawa and Co.,

Inc., passive ozone samplers6␣ (Koutrakis and others 1993). The samplers were

exposed to ambient air for a duration of 7 to 14 days. Measurements were carried

out in all forest locations from the beginning of August to the end of September.

In the Central Botanical Garden (Kiev) measurements of O3 with passive

samplers were accompanied by continuous monitoring with the Thermo

Environmental Model 49 ultraviolet absorption instrument (Cambridge, MA).

Results from passive ozone samplers were compared to those obtained from the

Thermo Environmental instrument, which had been calibrated before its use.

Ozone Phytotoxicity

Cultivated and native plants were surveyed for symptoms of ozone injury at all

five Carpathian forest locations and in Kiev. To determine the phytotoxicity of

ambient O3, two cultivars of tobacco (Nicotiana tabacum L.) were used: Bel-W3 (O3-

sensitive), and Bel-B (O3-tolerant) (Heggestad 1991). Before field exposures, all

seedlings were grown in charcoal filtered air to the 4-leaf stage of development in a

special growing substrate (pH = 6.5), consisting of peat, perlite, and gypsum (1:1:1

by volume).

The ozone-free air chamber made of transparent plastic (1.1 m3 volume) was

constructed in order to grow tobacco plants. The chamber was located in the

laboratory room. Two incandescent lamps (500 W) wrapped with aluminum foil

(for darkening) and a thermo-relay kept constant temperature at about 32 - 35 °C.

Air entering the chamber was pumped at a flow rate of 30 l/min. through a charcoal

filter. Cuvettes with water placed at the bottom of the chamber provided the

required humidity for proper growth of the tobacco plants. The plants were grown

under natural and luminescent lighting (light/dark regime — 16:8 hours).

Injury of the Bel-W3 tobacco plants in the Central Botanical Garden was

observed in two locations from early August to early September 1995. At one of

the research sites, measurements with O3 passive samplers and bioindicators were

accompanied by photometric monitoring of ambient O3 concentrations (near

Building No. 5, Department of Plant Physiology). At another location, (i.e., the

Palmetto Orchard) measurements were done by using only O3 passive samplers

and the bioindicator tobacco plants.

Results and Discussions

Ozone Concentrations

The 2-week long average concentrations in August to September in the Carpathian

Mountains sites ranged from 27.4 to 51.8 ppb, while at Lutezh, the lowland forest

location near Kiev, they ranged from 29.3 to 32.2 ppb. The 2-week long average

concentrations in Kiev (Botanical Garden) ranged from 22.0 to 31.5 ppb (table 1).

The highest 24-hour average concentration measured with the Thermo

Environmental instrument in August 1995 in the Botanical Garden was 46.7 ppb,

while the highest 1-hr average concentration reached 84.4 ppb. The lowest 1-hour

average concentrations were as low as 7.0 ppb (fig. 2).

In Kiev, typically, the lowest O3 levels occurred in the morning with highest

concentrations found in the afternoon. Average concentrations of O3 at night were

about 25 ppb, while the maximum values reached about 47 ppb (fig. 3).

In general, O3 concentrations in the five mountain forested locations in the

Ukrainian Carpathians were low or only moderately elevated. The determined

concentrations were similar or slightly lower than the values determined in the

forested locations of central and southern Poland (Bytnerowicz and others 1993,

Godzik 1996) and the Czech Republic (Bytnerowicz and others 1995). Ozone results

obtained with the Ogawa passive samplers in areas of low ozone concentrations

have about ± 20 percent sampling accuracy (Koutrakis and others 1993).

6 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Session I1 Ozone Air Pollution in the Ukrainian Carpathian Blum, Bytnerowicz, Manning, Popovicheva
48 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Table 1 — Concentrations of ozone determined with the Ogawa passive samplers in summer 1995 in the Ukrainian Table 1

locations (ppb).

 Location August 6-19 August 19-Sept. 4 September 4-18

1. Uzhoksky Pass 40.9 36.3 34.2

2. Synevir 39.5 38.3 27.4

3. Shiroky Lug 27.4 33.1 51.8

4. Yablumitsa - 36.1 -

5. Kryvopilja - 35.8 -

6. Kiev, Palmetto 22 31.5 -

7. Kiev, Bldg. No. 5 30 29.8 25.7

8. Lutezh 29.3 32.2 -

Ozone Phytotoxicity

Ozone injury symptoms were present on both Bel-W3 (ozone sensitive cultivar)

and Bel-B (ozone tolerant cultivar) tobacco plants exposed to ambient air in the

Botanical Garden (Kiev). The highest amount of O3 injury was found on the Bel-W3

plants at the Palmetto Orchard and near the Building No. 5 (table 2) reaching 62

percent of the foliar area on the first leaf of plants after 14-day exposure. Moderate

injury on the Bel-W3 plants (11-20 percent of the leaf area) occurred at the Building

No. 5 location after 7-day exposure. Slight O3 injury was also observed on the BelB tobacco plants (ozone tolerant cultivar, 1-4 percent of the leaf area) in the Building

No. 5 location after 14-day exposure. Higher degree of injury on the Bel-B plants

(up to 13 percent of the leaf area) occurred at the Palmetto Garden location

after 14-day exposure.

Figure 2 — Results of O3 monitoring in the Botanical Garden

in Kiev during the August 4 to September 21, 1995 period.

Results are shown as 24-hour averages, and 1-hour maximum

and 1-hour minimum values for every day during the

monitoring period (ppb).

Figure 3 — Diurnal dynamics of O3

 concentrations in the

Central Botanical Garden, National Academy of Sciences in

Kiev, shown as averages for the August 4 to September 21,

1995 period. Results are shown as mean, maximum, and

minimum values for every hour.

Ozone Concentration (ppb)

Date

Maximum

Average

Minimum

4.086.088.08

10.0812.0814.0816.0818.0820.0822.0824.0826.0828.0830.08

1.093.095.097.099.09

11.09

13.0915.0917.0919.0921.09

80

90

70

60

50

40

30

20

10

0

Hourly average ozone concentration (ppb)

Time

Maximum

Average

Minimum

0.302.304.306.308.30

10.3010.3014.3016.3018.3020.3022.30

80

90

70

60

50

40

30

20

10

0

Session I1 Ozone Air Pollution in the Ukrainian Carpathian Blum, Bytnerowicz, Manning, Popovicheva
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 49

Table 2 Table 2 — Injury of the Bel-W3 and Bel-B tobacco plants at two locations in the Central Botanical Garden, Kiev Table 2 

(percent of leaf area injured presented as means and standard deviation; n=4).

Location Leaf 7 days 14 days 7 days 14 days

 No. of exposure of exposure of exposure of exposure

(Aug. 7-14) (Aug. 7-14) (Aug. 7-21) (Aug. 7-21) (Aug. 22-29) (Aug. 22-29) (Aug. 22-Sept. 6) (Aug. 22-Sept. 6)

Bel-W3

Palmetto 1 37 ± 9 40 ± 11 30 ± 12 62 ± 1

Orchard 2 32 ± 14 35 ± 15 23 ± 8 57 ± 11

3 7 ± 3 25 ± 5 5 ± 3 11 ± 31

4 0 4 ± 3 0 2 ± 6

Building 1 11 ± 5 18 ± 6 19 ± 9 39 ± 16

No. 5 2 20 ± 9 27 ± 8 17 ± 7 40 ± 8

3 2 ± 1 6 ± 4 9 ± 5 18 ± 13

4 0 1 ± 1 0 0.5 ± 0.9

 Bel-B

Palmetto 1 0 0 6 ± 2 13 ± 6

Orchard 2 0 0 1 ± 1 11 ± 7

30 0 0 0

40 0 0 0

Building 1 0 0 0 4 ± 2

No. 5 2 0 0 0 1 ± 1

30 0 0 0

40 0 0 0

Ozone injury was found on morning glory (Ipomea purpurea) leaves at a

monastery complex in Kiev (table 2). Wild hop (Humulus lupulus) and native clematis

(Clematic vita alba) showed possible ozone injury at the Botanical Garden in Kiev.

At Uzoksky Pass, possible ozone injury was observed on leaves of red-fruited

elderberry (Sambucus racemosa), a common forest inhabitant. At Synevir, elderberry

also showed possible ozone injury. At Shiroky Lug, probable ozone injury was

observed on blackberry (Rubus hirtus), hazelnut (Corylus avellana), wild clematis

(Clematis hyb.), and vincetoxicum (Vincetoxicum sp.). At a forestry administration

unit near Rakhiv, probable ozone injury was noted on cultivated clematis, wild

hop, and elderberry.

Some native plants are potentially excellent detector indicators of ambient

ozone in the Carpathian Mountains of Ukraine:

● Actual Injury — Bel-W3 tobacco (Nicotiana tabacum), morning glory

(Ipomoea purpurea).

● Probable Injury — wild hop (Humulus lupulus).

● New Species — clematis (Clematis vita alba), clematis (Clematis hyb.),

hazelnut (Corylus avellana), Vincetoxicum (Vincetoxicum sp.).

● Known Species — blackberry (Rubus hirtus), elderberry

(Sambucus racemosa).

Their response to ozone needs to be verified so that they can be used as

bioindicators.

In general, the results of our pilot study indicated that ambient O3

 in Kiev and

some forest regions of Ukraine during some summer periods could injure plants.

The predicted increase of automobile traffic in Ukraine and the neighboring

countries and long-range transport of the air masses contaminated with

Session I1 Ozone Air Pollution in the Ukrainian Carpathian Blum, Bytnerowicz, Manning, Popovicheva
50 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

photochemical smog may cause further increase of O3 concentrations in the forested

areas of the Ukrainian Carpathian Mountains. Continuous monitoring of O3

concentrations and its phytotoxic effects in the Carpathian forests and other parts

of Ukraine is needed in order to signal the potential for deterioration of sensitive

plant species in this part of Europe.

Acknowledgments

This study was partially supported by the USDA International Cooperation and

Development and the Pacific Global Change Research Program, USDA Forest

Service. We thank Krystyna Grodzinska, Barbara Godzik, and Marek Krywult,

Institute of Botany, Polish Academy of Sciences, Krakow, Poland for their help in

the initial phase of this study. Many thanks go to the colleagues from the Central

Botanical Garden (Kiev): Vladimir Grakhov and Aleksey Chekhovich for computer

statistical processing of continuous ozone analyzer monitoring data and for figure

drawings. We also thank Laurie Dunn for technical editing of this manuscript.

References

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Bytnerowicz, Andrzej; Glaubig, Robert; Cerny, Martin; Michalec, Miroslav; Musselman, Robert; Zeller, Karl. 1995. Ozone

concentrations in forested areas of the Brdy and Sumava mountains, Czech Republic. Presentation in the 88th annual concentrations in forested areas of the Brdy and Sumava mountains, Czech Republic

meeting and exhibition, San Antonio, Texas. June 18-23; Air and Waste Management Association; 95-MP20.04: 1-12.

Bytnerowicz, Andrzej; Manning, William J.; Grosjean, Daniel; Chmielewski, Waldemar; Dmuchowski, Wojciech; Grodzinska,

Krystyna; Godzik, Barbara 1993. Detecting ozone and demonstrating its phytotoxicity in forested areas of Poland: a pilot

study. Environmental Pollution 80: 301-305. study

Derwent, R.G.; Jenkin, M.E. 1991. Hydrocarbons and the long-range transport of ozone and PAN across Europe. Atmospheric Hydrocarbons and the long-range transport of ozone and PAN across Europe

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Heggestad, Howard 1991. Origin of Bel-W3, Bel-C and Bel-B tobacco varieties and their use as indicators of ozone. Origin of Bel-W3, Bel-C and Bel-B tobacco varieties and their use as indicators of ozone. Environ- Origin of Bel-W3, Bel-C and Bel-B tobacco varieties and their use as indicators of ozone.

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using a nitrate-coated filter. Analytical Chemistry 65: 209- 214. using a nitrate-coated filter

Krupa, Sagar V.; Manning, William J. 1988. Atmospheric ozone: formation and effects on vegetation. Atmospheric ozone: formation and effects on vegetation. Environmental Pollu- Atmospheric ozone: formation and effects on vegetation.

tion 50: 101-137.

Laurila, T.; Latilla, H. 1993. Surface ozone exposures measured in Finland. Atmospheric Environment 27A. Surface ozone exposures measured in Finland

Logan, J. A. 1985. Tropospheric ozone: seasonal behavior, trends, and anthropogenic influence. Journal Geophysical Tropospheric ozone: seasonal behavior, trends, and anthropogenic influence

Research (Atmosphere) 90: 10,463-10,482.

National Report on Environment Conditions in Ukraine in 1993 (in Ukrainian). 1994. Ministry for environmental protection National Report on Environment Conditions in Ukraine in 1993

and nuclear safety of Ukraine. Rayevsky Publ.; 178 p.

Proyou, A. G.; Toupance G.; Perros, P. E. 1991. A two-year study of ozone behaviour at rural and forested sites in eastern

France. Atmospheric. Environment 25A: 2,145-2,153. France

Semenov, S.; Kouhta, B. 1996. Ozone influence on tree and crop growth: modelling the effects and estimating critical levels. Ozone influence on tree and crop growth: modelling the effects and estimating critical levels

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Session I1 Ozone Air Pollution in the Ukrainian Carpathian Blum, Bytnerowicz, Manning, Popovicheva
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 51

Abstract

From June to mid-October in 1995, the concentration of tropospheric ozone in 18 localities in the Krakow Province of

southern Poland was measured by using ultraviolet monitors and Ogawa passive samplers. At three active monitoring

stations, tropospheric ozone was recorded in the downtown and western part of Krakow and in Szarow, 30 km to the

east. The passive method was applied in a dozen or so localities distributed within and outside Krakow’s city limits. In

these locations two varieties of tobacco (Nicotiana tabacum L.), Bel-W3 and Bel-B, were exposed. In the Krakow

Province the mean 24-hour ozone concentration ( from active monitors) was higher than 30 μg/m3

 (24-hour Polish

standard) in June through July and part of September. The highest concentration was recorded in early afternoons and

the lowest between midnight and sunrise. The maximum 30-minute concentration of 205 μg/m3

 occurred in August.

However, great diversity of the tropospheric ozone concentration (measured with passive samplers) was recorded

in all locations. The lowest average concentration was recorded in Krakow and areas located west, northwest

and east of the city, whereas the highest average concentration was recorded in the north, northeastern,

southeastern, and southwestern parts of the Province. The highest average concentration of 100 μg/m3 for a 2-

week period of filter exposure was recorded in July and August from the sites Ratanica, located 40 km south, and

Goszcza, 25 km north of Krakow. The amount of damage to the leaves of the ozone-sensitive variety of tobacco

(Bel-W3) exposed at all sites was correlated (coefficient 0.69, p<0.000) with concentration results obtained

with passive samplers. Greater damage was recorded in the sites located outside the city than elsewhere, whereas

the damage that occurred within Krakow was markedly smaller.

Introduction

Tropospheric ozone is believed to be an important phytotoxic air pollutant (Fuhrer

and Achermann 1994, Guderian 1985, Stanners and Bourdeau 1995). Ozone levels

recorded in western Europe, such as France (Proyou and others 1991), Switzerland

(Ballaman 1993), or Austria (Dovland 1987), exceed the permissible concentrations

for that pollutant. In the countries located in northeastern Europe, such as Finland

(Laurila and Lattila 1993) or Lithuania (Girgzdiene 1991), concentrations of ozone

are relatively lower than in central Europe (Simpson 1993). Southern Poland is

among the most polluted areas of central Europe. The high input of sulphur, nitrogen

oxides, and heavy metals has originated from local and remote sources (Nowicki

1993). Recent studies performed in southern Poland (Upper Silesia Region, Beskidy

Mountains, Krakow Region) have shown that during the summer season a single

1-hour average ozone concentration may reach 120 ppb (Bytnerowicz and others

1993, Godzik and others 1994).

In the Krakow agglomeration, the main sources of pollution are a huge steel

mill, a cement factory, an electric power station, privately-owned coal stoves and

also rapidly increasing car traffic (Turzanski 1994). The highest concentration of

air pollutants in the Krakow Province occurs centrally in the city of Krakow

(Turzanski 1996).

This paper reports a study of ozone concentrations within the city of Krakow

in southern Poland and outside its limits; identifies which areas have the highest

ozone levels; and evaluates phytotoxic effects of ozone on the basis of responses

of bioindicator tobacco (Nicotiana tabacum L.) plants.

Study Area and Methods

The Krakow Province covers an area of 3,254 km2. In 1995 measurements of the

ozone concentration were made at 18 sites: 7 among them were located within the

Ozone Monitoring in the Krakow Province, Southern Poland1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Research Ecologist, W. Szafer Institute

of Botany, Polish Academy of Sciences,

46 Lubicz Str., 31-512 Krakow, Poland.

Barbara Godzik2
52 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

city limits, 11 were evenly distributed in the Krakow Province (fig. 1). Two sites

located within the city (Balice, Wieza Ratuszowa) and the one in Szarow (30 km

east of the city) were equipped with continuously working monitors (Thermo

Environmental Model 49 UV-photometric ambient O3 analyzer, Franklin, MA). 3 In

two stations within the city measurements were performed by the Voivodship

Inspectorate for Environmental Protection in Krakow. The correlation coefficient

for the conversion of ppb to μg/m3 were 1.93 for T=20 °C. All sites were furnished

with the Ogawa passive samplers (Ogawa and Company, Pompano Beach, FL)

(Koutrakis and others 1993). Passive sampler exposure was done from early June

to mid-October in 2- or 3-week intervals. The O3 concentration in the Krakow

Province was mapped by using Surfer 5.0 program. Isolines were made following

the kriging method for the parameters: component 1 = linear, scale C = 133, length

A = 40.7. Graphic description was made in CorelDraw 3.0 program.

In every monitoring site four plants of the Bel-W3 and four plants of the Bel-B

tobacco (Nicotiana tabacum L.) varieties were exposed during two periods (07/20-

08/08 and 08/11-09/06). Damage to leaves was measured at 1-week intervals

according to procedures described by Manning and Feder (1980). Four leaves on

each plant were evaluated for injury by the sum of percentage area showing

symptoms on each leaf. The correlation between percentage of leaf damage and

ozone concentration was analyzed by Spearman rank test.

Results

Thirty-minute average concentrations of ozone measured with continuous monitors

varied predictably during a 24-hour period (especially in July and August) and

between months in all sites. The ozone concentration at night decreased to several

μg/m3, and it increased to 160 μg/m3 during the day (fig. 2). According to the Polish

standard, the 30-minute concentration should not be greater than 100 μg/m3. About

100 hours in June and July and more than 200 hours in August showed

concentrations higher than the standard. In September excessive ozone

concentrations occurred less frequently (fig. 3).

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik

Figure 1 — Distribution of

ozone-monitoring sites in the

Krakow city (A) and the Krakow

Province (B). 1—stations

equipped with monitor and

passive samplers, 2—stations

equipped with passive samplers.

11

10

9 8

18

16

17

15

14

13 12

1

2

7 3 6

4

5

1 2

KRAKOW

Krowodrza

Pcim

Polanka

Haller

Ratanica

Wola

Filipowska Radwanowice

Wista Szarów

Wawrzen'czyce

Goszcza Lyszkowice

Ojeów

Zadroze

54°

52°

50°

16° 20° 24°

KRAKOW

Nowa

Huta

Padogorze

A

B
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 53

The maximum 30-minute ozone concentration for subsequent months occurred

in August (table 1). The highest ozone concentration occurred in August. The lowest

concentration was recorded between midnight and sunrise and the highest in early

afternoons (fig. 4).

In July and August the maximum mean 24-hour concentrations at the Balice

research station were higher than in June. These values were markedly lower in

September than in the previous months (table 2). The mean 24-hour concentrations

of ozone at the Wieza Ratuszowa station were similar in subsequent months to the

concentrations recorded in the Balice station. In general, however, the minimum

values for subsequent months were lower and the maximum values were higher

than the ones recorded from the Balice station (table 2). The mean 24-hour

permissible ozone concentration, which follows the Polish standard, should not

be higher than 30 μg/m3

. In June, July, August, and part of September the mean

24-hour concentration in Krakow was markedly higher than the Polish standard.

During the period of measurements from June through September, the mean

monthly concentrations measured by active monitor were higher at the Balice

station (western part of the city) than at the Wieza Ratuszowa station (center of

the city). The highest mean monthly value for August was recorded from the Szarow

station (30 km east of Krakow) (table 2).

Table 1 — Table 1 — Highest 30-minute ozone concentration (μg/m3 ) at three sites in southern Poland, by month of

measurement.

June July August September

Balice 152 177 1159 120

Wieza Ratuszowa 148 189 205 126

Szarow - - 180 129

Figure 2 — Diurnal pattern of

ozone concentration (μg/m3

) in

July 1995 at the Balice monitoring

station (western part of the city).

(1μg/m3 O3

= 0.51 ppb O3 at 25oC,

760 mm of Hg.)

Figure 3 — Diurnal pattern of

ozone concentration (μg/m3

) in

September 1995 at the Balice

monitoring station (western part

of the city).

1 Data incomplete for a whole period

g/m3

Days (July)

Polish standard

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

200

150

100

50

0

g/m3

Days (September)

Polish standard

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30

150

100

50

0

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
54 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

In all periods of measurements made with passive samplers, the average

ozone concentration was markedly different among selected sites and exposure

periods (table 3). In the first two 2-week periods of measurement (including

June) and in the last two (September and October) the mean ozone concentration

was low (table 3). In July and August it increased dramatically. The maximum

values for a 3-week exposure (June, July) were recorded from Goszcza and

Ratanica research site (about 100 μg/m3

(table 3). During the same period, a

high average ozone concentration occurred also in Szarow, Polanka Haller, and

Trzyciaz. In all measurement periods the lowest average concentration of ozone

was recorded from the sites located in the city center (Krasinski Avenue,

Helclow Street) (table 3). All sequences of measurements yielded a similar

pattern of spatial differentiation of the ozone concentration in the Krakow

Province. During all exposure periods the lowest average ozone concentration

occurred in the center of the Krakow Province (Krakow city and the areas located

west, northwest, and east of the city) and in its southern part (figs. 5, 6, 7). The

highest concentration was determined in the north, northeastern, southeastern,

and southwestern parts of the Krakow Province (figs. 5, 6, 7).

The amount of damage to the leaves of the ozone-sensitive variety of tobacco

(Bel-W3) exposed at all sites was correlated with results obtained through

passive samplers. This coefficient (=0.69, p<0.000) was statistically significant.

During two exposition periods of tobacco (Nicotiana tabacum L.) the lowest injury

to the leaves of the ozone-sensitive (Bel-W3) variety of tobacco was recorded

at the sites located in the center of Krakow (Krasinski Avenue, Helclow Street)

Figure 4 — Monthly averages of

diurnal dynamics of ozone

concentrations (μg/m3

) at the

Balice monitoring station.

Table 2 — Table 2 — Average daily minimum (min) and maximum (max) and average monthly (x) ozone concentrations Table 2 — 

(μg/m3

) at the Balice, Wieza Ratuszowa, and Szarow Research Station.

 Month Balice Wieza Ratuszowa Szarow

min 30.21 25.11 -

June max 69.71 78.43 -

 x 57.95 55.06 -

min 46.66 45.57 -

July max 83.02 87.72 -

x 66.46 65.28 -

min 30.13 21.49 28.16

August max 92.08 98.41 113.96

x 58.98 53.86 68.68

min 10.51 5.25 10.28

September max 58.96 52.65 61.48

 x 34.06 27.64 30.41

c

August

July

September

June

g/m3

Hours

120

100

80

60

40

20

0

0 3 6 9 12 15 18 21

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 55

and at the sites located west of the city (Radwanowice, Ojcow, Wola Filipowska)

(fig. 8). The most severe injury to the tobacco leaves were noted in sites located in the

northern, southeastern, and southwestern parts of the Krakow Province (Trzyciaz,

Goszcza, Szarow, Ratanica, Polanka Haller) (fig. 8). No leaf injury was observed in the

ozone-tolerant tobacco variety (Bel-B) at any of the study sites.

Discussion

Ozone has been recognized as a major phytotoxic constituent of urban and

suburban air pollution (Stanners and Bourdeau 1995). Concentrations of ozone

at average and high latitudes of the northern hemisphere doubled during the

last century (Anfossi and others 1991, Volz and Kley 1988). Over the last decade

an increase in ozone concentration has also been observed in the countries of

north-eastern Europe. For example, rural Lithuania has been affected by the

long-range transport from central Europe (Girgzdiene 1995).

Ozone causes the reduction of yield in crop species in North America and

Europe (Fuhrer and Ascherman 1994), and these effects become more

apparent as the phytotoxic concentrations of ozone increase in the rural

areas (Colbeck and Mackenzie 1994). Ozone is also one of the factors causing

recent forest decline on these continents (Ashmore and others 1985,

Chevone and Linzon 1988).

Studies made in the Krakow Province found that the concentration of ozone

in this area is markedly higher from the levels generally accepted as natural

(Guderian 1985). During summer the concentrations of ozone in the area

surrounding Krakow are higher than in unpolluted areas of central Europe

(Girgzdiene 1991, Hakoala and others 1991), and they are similar to the ozone

concentrations recorded from western European countries (Ballaman 1993, Dovland

Table 3 — Table 3 — Average ozone concentrations ( Table 3 — μg/m3

) during different exposure periods in the Krakow city and the

Krakow Province.

Station Exposure periods

5/31 6/14 6/27 7/17 8/7 8/30 9/21

-6/13 -6/27 -7/17 -8/7 -8/30 -9/21 -10/17

Krakow city: Krakow city:

1. Balice 54.5 43.3 72.9 78.5 55.8 32.1 32.2

2. Krasinskiego - - 19.3 14.7 17.1 12.7 10.8

3. Helclow - 4.9 25.7 25.1 23.5 12.7 7.6

4. Strzelcow - - 50.2 57.9 45.0 28.5 18.5

5. Wallenroda - - 71.5 65.0 46.3 35.3 22.3

6. Zeromskiego - 43.6 71.1 66.9 48.4 26.6 19.3

7. Zielone - - 57.2 49.3 38.7 23.2 17.4

Krakow Province: Krakow Province:

8. Wola Filipowska 44.0 30.3 55.1 49.8 39.1 24.9 19.5

9. Radwanowice 47.7 31.4 49.8 55.8 43.8 24.8 21.7

10. Ojcow 29.8 12.4 32.9 37.9 29.1 19.1 16.8

11. Zadroze 60.7 57.3 86.2 88.0 67.1 45.4 39.3

12. Goszcza 65.0 57.2 100.1 101.5 61.4 48.1 35.3

13. Lyszkowice 54.3 48.1 77.9 81.5 60.8 42.3 29.4

14. Wawrzenczyce 40.3 17.9 50.5 50.0 39.3 31.8 19.8

15. Szarow 58.8 51.1 81.5 89.1 70.5 39.1 45.9

16. Ratanica 69.5 52.6 98.1 107.6 73.3 56.2 32.1

17. Pcim 35.4 50.8 50.0 47.4 35.8 26.5 20.3

18. Polanka Haller 54.3 48.2 86.6 85.9 63.5 44.2 41.4

c

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
56 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

1987, Proyou and others 1991). The highest monthly ozone concentration in

southern Poland occurred in July and in the beginning of August. In Upper Silesia

(80 km west of Krakow) the ozone concentration recorded in 1994 was similar to

the values recorded in Krakow and were higher than the values recorded from

these areas in 1995 (Godzik and others 1994, Godzik and Grodzinska 1996).

Although the highest concentration of ozone occurred in the city of Katowice, a

high concentration was not recorded at that time in other sites in Silesia. This was

probably caused by the presence of appropriately high concentrations of ozone

precursors in the air (Godzik and others 1995). Ozone concentrations within the

Krakow Province differed: higher concentrations were recorded in the northern

and southern sites that are at higher elevations. In these localities, in contrast to

the urban areas, small changes in the ozone concentration over 24 hours have

been observed (absence of zero ozone concentration in the night) (Godzik and

others 1995). It is impossible to give definite reasons for the occurrence of the highest

ozone concentration in the northern and southern parts of the Krakow Province

because data are not available on the proportion of nitrogen oxides and

hydrocarbon or sun exposure regimes for the study sites.

Figure 5 — Spatial ozone

distribution (μg/m3

) obtained

from passive samplers in the

Krakow Province for the period

27 June to 17 July 1995.

Figure 6 — Spatial ozone

distribution (μg/m3

) obtained

from passive samplers in the

Krakow Province for the period

17 July to 7 August 1995.

Figure 7 — Spatial ozone

distribution (μg/m3

) obtained

from passive samplers in the

Krakow Province for the period

21 September to 16 October

1995.

5

6 7

Zadroze

Ojców

Radwanowice

Polanka

Haller

Ratanica

< 58

58 - 65

> 65 Pcim

Szarów

Strzelców

Helclów

Krasinskiego

Wallenroda

Balice Zielone

65

65

65

65

65

65

58

58

58

58

58

Zeromskiego

Wola

Filipowska Wawrzenczyce

Gaszca Lyszkowice

Zadroze

Ojców

Radwanowice

Polanka

Haller

Ratanica

< 58

58 - 65

> 65 Pcim

Szarów

Strzelców

Helclów

Krasinskiego

Wallenroda

Balice Zielone

65

65

65

65

65

65

65

58

58

58

58

58

Zeromskiego

Wola

Filipowska Wawrzenczyce

Gaszca Lyszkowice

Zadroze

Ojców

Radwanowice

Polanka

Haller

Ratanica

< 23

23 - 27

> 27 Pcim

Szarów

Strzelców

Helclów

Krasinskiego

Wallenroda

Balice Zielone

27

27

27

23

27

27

23

27

23

29

23

27

23 Zeromskiego

Wola

Filipowska Wawrzenczyce

Gaszca Lyszkowice

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 57

The ozone concentrations in the Krakow Province were high enough to cause

leaf injury of the ozone-sensitive variety of tobacco (Bytnerowicz and others 1993,

Spellerberg 1991). Similar concentrations in such amounts may also cause injuries

in other plant species (Chappelka and Chevone 1992, Runeckles and Chevone 1992).

In the Krakow Province the levels of other pollutants are high, including sulfur

dioxide, particulate pollution heavily contaminated with heavy metals, etc. These

pollutants originate either from the local sources or from the transboundary

transport (Nowicki 1993). Synergistic effects of these compounds and ozone may

result higher damage of plants than by single pollutants (Guderian 1985). The

results of ozone measurement concentration in southern Poland (Upper Silesia,

Krakow region) and evaluation injury of the test-plants have shown that the cooccurence of elevated ozone concentration and high physiological potential of

native plants can cause their injury. Ozone injury to native plants in Beskid Slaski

of southern Poland has been observed but has not been verified in controlled ozone

exposures (Godzik and others 1995). Similarly, the occurrence of ozone injury

described in the literature has been recorded in the area surrounding Krakow.

Among the affected species are: Ulmus scabra, Ulmus laevis, Sambucus racemosa,

Rubus hirtus.

4

 However, the above observations need to be confirmed in the

controlled ozone exposures.

The number of motor vehicles has rapidly increased since 1990 in the Krakow

region (Turzanski 1994) and has resulted in increased emission of nitrogen oxides

and hydrocarbon. These compounds, known as ozone precursors, can cause further

increase in ozone concentrations in the future.

Figure 8 — Leaf injury to

bioindicator plants (Bel-W3

cultivar of tobacco) exposed in

Krakow (sites 1-7) and outside

the city (sites 8-18) in the period

07/20-08/08/1995. The amount of

injury is presented as the mean

sum for four evaluated leaves

from each of the four plants; data

accumulated for subsequent

exposition weeks. Research sites

have been labelled as in table 3.

c

4 Unpublished data on file at the Polish

Academy of Sciences.

pct

Exposure: 7/26 - 7/31/95

Sites

0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

0

50

100

150

pct

Exposure: 7/31 - 8/8/95

Sites

0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

0

50

100

150

pct

Exposure: 7/20 - 7/26/95

Sites

0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

0

50

100

150

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
58 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Acknowledgments

This study was jointly funded by the USDA Office of International Cooperation

and Development and the USDA Forest Service program “Effects of atmospheric

deposition and climate change of forest ecosystem in Eastern Europe and the United

States,” by the Voivodship Environmental Protection and Water Management Fund

in Krakow, and by the Wladyslaw Szafer Institute of Botany, Polish Academy of

Sciences. I thank Elzbieta Chrzanowska and Stanislaw Braniewski for their technical

assistance. I thank Laurie Dunn for technical editing of this manuscript.

References

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Geophysical Research 96: 17,349-17,352.

Ashmore, M; Bell, M; Rutter, J. 1985. The role of ozone in forest damage in West Germany. The role of ozone in forest damage in West Germany. Ambio 14: 81-87. The role of ozone in forest damage in West Germany.

Ballaman, R. 1993. Transport of ozone in Switzerland. Transport of ozone in Switzerland. Science of the Total Environment 134: 103-115. Transport of ozone in Switzerland.

Bytnerowicz, A.; Manning, W.J.; Grosjean, D.; Chmielewski, W.; Dmuchowski, W.; Grodzinska, K.; Godzik, B. 1993.

Detecting ozone and demonstrating its phytotoxicity in forested areas of Poland: a pilot study. Detecting ozone and demonstrating its phytotoxicity in forested areas of Poland: a pilot study. Environmental Detecting ozone and demonstrating its phytotoxicity in forested areas of Poland: a pilot study.

Pollution 80: 301-305.

Chappelka, A.H.; Chevone, B.I. 1992. Tree responses to ozone. Tree responses to ozone. In: Lefohn, A.S., ed. Surface level ozone exposures and their Tree responses to ozone.

effects on vegetation. Chelsea, MI: Lewis Publishers; 271-324.

Chevone, B.I.; Linzon, S.N. 1988. Tree decline in North America. Tree decline in North America. Environmental Pollution 50: 87-99. Tree decline in North America. 

Colbeck, I; Mackenzie, A.R. 1994. Air pollution by photochemical oxidants. Air pollution by photochemical oxidants. Air Quality Monographs, Vol. 1. Amsterdam- Air pollution by photochemical oxidants.

London-New York-Tokyo: Elsevier; 376 p.

Dovland, H. 1987. Monitoring European transboundary air pollution. Monitoring European transboundary air pollution. Environment 29: 10- 27. Monitoring European transboundary air pollution.

Fuhrer, J.; Ascherman, B., eds. 1994. Critical levels for ozone - a UN-ECE workshop report. Critical levels for ozone - a UN-ECE workshop report. Liebefeld. Schriftenreihe Critical levels for ozone - a UN-ECE workshop report.

FAC 16; 328 p.

Girgzdiene, R. 1991. Surface ozone measurements in Lithuania. Surface ozone measurements in Lithuania. Atmospheric Environment 25A: 1,791-1,794. Surface ozone measurements in Lithuania. 

Girgzdiene, R. 1995. Surface ozone concentrations and exposures during growing season at the Lithuanian rural site. Surface ozone concentrations and exposures during growing season at the Lithuanian rural site. Water, Surface ozone concentrations and exposures during growing season at the Lithuanian rural site.

Air, and Soil Pollution 85: 1,509-1,514.

Godzik, Barbara; Grodzinska, Krystyna. 1998. Ozone concentrations in Krakow region, Poland—a three-year study. Ozone concentrations in Krakow region, Poland—a three-year study. In: Ozone concentrations in Krakow region, Poland—a three-year study.

Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan, technical coordinators. Proceedings of the international

symposium on air pollution and climate change effects on forest ecosystems. 1996 February 5-9; Riverside, CA. Gen.

Tech. Rep. PSW-GTR-166. Albany, Ca: Pacific Southwest Research Station, USDA Forest Service [this volume].

Godzik, Stefan; Hlawiczka, Stanislaw; Poborski, Piotr. 1995. Smog-Przyczyny-Skutki- Przeciwdzialania. Biblioteka Monitoringu Smog-Przyczyny-Skutki- Przeciwdzialania

Srodowiska. PIOS. Warszawa; 58 p.

Godzik, Stefan; Szdzuj, Jerzy; Osrodka, Leszek; Stawiany, Wojciech. 1994. Time and spatial differentiation of ground level Time and spatial differentiation of ground level

ozone concentration from four monitoring stations in the Upper Silesia Region. ozone concentration from four monitoring stations in the Upper Silesia Region. Seminar: Ozone—a regional and global ozone concentration from four monitoring stations in the Upper Silesia Region.

problem. Katowice, 21 November 1994. Polish National Committee, Global IGBP Change. Institute for Ecology of Industrial Areas, Katowice: 41-50.

Guderian, R., ed. 1985. Air pollution by photochemical oxidants, formation, transport, control and effects on plants. Air pollution by photochemical oxidants, formation, transport, control and effects on plants. Berlin: Air pollution by photochemical oxidants, formation, transport, control and effects on plants.

Springer Verlag; 247-250.

Hakola, H.; Joffre, S.; Lattila, H.; Taalas, P. 1991. Transport, formation and sink processes behind surface ozone variability in

north European conditions. north European conditions. Atmospheric Environment 25A: 1,437-1,447. north European conditions. 

Koutrakis, Petros; Wolfson, Jack M.; Bunyaviroch, Arnold; Froelich, Susan E.; Hirano, Koichiro; Mulik, James D. 1993. Measurement of ambient ozone using a nitrate-coated filter. surement of ambient ozone using a nitrate-coated filter. Analytical Chemistry 65: 209-214. surement of ambient ozone using a nitrate-coated filter. 

Laurila, T.; Lattila, H. 1993. Surface ozone exposures measured in Finland. Surface ozone exposures measured in Finland. Atmospheric Environment 27A. Surface ozone exposures measured in Finland. 

Manning, W.J.; Feder, W.A. 1980. Biomonitoring air pollutants with plants. Biomonitoring air pollutants with plants. London: Applied Science Publishers; 142 p. Biomonitoring air pollutants with plants.

Nowicki, Maciej. 1993. Environment in Poland. Issues and solutions. Environment in Poland. Issues and solutions. Dordrecht/Boston/London: Kluvier Academic Environment in Poland. Issues and solutions.

Publishers; 191 p.

Proyou, A.G.; Toupance, G.; Perros, P.E. 1991. A two-year study of ozone behaviours at rural and forested sites in eastern

France. France. Atmospheric Environment 25A: 2,145-2,153. France. 

Runeckles, V.C.; Chevone, B.I. 1992. Crop response to ozone. Crop response to ozone. In: Lefohn, A.S., ed. Surface level ozone exposures and their Crop response to ozone.

effects on vegetation. Chelsea, MI: Lewis Publishers; 189-271.

Simpson, D. 1993. Photochemical model calculations over Europe for two extended summer periods: 1985 and 1989. Model Photochemical model calculations over Europe for two extended summer periods: 1985 and 1989. Model

results and comparision with observations. results and comparision with observations. Atmospheric Environment 27A: 921-943. results and comparision with observations. 

Spellerberg, I.F. 1991. Monitoring ecological change. Monitoring ecological change. Cambridge, U.K: Cambridge University Press. Monitoring ecological change.

Stanners, Dawid; Bourdeau, Philippe, eds. 1995. Europe’s environment. The Dobris assessment. Europe’s environment. The Dobris assessment. Copenhagen: European En- Europe’s environment. The Dobris assessment.

vironment Agency; 547-551.

Turzanski, Konrad P., ed. 1994. Problemy zanieczyszczen komunikacyjnych. Problemy zanieczyszczen komunikacyjnych. Biblioteka Monitoringu Srodowiska. Krakow. Problemy zanieczyszczen komunikacyjnych.

PIOS, WIOS; 82 p.

Turzanski, Konrad P., ed. 1996. Raport o stanie srodowiska w wojewodztwie krakowskim w 1994 roku. Raport o stanie srodowiska w wojewodztwie krakowskim w 1994 roku. Biblioteka Monitoringu Raport o stanie srodowiska w wojewodztwie krakowskim w 1994 roku.

Srodowiska. Krakow. PIOS, WIOS; 60 p.

Volz, A.; Kley, D. 1988. Evaluation of the Montsouris series of ozone measurements made at the nineteenth century. Evaluation of the Montsouris series of ozone measurements made at the nineteenth century. Nature Evaluation of the Montsouris series of ozone measurements made at the nineteenth century.

332: 240-242.

Session I1 Ozone Monitoring in the Krakow Province, Southern Poland Godzik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 59

Stefan Godzik2

Abstract

In the Beskidy Mountains at Brenna, Poland, and several other locations in the Katowice administrative district,

plants were used as bioindicators to determine ozone concentration measurements in 1994 and 1995. Results showed

that ozone is the only gaseous air pollutant significantly exceeding the permissible concentrations and causing foliar

injury to both test and some native plant species. For all of the tests using plants, the most severe ozone injuries, were

found for the Brenna location. The first signs of leaf injury were found on beans (Phaseolus vulgaris L.). Tobacco

(Nicotiana tabacum L.) Bel W3 also showed leaf injury at this time. Less sensitive plants included the tomato

(Lycopersicum esculentum) and soy bean (Glycina max). More severe injuries to the same plant species (cultivars)

were not found with increased elevation. In the Brenna region only, foliar injury was also found on field-grown plant

species: Achillea sp., Asceplias syriaca L., Impatiens parviflora DC., Sambucus nigra L., S. racemosa L., Fagus

silvatica L., Prunus avium L., and Rubus sp. The mean ozone concentrations at Brenna were 85.2 and 77.4 μg/m3

in the summers of 1994 and 1995, respectively. For the locations in the Katowice area, mean concentrations were 50.3

and 44.5 μg/m3

. AOT40 calculated on a 24- hour or day-hours (9.00 - 17.00) basis exceeded the critical values of

5,300 ppb.h and 1 ppm.h recommended by the United Nations ECE for crops and forest tree species. Thus, on

the basis of these results ozone is a contributing factor to poor forest health conditions in the Upper Silesia

Region of Poland.

Introduction

Leaf injury to native or introduced plant species, such as horse-chestnut (Aesculus

hippocastanum L.), rose (Rosa sp.), and some others, was caused by air pollutants of

photochemical origin in the vicinity of coke ovens in the mid-1960’s (Godzik and

Piskornik 1966). However, injury of tobacco (Nicotine tabacum L.) Bel W3 was not

the same as that caused by ozone. Nonetheless, air pollutants of photochemical

origin can cause leaf injury in locations where sulfur dioxide, nitrogen oxides, and

particulate matter were seen as the only “real” problem to the environment.

Exposure started later for the Bel W3 and Bel B tobacco, and for radish

(Raphanus sativa cv Cherry Bella) with and without ethylene diurea (EDU)

treatments in the mountain region (Godzik and others 1991). Injuries to Bel W3

and differences in the response of radish with no EDU treatments have shown

that ozone has caused adverse effects, including injury to test plants. Injury to

some native or introduced field grown plant species has been observed

simultaneously.

A 6-month series of measurements using passive samplers was performed in

1990 for the Rozdroze Izerskie (Sudety Mountains) and Brenna area (Beskidy

Mountains). These measurements were completed by 1-week continuous

measurements of ozone in these two locations.3

 Data have shown that

concentrations are similar to those found in other European countries.

First, short-term ozone concentration measurements using ozone analyzers

were done in Brenna in 1990.3 The highest 1-hour mean concentration was 110

μg/m3. Next, more regular measurements of ozone concentration and experiments

with several test plants were done. In 1994 an extensive air pollution monitoring

program started in the Katowice area by using test plants as bioindicators, including

tobacco (Nicotiana tabacum L.), Bel W3 and Bel B, bean (Phaseolus vulgaris L. cv Lit),

clover (Trifolium subtereanum cv crimsoni), tomato (Lycopersicum esculentum L. cv

Tiny Tim), and soy bean (Glycina max). These were grown in containers with

standardized soil and in the local soil (Godzik and others 1995, 1996 a, b).

Ozone in the Upper Silesia Region — Concentration and

Effects on Plants1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Natural Sciences, Institute

for Ecology of Industrial Areas,

Kossutha 6, 40-833 Katowice, Poland

3 Unpublished data supplied by the

author.
60 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Table 1 Table 1 – Concentration of sulfur dioxide, nitrogen dioxide, and ozone at some monitoring stations in the Upper Table 1 

Silesia Region, Poland. 1

Pollutant Pollutant Katowics Kuznia Nieb. Kuznia Nieb. Slawkow Brenna Standard

 and year value

D30

D24

Da

Nitrogen dioxide

500

1994 50.1 22.7 12.1 3.1 150

1995 23.5 17.0 16.8 5.3 50

 Sulfur dioxide

600

1994 36.7 14.1 27.8 6.5 200

1995 27.6 10.4 21.5 24.0 32

 Ozone

100

1994 45.9 50.3 45.8 85.2 30

1995 47.3 44.5 - 77.4 -

The objective of this study was to assess the effect of ambient ozone

concentration in the Upper Silesia Region of Poland by using plants as bioindicators

exposed at locations that differed in climate and air pollution.

Materials and Methods

In the growing seasons of 1994 and 1995 ozone concentrations were measured by

using analyzers, including Environics series 300, MLU and Thermo Environmental.4

Before each measuring season analyzers were calibrated. Seeds of all but one test

plant (tobacco) were provided by the International Cooperative Program (ICP) Crops

Coordination Center of Nottingham University, England. Seeds of Bel W3 and

Bel B tobacco were provided by the University of Massachusetts.

Clover and bean in standardized soil and containers were exposed in Brenna,

Katowice, Kuznia Nieborowicka, and Slawkow. Before 1994 the number of test

plants was limited to clover and tobacco only. Radish, treated and non-treated

with EDU, was used during two growing seasons.

Bean, clover, and Bel W3 and Bel B tobacco were grown in standardized soil,

in self-watering containers. Procedures were based on guidelines elaborated by

the ICP Crops, under the UN ECE Convention on Long Range Transboundry Air

Pollution. At Brenna and Katowice, beans, tomato (Lycopersicum esculentum L.),

and soy bean (Glycina max) were also grown in local soils. Foliar injury of both test

and native plant species were compared with those given in the literature and

photographs5 (Manning and Feder 1980, Sanders and Benton 1995).

Results and Discussion

Mean concentration of ozone for Brenna in 1994 was 85.2 μg/m3, and for 1995 was

77.4 μg/m3(table 1). Mean concentrations for the same period for Katowice region

were 50.3 μg/m3 and 44.5 μg/m3, respectively. The highest ozone concentration

was measured in Katowice, June 1994 — 232 μg/m3. Other locations did not show

high concentrations (Godzik and others 1995, 1996) compared to the data for the

years 1994 and 1995. According to the national standards, this finding suggests

Session I1 Ozone in the Upper Silesia Region – Concentration and Effects on Plants Godzik

1 Mean values for the period: May 5 – Oct. 31, 1994 and May 5 – Aug. 31, 1995 in comparision to national standards (μg/m3

).

Calculations are based on 24-h mean values (Godzik and others 1996b).

4 Mention of trade names or products is

for information only and does not

imply endorsement by the U.S. Department of Agriculture.

5 Unpublished data supplied by the author.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 61

that ozone concentrations found in the Brenna region and the Katowice area are

high enough to cause foliar injury to plants (Council Directive 1992).

In the Brenna and Katowice locations, ozone concentrations set by the UN

ECE convention as critical levels were exceeded (Ashmore 1994, Benton and others

1995, Fuhrer and Achermann 1994 ). These levels were defined as accumulated

dose over the threshold of 40 ppb (AOT40), which is the sum of hours during the

day of ozone concentration above 40 ppb (Benton and others 1995, Fuhrer and

Achermann 1994, and Legge and others 1995).

The patterns of daily ozone concentration differ at locations in the Katowice

area and Brenna. Peak day values are higher for the Katowice area, but mean

concentrations are higher at the Brenna site. This difference is caused by higher

ozone concentrations at night at the Brenna site compared to the Katowice area

measurements. On most occasions the concentration at Brenna during the night

are in the range of 15 μg/m3 (figs. 1-2, table 1). Both for 1994 and 1995 the highest

AOT values are for Brenna, not for Katowice, where the highest 1-hour mean values

were measured (figs. 3,4).

Foliar injury of the test plants—clover, tobacco, bean, soy bean, tomato, and

native plant species—corresponded to that described in the literature (Arndt and

others 1987, Manning and Feder 1980, Sanders and Benton, 1995). The first signs

of injury were found at the Brenna location and were noted for Bel W3 and bean in

both local soil and standardized soil. They appeared after about 2 weeks of

Figure 1 — Daily pattern of

ozone concentrations at the

Brenna, Poland monitoring station,

summer 1994. (1μg/m3 O3

= 0.51

ppb O3

 at 25oC, 760 mm of Hg.)

Figure 2 — Daily pattern of

ozone concentrations at the

monitoring station of the Institute

for Ecology of Industrial Areas,

Katowice, 1994.

200.00

180.00

160.00

140.00

120.00

100.00

80.00

60.00

40.00

20.00

0.00

- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94

 - 94

- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94

-94

- 94- 94- 94- 94

May June July

03 -05 -07 -09 -

11 -

13 -15 -17 -19 -21 -23 -25 -27 -29 -31 -02 -04 -06 -08 -10 -12 -14 -16 -18 -20 -22 -24 -26 -28 -30 -02 -04 -06 -08 -10 -12 -14 -16 -18 -20 -22 -24 -26 -28 -

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

mean min. max

_

_

_

_

_

_

_

_

_

_

_

[ug/m ] 3

250

200

150

100

50

0

mean min. max

- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94- 94

30 -05 -10 -15 -20 -25 -30 -04 -09 -14 -19 -24 -29 -04 -09 -14 -19 -24 -29 -03 -08 -13 -18 -23 -28 -02 -07 -12 -17 -22 -27 -02 -07 -12 -17 -22 -27 -01 -06 -

May June July August September October 

–––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

–

–

–

–

–

–

[ug/m ] 3

Session I1 Ozone in the Upper Silesia Region – Concentration and Effects on Plants Godzik
62 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

exposure. Intensity of injuries increased during the growing season. In the Brenna

location all test plants—species or cultivars—showed foliar injury. Statistically

significant differences were not found in dry mass production of test plants treated

and non-treated with EDU and grown in standardized soil at different locations.

The number and weight of pods of bean cultivar Lit grown in local soil in both Brenna

and Katowice was lower than cultivars of Stella and Grophy. Crops of pods of beans

cultivars Stella and Grophy were very similar (Godzik and others 1996b).

Differences were found in plant responses in 1994 and 1995 between elevations

in the Brenna region and the Katowice area. At Salmopol less severe injuries were

observed in 1995. The opposite was found for the Katowice area. The reason for

these differences was undetermined. These 2 years differed in climatic conditions,

with air temperatures higher for 1994; but they also differed in concentrations of

other air pollutants (Godzik and others 1996a, 1996b; table 1). Air temperatures

close to and above 30 °C at the Brenna location were measured in June and July in

1994 for several days. A similar phenomenon was not found in the same period of

1995. Temperatures in the Katowice area for the same days were higher by a few

degrees Celsius (Godzik and others 1996a, 1996b).

Results of other investigations performed in the Brenna region in forests suggest

that climatic parameters are important factors to include when determining severity

Figure 3 — Accumulated dose

over the threshold of 40 ppb

(AOT40) for locations in Brenna

and Katowice district. Values for

the daily hours (9:00 – 17:00)

were used for calculations.

Figure 4 — Accumulated dose

over the threshold of 40 ppb

(AOT40) for Brenna and

Katowice growing season, 1995.

Twenty-four hour values were

used for calculations.

16000

14000

12000

10000

8000

6000

4000

2000

0

[ Date ]

1.05.945.05.949.05.94

13.05.9417.05.9421.05.9425.05.9429.05.94

2.06.946.06.94

10.06.9414.06.9418.06.9422.06.9426.06.9430.06.94

4.07.948.07.94

12.07.9416.07.9420.07.9424.07.9428.07.94

1.08.945.08.949.08.94

13.08.9417.08.9421.08.9425.08.9429.08.94

–

–

–

–

–

_

_

–

–

Kuznia Slawkow Katowice Brenna critical level

[ ppb h ]

–––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

18000

16000

14000

12000

10000

8000

6000

4000

2000

0

–

–

–

–

–

_

_

–

_

_––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––

[ ppb h ]

29.04.95

4.05.959.05.95

14.05.9519.05.9524.05.9529.05.95

3.06.958.06.96

13.06.9518.06.9523.06.9528.06.95

3.07.958.07.95

13.07.9518.07.9523.07.9528.07.95

2.08.957.08.95

12.08.9517.08.9522.08.9527.08.95

1.09.956.09.95

11.09.95

16.09.9521.09.9526.09.95

[ Date ]

Kuznia Katowice Brenna critical level

Session I1 Ozone in the Upper Silesia Region – Concentration and Effects on Plants Godzik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 63

of injury (Godzik and others, this volume). Ozone-like foliar injuries have been

frequently found on field-grown plant species: Achillea sp., Asceplias syriaca L.,

Impatiens parviflora DC., Sambucus nigra L., S. racemosa L., Fagus silvatica L., Prunus

avium L., and Rubus sp. Except Asceplais syriaca grown in the Katowice area, all

other species mentioned have shown signs of injury in the Brenna region only.

These findings have not been confirmed by controlled O3 fumigations.

Foliar injury on all test plants appeared in late June and increased during the

growing season. Foliar injury on Impatiens parviflora, Achillea sp., Asclepias syriaca

and Rubus sp. were noted from mid-July. On other tree or shrub foliage, signs of

injury were found in August.

Conclusion

Severe ozone injury has been observed for tobacco Bel W3 and beans grown in the

Brenna and Salmopol locations. Injuries of other plant species such as soy bean and

tomato were less severe. Ozone injury to the same plant species grown in the Upper

Silesia Industrial Region (USIR) were observed less frequently and differed between

1994 and 1995.

Ozone concentration during the day was the highest in the USIR, but the mean

concentration was the highest in the Brenna location. The accumulated dose over

the threshold of 40 ppb (AOT40) was also much higher at the Brenna. Ozone of

photochemical origin is likely the only air pollutant in southern Poland causing

foliar injury to both test and some field grown native or introduced plants, including

some broad leaf tree species.

The effects of climatic parameters and other air pollutants, such as sulfur

dioxide and nitrogen dioxide, to plant injury are not yet known. The role of air

pollutants on crop of test plants used in our experiments remains uncertain. Data

from experiments performed were not consistent with ozone concentration. For a

few locations in the Katowice area, dry mass production was the highest where

N-NH4 in bulk deposition was found to be the highest. However, more data are

needed to confirm these results.

Acknowledgments

Investigations on ozone effects on plants performed at the Beskidy site—Brenna

and Salmopol—were funded by the National Science Committee, Poland, by grant

4 S401 014 05, and the USDA Foreign Agriculture Service, International Cooperation

and Development grant “Effects of Atmospheric Deposition and Climate Change

on Forest Ecosystems in Poland and the United States.”

References

Arndt, U.; Nobel, W.; Schweizer, B. 1987. Bioindikatoren. Moeglichkeiten, Grenzen und neue Erkentnisse. Bioindikatoren. Moeglichkeiten, Grenzen und neue Erkentnisse. Verlag Eugen Ulmer. Bioindikatoren. Moeglichkeiten, Grenzen und neue Erkentnisse.

Stuttgart.

Ashmore, M.R. 1994. Critical levels for ozone. Critical levels for ozone. In: Ashmore, M.R.; Wilson, R.B., ed. 1994. Critical levels of air pollutants for Critical levels for ozone.

Europe. London: Department of Environment; 20-47. 

Benton, J.; Fuhrer, J.; Gimeno, B.S.; Skarby, L.; Balls, G.; Palmer-Brown, P.; Roadknight, C.; Sanders, G. 1995. ICP-crops and

critical levels of ozone for injury development. critical levels of ozone for injury development. In: Exceedances of critical loads and levels. Vienna: 1995 November 22-24; critical levels of ozone for injury development.

Austrian Research Centre Seibersdorf; 1-15.

Council Directive. 1992 92/72/EEC of 21 September 1992 on air pollution by ozone. EEC of 21 September 1992 on air pollution by ozone. Official Journal of the European Com- EEC of 21 September 1992 on air pollution by ozone.

munities, No L 297/1-7, 13.10.92.

Fuhrer, J.; Achermann, B. eds. 1994. Critical levels for ozone: Critical levels for ozone: a UN-ECE workshop report. Schriftenreihe der FAC 16. Libefeld. Critical levels for ozone:

Godzik, S.; Piskornik, Z. 1966. Transpiration of Transpiration of Aesculus hippocastanum L. leaves from air polluted and rural areas. L. leaves from air polluted and rural areas. Bulletin L. leaves from air polluted and rural areas.

Acad Polon. Sci., Ser.Sci. Biol. 14: 181-184.

Godzik, S.; Staszewski, T.; Kubiesa, P.; Szdzuj, J. 1996a. Monitoring biologiczny wojewodztwa katowickiego (Biological moni- Monitoring biologiczny wojewodztwa katowickiego (Biological monitoring of the Katowice administrative district). toring of the Katowice administrative district). Report 546/DN/95. Katowice: Instytut Ekologii Terenow toring of the Katowice administrative district).

Uprzemyslowionych.

Session I1 Ozone in the Upper Silesia Region – Concentration and Effects on Plants Godzik
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Godzik, S.; Staszewski, T.; Kubiesa, P.; Szdzuj, J. 1996b. Wykorzystanie roslin do oceny oddzialywania ozonu na srodowisko

(Use of plants in assessing ozone effects on environment). (Use of plants in assessing ozone effects on environment). Report 4 S401 014 05. Katowice: Instytut Ekologii Terenow (Use of plants in assessing ozone effects on environment). 

Uprzemyslowionych.

Godzik, S.; Szdzuj, J.; Osrodka, L.; Stawiany, W. 1995. Time and spatial differentiation of ground level ozone concentration

from four monitoring stations in the Upper Silesia Region - summer 1994. from four monitoring stations in the Upper Silesia Region - summer 1994. In: Godzik, S.; Starkel, L., eds. Ozone — a from four monitoring stations in the Upper Silesia Region - summer 1994.

global and regional problem. Polish National Commission on Global Change. Katowice: Institute for Ecology of Industrial Areas; 41-50.

Godzik, S.; Szdzuj, J.; Pawlinska, R. 1991. Proby oceny bezposredniego i posredniego udzialu zanieczyszczen powietrza w

uszkadzaniu drzewostanow poza rejonami przemyslowymi (An attempt to assess the direct and indirect effects of air

pollutants on trees outside industrial areas). pollutants on trees outside industrial areas). In: Poznawcze i praktyczne efekty badan prowadzonych w CPBP 03.11 w pollutants on trees outside industrial areas).

latach 1986 - 1990. Zabrze: PAN IPIS.

Legge, A.H.; Gruenhage, L.; Nosal, M.; Jaeger, H.J.; Krupa, S.V. 1995. Ambient ozone and adverse crop response: an evalua- Ambient ozone and adverse crop response: an evaluation of North American and European data as they relate to exposure indices and critical levels. tion of North American and European data as they relate to exposure indices and critical levels. In: Exceedances of critical tion of North American and European data as they relate to exposure indices and critical levels.

loads and levels. Vienna 1995 November 22-24, Austrian Research Centre Seibersdorf; 1-35.

Manning, W.J.; Feder, W.A. 1980. Biomonitoring of air pollutants with plants. London: Applied Science Publishers. Biomonitoring of air pollutants with plants. London: Applied Science Publishers.

Sanders, G.; Benton, J. 1995. Ozone pollution and plant response in Europe — an illustrated guide. The Nottingham Trent

University, Nottingham: UN ECE. The ICP-Crops Coordination Centre.

Session I1 Ozone in the Upper Silesia Region – Concentration and Effects on Plants Godzik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 65

Abstract

Ozone measurements are presented from the Bulgarian Govedartsi ecosystem study site, Ovnarsko No. 3. The site is

located on the south slope of the Govedartsi Valley in the northwestern part of the Rila Mountain area. The Rila is the

highest mountain in the Balkan peninsula and is representative of rural conditions in that part of Europe. The

experimental site, Ovnarsko, is situated at 1,600 m above sea level and about 10 km west of the village of Govedartsi.

There are no anthropogenic sources for ozone precursors for at least 30 km in all directions. Ozone concentrations

within the Govedartsi Valley are strongly influenced by thermally driven local winds from late spring to summer that

occur during high pressure synoptic weather conditions. In these cases a thin stable cold layer of downslope wind is

experienced during the night, separating surface air from the higher boundary layer air. This helps the reduction of

ozone concentrations from dry deposition measured at the Ovnarsko site. During the 1994 and 1995 measurement

periods no high ozone concentration episodes were detected. This may be because of either an absence of significant

anthropogenic sources of ozone precursors in this part of the Balkan peninsula or to the shielding of the Valley during

high pressure conditions. In general, ozone concentrations at Ovnarsko were below 35 ppb.

Introduction

In September 1991, an air-pollution study was launched at the Govedartsi in the

highest mountain range in the Balkans. The site—designated as Ovnarsko No. 3—

is located on the south side of the Govedartsi Valley on the northwestern part of

the Rila Mountains (Zeller 1991). The Valley is bordered on the south and west by

high ridges (~2,600 m), and on the north by a lower and more fractional ridge

(highest point ~1,800 m). The Valley opens to the northeast with a narrow mountain

pass, through which the Iskar river flows toward the Samokov area. The

experimental site Ovnarsko is situated at 1,600 m above sea level and about 10 km

west of the village of Govedartsi, which is on the less steep part of the northern

slope of Rila Mountain. The terrain is higher in the western part and lower in the

central part of Govedartsi Valley (fig.1). The greater part of Rila mountain lies to

the south of the Valley area.

There are no anthropogenic sources for ozone precursors for at least 50 km in

the south direction and for about 30 km in the west direction where the mountain

ridge is a high obstacle for west-to-east transport of air pollutants. Weekly average

measurements of SO4 

2-, NO3-

, and some metals taken in 1991 (Zeller and others

1992) and 1992 demonstrated increasing air pollutant concentrations after each

well-marked western air outbreak (strong winds accompanied by rain during

cyclonic synoptic weather). However, during high-pressure weather systems (low

wind and clear sky), decreased SO4 

2- and NO3-

 concentrations were observed. Under

the latter weather conditions mountain-valley gradient wind flows predominate,

allowing local natural sources of trace gases to determine the measured air quality.

During such periods the Govedartsi Valley is probably screened from

anthropogenic air pollutants from outside. In these cases the mountain ridges,

surrounding the Valley, are mostly above the height of the planetary boundary

layer causing the measured wind speeds at the experimental site to be low and wind

directions to follow a diel cycle of mountain-valley circulation.

This paper presents data that the Ovnarsko experimental site is not impacted

by ozone or its precursors of anthropogenic origin under high-pressure weather.

This phenomenon is associated with so called episodes of high ozone concentrations

Ozone Concentrations at the Bulgarian Govedartsi Ecosystem

Site in Early Summer of 1994 and 19951

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor, Sofia University, Department of Meteorology, Faculty of Physics, 5 James Baucher Blvd., 1126

Sofia,Bulgaria.

3 Research Scientist, Rocky Mountain

Forest and Range Experiment Station,

USDA Forest Service, 240 W. Prospect,

Fort Collins, CO 80526 USA.

4 Deputy Director, Forest Research Institute, Bulgarian Academy of Sciences,

132 Liment Ochridsky Street, 1080

Sofia, Bulgaria.

Evgeny Donev,2 Karl Zeller,3 and Christo Bojinov4
66 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

in rural areas in other regions of the world (Cox and others 1975, Logan 1985).

Many studies (Attmannspacher and others 1984, Guicherit and van Dop 1977,

Mueller 1994, Prior and others 1981, Vukovich and others 1977) relate the high

ozone levels to the presence of warm and slowly moving synoptic-scale high

pressure systems, which was not observed at the Ovnarsko site.

Data and Instrumentation

The reason for monitoring ozone at the Ovnarsko site was to determine the extent

to which upwind anthropogenic sources influenced Govedartsi Valley ozone

concentrations and whether the Ovnarsko site was representative as a background

station for this part of Europe. The site location allows for separating synoptic from

mesoscale meteorological wind regimes, isolating source regions for measured air

pollutants. Because of difficult site access during winter and limited technical

support after late summer, ozone concentrations were only measured from late

spring to summer: May 28 to July 9, 1994 (Julian Days 148 to 190) and June 12 to

July 30, 1995 (Julian Days 163 to 212). Wind speed and direction, air temperature

and humidity at 5 m height, total solar radiation, wetness, precipitation, soil

temperature and soil heat flux were measured through autumn. Hourly mean and

standard deviations of the measured quantities were recorded with a Campbell

Scientific 21x data logger (table 1).5␣ Ozone was sampled through a teflon filter at

2 m height. The ozone instrument was calibrated in the United States before

shipment to Bulgaria. It appeared to function normally; however, a post calibration

was not made. Because of the inherent stability of the ultraviolet adsorption

technique, the ozone concentrations were within ±5 ppb. The teflon filter used to

remove foreign objects in the sample air was changed approximately every 15 days.

Table 1 Table 1 — Parameters measured hourly at the Ovnarsko site. Table 1 

 Sensor Parameter Units

Teco 49 O3 ppb

Metone Wind speed m s-1

Metone Wind direction degree

Eppley Solar radiation watt m-2

MRI Inc. Precipitation mm

Metone Temperature °C

Metone Relative humidity pct

Rad. energy Soil heat flux watt m-2

thermocouple Soil temperature °C

thermocouple Soil temp Gradient °C

Campbell Sc. Leaf wetness wet/dry

Figure 1 — The Govedartsi

Valley from the northeast on a

100 m grid. The x axis, west-east,

is 15 km and the y axis, southnorth, is 10 km.

Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov

5 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

3000

2000

1000

2

4

6

8

10

4

6

8

10

12

14

2

South-North(km)

West-East(km)
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 67

Results and Discussion

A difference in diurnal ozone concentration cycle patterns was observed after sorting

the data by meteorological phenomena. Air laden with ozone and ozone precursors

is frequently transported from urban areas to cleaner mountain and rural areas

during periods with high atmospheric pressure and low wind speeds (Cox and

others 1975, Vukovich and others 1977, Wolff and others 1977). The days with such

weather are categorized: “days with clear skies and local winds” for this paper. On

the other hand, cyclonic weather conditions with strong winds, and turbulent

boundary layer air cause intense vertical mixing and potential dilution of polluted

air. In the Govedartsi Valley, however, this same vertical mixing appears to cause

downward ozone transport from higher ozone-laden layers of the troposphere. In

remote areas higher mean ozone concentrations are often experienced under these

more turbulent conditions when air, transported from distant upwind sources, is

mixed downward (Zeller and others 1977). For Ovnarsko the data were separated

into the two types of weather by using wind speed and direction and solar radiation

as the main criteria. Precipitation, air, and soil temperatures were used as additional

criteria when necessary.

Meteorology

The hourly and 24-hour mean wind direction and 24-hour wind speed sums for

the 1994 and 1995 measurement periods were obtained through summation of mean

values for each hour (fig.2a,b). There are two distinct meteorological regimes affecting

wind flow as measured at Ovnarsko: locally generated winds and synoptic winds.

During local wind days regional synoptic pressure gradients are weak and local

mountain-valley circulation predominates. On such days the wind direction follows

Figure 2 a, b — Wind direction

for each hour (line), mean wind

direction (squares), and velocity

for each day (diamonds) for the

two measurement periods in 1994

(a) and 1995 (b).

Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov
68 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

a typical diurnal cycle: southeast at night (mountain catabatic winds) and northeast

during the day (upslope winds). On days influenced by synoptic weather the diurnal

cycle of mean wind direction does not follow a predictable course but varies from

south to west demonstrating the effect of upwind regional air intruding into the

Govedartsi Valley. The strongest of these winds are from southwest and west during

such days and tending to return to southeast at night demonstrating the surface

layer decoupling from upper synoptic winds on most but not all synoptic nights.

The total solar radiation was calculated for each hour in addition to the daily

mean (fig. 3a,b). Days with high solar radiation (indicating few or no clouds) are

typically days when local winds occur (“days with clear skies and local winds”)

and vice versa: lower solar radiation (cloudy days) are the days with stronger

westerly winds. The 1994 study days determined as “days with clear skies and

local winds” are: Julian days 152-154 and 174-182, 12 days out of the 42-day

measurement session. For 1995 the study days are the Julian days 163, 183, 184,

194, 195 and 202-207—11 days out of the 50-day measurement session.

Ozone Concentration

Hourly and daily mean ozone concentrations from the 1994 and 1995 measurement

periods were plotted (fig. 4a,b). These values, adjusted to mean sea level, rarely

exceeded 35 ppb during the day and drop to 15 to 20 ppb at night. This diurnal

swing in concentration is similar to summertime results taken in a spruce-fir forest

in Snowy Range, Wyoming above 3,000 m elevation at 3 m above ground level but

below canopy height (Wooldridge and others 1994, 1995). Ozone concentrations

taken at 30 m above ground (13 m above canopy height) at the same Wyoming site

only vary 2 to 3 ppb from day to night (Zeller 1995). At Ovnarsko, higher nighttime

ozone concentrations were measured during the synoptic wind days when air from

Central or Southern Europe apparently entered the Valley (e.g., Julian days 156

Figure 3 a, b — Total solar

radiation (watts/m2

) for each hour

(line) and daily mean values

(squares) for the two measurement periods in 1994 (a) and

1995 (b).

Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 69

and 170 from 1994 and days 176 and 186 in 1995). There were no high ozone

concentration episodes during either classification period in 1994 and 1995. With

the exception of Julian days 154 in 1995 and 195 in 1996, periods when outer air

masses penetrated into the Valley (i.e., days without clear skies and local winds)

tended to be associated with similar daily maximum ozone concentration; however,

the nighttime minimums on such days were much higher, causing larger daily

averages. These data suggest that strong exchange of air within the whole boundary

layer during these periods enhanced vertical ozone transport, advected from distant

upwind sources, and maintained an above ridge-height ozone source preventing

the Valley from experiencing lower nighttime ozone concentrations. The lower

nighttime minimums during days with clear skies and local winds were from ozone

deposition to surfaces and the lack of a transport mechanism to vertically transport

the upper ozone to maintain higher surface concentrations. This explains the strong

influence of the local winds (mountain-valley circulation) on the diurnal cycle of

ozone concentrations in the Govedartsi ecosystem.

The existence of well-manifested thermally driven local wind system in the

Valley around the Ovnarsko site gives us the opportunity to compare the influence

of upslope and downslope winds on the diurnal variations of ozone concentration.

The mean diurnal cycle of ozone concentration, temperature, wind speed, and

direction (degree/10) was calculated for the chosen days with clear skies and local

winds. The thermally driven mountain-valley circulation is clearly seen especially

during the night when the cold downslope/down valley winds along the northnorthwest slope are constantly east-southeast to east in direction from 22:00 local

standard time until 8:00 LST. Northeast upslope winds begin after 8:00 LST until

20:00 to 21:00 LST. Diurnal ozone concentrations show a typical diurnal cycle for

the clear skies days in rural areas (Bottenheim and others 1994, Meagher and others

Figure 4 a, b — Hourly mean

ozone concentration (line) and

daily mean values (squares) for

the two measurement periods in

1994 (a) and 1995 (b).

Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov
70 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

1987, Prevot and others 1993): minimum values in the morning until upslope

conditions take over, and maximum values in the afternoon. During a typical

afternoon the east to northeast wind passes through the agrarian Govedartsi village

10 km E (upwind at this time) of Ovnarsko. There is little automobile traffic and

few farm vehicles that are not horse-drawn. It is possible some anthropogenic

ozone precursors come from Samokov, a larger city about 15 km farther east,

however, in general there is a lack of so called high ozone concentration episodes,

frequently experienced in the eastern part of the USA and in Western Europe under

similar weather conditions (Aneja and Li 1992, Aneja and others 1994, Davies and

others 1992, Mueller 1994, Trainer and others 1993).

In mountainous terrain, Brother and Gydax (1985) found an increase of ozone

during the day caused by upslope air transport with photochemical ozone

production. During the night ozone concentrations decreased on the slope due to

dry deposition. In another case Prevot and others (1993) concluded that thermally

driven winds defined the origin of air masses of the Reuss Valley in the summer

during high pressure systems. During the night the thickness of the downslope

wind layer experiencing efficient dry ozone deposition seemed to be very thin on

the slopes. They found that ozone concentrations at 2 m height were significantly

lower than those at 15 m height during the night similar to Wooldridge and others

(1994, 1995). The nocturnal concentrations measured at Ovnarsko at the same 2 m

height during downslope winds were rather low (fig. 5a, b) for a site at 1,600 m

elevation. The low nighttime Ovnarsko ozone values may be a result of dry

deposition of ozone to surfaces, including reactions with soil emitted NO (Brother

and Gydax 1985) within the cold stable downslope flow. The downslope flow

concentrated within a thin layer near the earth will not interact with the air mass

above, where ozone concentration is most likely higher and not effected by dry

deposition mechanisms. Mueller (1994) showed a clear tendency for diurnal ozone

amplitudes to diminish at rural mountain sites as site elevation increases. For some

mountain experimental sites a tendency for a reversed diurnal cycle of ozone

Figure 5 a, b — Mean diurnal

cycle of ozone concentration

(squares: parts per billion), wind

speed (diamonds: m/s), wind

direction (+: degree x 10-1), and

air temperature (x: °C) for the

two measurement periods in

1994 (a) and 1995 (b).

Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 71

concentration has been detected when maximum values are measured during the

night (Aneja and Li 1992). This is not the case for sites under the influence of a

mountain-valley circulation.

Apparently ozone concentrations are lower in this rural part of Bulgaria

compared to more populated areas. For instance, within southeast Sofia during

the end of the summer 1-m height, 10-day daily mean ozone concentrations were

between 40 and 50 ppb, with maximum values reaching 70 to 80 ppb. These Sofia

values are similar to those in metropolitan Denver, Colorado (Zeller 1995). Ozone

values found in forested areas of Poland are predominantly above 60 ppb and

often exceed 80 ppb (Bytnerowicz and others 1993), unlike the Ovnarsko results.

Conclusion

Ozone concentrations within the Govedartsi Valley are strongly influenced by

thermally driven local winds from late spring to summer during high pressure

systems. In these cases, a thin stable cold layer of downslope wind is experienced

during the night, which separates surface air from the higher boundary layer air

causing reduction of ozone concentrations from dry deposition measured at the

Ovnarsko site. During the two measurement periods, no high ozone concentration

episodes were detected. This condition may be a result of the absence of significant

anthropogenic sources of ozone and its precursors in this part of the Balkan peninsula

or from the shielding of the Valley during high pressure conditions where the

surrounding mountain ridges are above the boundary layer height and the air in

the valley flows as thermally-driven local circulation.

Acknowledgments

We thank D. Ivanov, A. Avramov and A. Tanev for technical assistance. The project

would have been impossible without support and instrumentation from Rocky

Mountain Forest and Range Experiment Station, USDA Forest Service, Fort Collins,

Colorado and USDA/Office of International Cooperation and Development,

Washington DC. We also thank Laurie Dunn for technical editing of this manuscript.

References

Aneja, V. P.; Zhen Li. 1992. Characterization of ozone at high elevation in the eastern U.S.: trends, seasonal variations and

exposure. Journal Geophysical Research 97: 9873-9888. exposure

Attmannspacher, W.; Hartmannsgruber, R.; Lang, P. 1984. Long period tendencies of atmospheric ozone based on

ozone measurements started in 1967 at the Hohenpeissenberg Meteorological Observatory. Meteorologische ozone measurements started in 1967 at the Hohenpeissenberg Meteorological Observatory

Runschau 37: 193-199.

Bottenheim, J. W.; Siros, A. Brice K. A.; Gallant, A. J. 1994. Five years of continuous observations of PAN and ozone at rural . Five years of continuous observations of PAN and ozone at rural

location in eastern Canada. location in eastern Canada. Journal Geophysical Research 99: 5333-5352. location in eastern Canada.

Brother, B.; Gydax H. A. 1985. The influence of locally induced wind systems on the effectiveness of nocturnal dry deposition

of ozone. of ozone. Atmospheric Environment 19: 1627-1637. of ozone.

Bytnerowicz, A.; Manning, W.J.; Grosjean, D.; Chmielewski, W.; Dmuchowski, W.; Grodzinska, K.; Godzik, B. 1993.

Detecting ozone and demonstrating its phytotoxity in forested areas of Poland: a pilot study. Environmental Detecting ozone and demonstrating its phytotoxity in forested areas of Poland: a pilot study

Pollution 80: 301-305.

Cox, R. A; Eggleton, A. E. J.; Derwent, R. G.; Lovelock; J. E.; Pack, D. E. 1975. Long-range transport of photochemical ozone in

north-western Europe. north-western Europe. Nature 255: 118-121. north-western Europe.

 Davies, T.; Kelly, D; P. M.; Low, P. S.; Pierce, C. E. 1992. Surface ozone concentrations in Europe: Links with the regional-scale

atmospheric circulation. Journal Geophysical Research 97: 9819-9832. atmospheric circulation

Guicherit, R.; van Dop, H. 1977. Photochemical production of ozone in western Europe (1971-1975) and its relation to meteo- Photochemical production of ozone in western Europe (1971-1975) and its relation to meteorology. Atmospheric Environment, 11: 145-155. rology

Logan, J. A. 1985. Tropospheric ozone: Seasonal behavior, trends and anthropogenic influence. Journal Geophysical Research Tropospheric ozone: Seasonal behavior, trends and anthropogenic influence

90: 10493-10482.

Meagher, J. F; Lee, N. T.; Valente, R. J.; Parkhurst, W. J. 1987. Rural ozone in the southern United States. Atmospheric Environ- Rural ozone in the southern United States

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Mueller, S. F. 1994. Characterization of ambient ozone levels in the Great Smoky Mountains National Park. Journal Applied Characterization of ambient ozone levels in the Great Smoky Mountains National Park

Meteorology 33: 465-472.

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Prevot, A. S. H.; Staenelin, J.; Richer, H.; Griesser, T. 1993. A thermally driven wind system influencing concentrations of A thermally driven wind system influencing concentrations of

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Trainer, M; Parish, D. D.; Aneja, V. P; Zhen, Li; Mita, Das. 1994. Ozone case studies at high elevation in the eastern U.S. Ozone case studies at high elevation in the eastern U.S

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Session I1 Ozone Concentrations at the Bulgarian Govedartsi Ecosystem Site Donev, Zeller, Bojinov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 73

Abstract

Ozone concentrations have been monitored at the Glacier Lakes Ecosystem Experiment Site (GLEES) in the Snowy

Range of the Medicine Bow Mountains 55 km west of Laramie, Wyoming, USA. The site is located at 3,186 m

elevation in a large subalpine meadow of a mature subalpine forest near timberline. Continuous ozone and

meteorological monitoring are a part of the GLEES research to determine the effects of atmospheric deposition on

alpine and subalpine ecosystems. Ozone monitoring has shown specific summer and winter diel ozone distribution

patterns, typical of remote, rural, high elevation sites. The data show relatively high background ozone concentrations

with little diel variation in winter. Stratospheric intrusions contribute to the ground-level ozone concentrations in

the spring. Summer ozone patterns show diurnal photochemical peaks and significant night time concentrations

suggesting lack of scavenging. The relationship of modeled leaf stomatal conductances, boundary layer conductances,

canopy conductances and photosynthesis rates to ambient ozone concentrations and measured ozone flux was

examined by using regression analysis. Ozone concentrations were also compared with wind, radiation, temperature,

and humidity conditions at the site by using regression analysis. The data provide information on ozone

concentration and environmental conditions at a remote, high elevation site typical of many wilderness areas of

the western United States.

Introduction

The Glacier Lakes Ecosystem Experiment Site (GLEES) is a 600-ha research site in

southeastern Wyoming, where atmospheric deposition and its effects on terrestrial

and aquatic ecosystem processes are being studied (Musselman 1994). The site is

located at 3,100 to 3,400 m asl elevation in the Snowy Range of southeastern

Wyoming, about 40 km west of Laramie. The research site is in the Medicine Bow

National Forest at the alpine/subalpine ecotone. Much of the atmospheric

monitoring is conducted at the Brooklyn Lake site, a forest stand dominated by

Engelmann spruce (Picea engelmannii (Parry) Engelm.) and subalpine fir (Abies

lasiocarpa (Hook.) Nutt.). The Brooklyn site is occupied by patchy forest with large

subalpine meadow openings.

This paper discusses the results of studies to determine the characteristics of

ozone deposition in alpine and subalpine ecosystems in southeastern Wyoming.

Methods

The Brooklyn meteorological and air quality monitoring site at the GLEES is located

in a small subalpine forest opening (30-m diameter). Electric power is available at

the site. Standard meteorological measurements are monitored continuously at 10

m and 29 m above the ground and recorded on a data logger. Parameters measured

at the site include wind speed and wind direction, soil and air temperature, radiation,

wetness, and relative humidity. Precipitation is monitored at 2 m above the ground

with an Alter shielded Belfort rain gauge.3 Air quality measurements include

ambient ozone and dry and wet deposition.

Dry deposition is monitored as part of two national networks, the National

Dry Deposition Network (NDDN) Centennial site, and the Interagency Monitoring

of Protected Visual Environments (IMPROVE) Brooklyn Lake site. The NDDN

site is part of a United States Environmental Protection Agency-sponsored national

network of dry deposition monitoring stations. Protocols for both dry deposition

monitoring networks use a filter pack for collection of dry particles and gases, but

Ozone and Modeled Stomatal Conductance at a High

Elevation Subalpine Site in Southeastern Wyoming1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside

California.

2 Plant Physiologist, Meteorologist, and

Research Associate, Rocky Mountain

Forest and Range Experiment Station,

USDA Forest Service, 240 West Prospect Road, Fort Collins, CO 80526-

2098.

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Robert C. Musselman, Karl F. Zeller, and Nedialko T. Nikolov2
74 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

they each conduct different analyses. The NDDN site is located in a larger opening

about 120 m southwest of the Brooklyn meteorological monitoring. The IMPROVE

monitor is located at the Brooklyn monitoring site meteorological tower.

Wet deposition is monitored as a part of the U.S. National Atmospheric

Deposition Program (NADP) national network, site Brooklyn Lake, WY95, which

monitors wet chemistry of precipitation. The NADP wet deposition station is also

located at the Brooklyn Lake Monitoring site, about 80 m north of the NDDN site.

Ozone has been monitored at 3,180 m asl at the GLEES Brooklyn Lake NDDN

monitoring site since 1989. Monitoring and calibrations follow standard NDDN

protocols. The station monitors ozone continuously at 3-m height above the ground.

Ozone flux measurements were also made at the site by using an eddy correlation

system (Zeller and Hehn 1996).

The FORFLUX model (Nikolov 1995) was used to estimate canopy

photosynthesis and stomatal CO2 conductance by using meteorological (March to

June 1990) data from the site. FORFLUX estimates canopy fluxes by numerical

integration of LEAFC3 model (Nikolov and others 1995) over stand leaf area index.

Leaf area index of 3.5 was used for this study. The modeled photosynthesis and

CO2 conductances were compared with values of ambient ozone using regression

analyses. The relationship was examined between the modeled canopy conductance

and (1) ambient ozone concentration or (2) ozone flux measurements. The ambient

ozone concentration data were also compared to measured meteorological data

by using regression analysis.

Results and Discussion

Typical ozone distribution patterns at the GLEES are characteristic of high elevation

remote sites. These patterns show low diel variation in ozone concentration, a result

of minimal daytime increase from photochemical production and minimal night

time decrease due to scavenging. The site also shows evidence of stratospheric

intrusion of ozone (Wooldridge and others 1996). Characteristics of the ozone

deposition at the GLEES have been described elsewhere (Wooldridge and others 1996,

Zeller and Hehn 1995). There are three main characteristics of the ozone distribution

for this remote site. First, concentrations of ozone are relatively high at the site (table 1).

Concentrations above 60 ppb occur frequently during the spring and summer

(Wooldridge and others 1996). Concentrations above 50 ppb are common.

A second characteristic is the diel variation of ozone concentration shows very

little amplitude. Diel concentrations vary as little as 1 to 3 ppb in January and

February (44 to 47 ppb). Diurnal variation and peak daily ozone concentration

increase monthly from March (minimum 48 ppb, maximum 51 ppb) to May

(minimum 51 ppb, maximum 58 ppb), with typical afternoon photochemical peaks.

The diel pattern in May and June is more pronounced, typical of increased

photochemical generation of ozone during the day and greater amount of ozone

scavenging by nitrogen oxides and surfaces at night. This difference indicates the

Session I1 Ozone and Modeled Stomatal Conductance at a High Elevation Subalpine Site Musselman, Zeller, Nikolov

Table 1 Table 1 — 1990-1994 average monthly ozone concentration at the GLEES Brooklyn monitoring site (Wooldridge and Table 1 

others 1996).

 Month Mean ozone concentration, ppb

January 44.6

February 47.5

March 50.5

April 53.9

May 54.6

June 49.8
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 75

importance of the lack of night time nitrogen oxide scavenging at this site during

the winter. In addition, there is very little daily photochemical generation of ozone

in winter. The lack of a diurnal peak is likely because of the limited amount of

precursors, and the very low temperatures that are not conducive for chemical

reactions. Temperatures at the site in January average less than -10 °C. The lack of

night time scavenging is likely due to the snow cover. Snow has been shown to be

a poor sink for ozone (Stocker and others 1995, Zeller and Hehn 1995).

The third unique characteristic is relatively high ozone concentration occurring

at this site even during midwinter, considering its remoteness, and the change in

average ozone concentration as the season progresses (Wooldridge and others 1996)

(table 1). Monthly average ozone concentration increases from January to May,

then decreases in June. There is evidence that stratospheric intrusion causes the

April/May ozone peak (Wooldridge and others 1996). Examination of the data

indicates high concentration ozone episodes with levels above 60 ppb that can last

for several days. These episodes are associated with very low (less than 25 percent)

relative humidity, suggesting downward intrusion of air from the stratosphere or

upper troposphere. The episodes occur frequently during April and May coincident

with passage through the area of cutoff low pressure centers. Ozone concentration

levels, during times of the month when these episodes occur, show almost no

evidence of diurnal photochemical ozone generation.

The June pattern has the highest diel variation (41 ppb to 52 ppb), yet the

average ozone concentration is reduced as the weather patterns for intrusions are

greatly reduced. Diurnal photochemical generation of ozone is more likely to occur

during June, and lack of a snow cover increases night time scavenging. Yet, average

low concentrations in June do not fall below 41 ppb, indicating considerably less

scavenging than in urban areas.

Output from the FORFLUX simulation model and ozone concentration was

examined, and no significant relationships were found between ambient ozone

concentration and modeled canopy photosynthesis or CO2

 conductance (fig 1).

Variability in the modeled parameters was high. There was no significant

relationship between solar radiation, temperature, wind speed, and relative

humidity and ambient ozone concentration. We have previously shown a

relationship between low humidity and high ozone concentrations (Wooldridge

and others 1996), and ambient ozone has been shown to be related to temperature

(Flaum and others 1996). There did appear to be a slight trend for increasing ozone

concentration with increasing solar radiation and temperature. These relationships

would reflect meteorological conditions favorable to ozone production at the site,

and would be expected to be stronger with summer data than with the spring data

presented here.

Plants often have a closer relationship to ozone flux than to ambient ozone

concentration (Musselman and others 1994). Modeled canopy CO2

 conductance

showed some relationship to measured ozone flux at GLEES site (fig. 2), in contrast

to the lack of relationship between conductance and ambient ozone concentration.

There are several reasons for the lack of a relationship between modeled

physiological parameters and ambient ozone. First, the model was not specifically

designed to simulate physiological response to ozone. Our objective was simply

to examine any possible relationship. Second, ozone concentrations are near the

lower threshold where plant response should occur; thus, the impact of ozone on

plant physiological response was limited. Third, these data represent early season

meteorological data. Greater effects should be expected later in the growing season

when plants are physiologically more active. CO2 uptake and conductance increase

as the growing season progresses (fig. 1). Although the ambient changes in ozone

concentration are real, the magnitude of the change as the season progresses is

comparatively small and cannot account for the changes in canopy photosynthesis

and conductance.

A relationship was found between canopy conductance and measured ozone

flux, although the r2

 value was low indicating high variability in the data. We

Session I1 Ozone and Modeled Stomatal Conductance at a High Elevation Subalpine Site Musselman, Zeller, Nikolov
76 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

expected that a greater relationship would be found between ozone flux and

conductance than between ambient ozone and conductance.

The lack of a significant relationship between ambient ozone and

meteorological factors is not surprising. The interaction of diurnal photochemical

production of ozone with the longer duration episodes of stratospheric input of

ozone and subsequent mixing complicate the dynamics of ambient ozone

concentration at any given time, particularly when examined for only 1 year. Multiyear data were necessary to determine the relationships previously reported

(Wooldridge and others 1996). Lack of significant soil and canopy surface sinks

and atmospheric chemical sinks at the site also tend to stabilize ozone concentration

during changing meteorological conditions. The relationship between ozone

concentration and meteorological parameters would be expected to be greater in

the summer when soil and canopy sinks are more pronounced and conditions for

photochemical production and atmospheric chemical scavenging are greater.

Figure 1 — FORFLUX model

simulation for GLEES, canopy

CO2

 conductance from midMarch through June, 1990 (A);

FORFLUX model simulation for

GLEES, canopy CO2

 uptake from

mid-March through June 1990

(B); and ambient ozone

concentration from mid-March

through June 1990 (C).

Ambient O3 Concentration (ppb) Canopy CO2 Uptake ( 

μmol m-2 s-1 ) Canopy CO2 Cond. ( mm m-2 s-1 )

FORFLUZ Model Simulation A

B

C

FORFLUZ Model Simulation

Measured

70 80 90 100 110 120 130 140 150 160 170 180 190 

Julian Day ( 1990 )

110

100

90

80

70

60

50

40

30

20

10

5

0

- 5

- 10

- 15

- 20

- 25

500

400

300

200

100

0

Session I1 Ozone and Modeled Stomatal Conductance at a High Elevation Subalpine Site Musselman, Zeller, Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 77

Conclusions

Ozone monitoring at a relatively remote high-elevation Rocky Mountain site showed

high concentrations of ambient ozone. Diel variation was slight during winter

months indicating little diurnal photochemical generation of ozone; but, diel

variation increased as the spring season progressed. Little night time scavenging of

ozone was evident when snow cover was present. There was strong evidence of

stratospheric intrusion of ozone during spring months, particularly during April

and May, associated with the passage of upper level low pressure troughs. Modeled

canopy physiological responses were not related to ozone concentration, but

modeled canopy CO2 conductance was related to ozone flux. There was no

significant relationship between meteorological parameters and ambient ozone

concentration for the 1990 data.

Figure 2 — Regression analysis

(A) between measured O3

 flux (B)

and modeled canopy CO2

conductance (C).

r2

 = correlation coefficient;

S = slope, and

I = intercept.

120

100

80

60

40

20

0

1.0

0.5

0.0

- 0.5

- 1.0

Measured O3 Flux ( μg m-2 s-1 )

0.2

0.0

- 0.2

- 0.4

- 0.6

 - 0.8

- 1.0

0 10 20 30 40 50 60 70 80 90

r

2 = 0.34

S = - 3.61

I = - 0.10

Modeled Canopy CO2 Conductance (mmol m-2 s-1)

Ozone flux ( μg m-2 s-1 )

B

A

Canopy CO2 Cond. ( mmol m-2 s-1 )

C

184 185 186 187 188 189 190 191 192 

Julian Day (1992)

Session I1 Ozone and Modeled Stomatal Conductance at a High Elevation Subalpine Site Musselman, Zeller, Nikolov
78 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Acknowledgments

We gratefully acknowledge the work of Dr. Gene L. Wooldridge in the original

summarization of the Brooklyn ozone data, and his analysis regarding stratospheric

intrusion of ozone at the GLEES. We thank Laurie Dunn for technical editing of this

manuscript.

References

Flaum, J.B.; Rai, S.T.; Zurbenko, I.G. 1996. Moderating the influence of meteorological conditions on ambient ozone concen- Moderating the influence of meteorological conditions on ambient ozone concentrations. trations. Journal of Air and Waste Management Association 46: 35-46. trations. 

Musselman, Robert C., ed. 1994. The Glacier Lakes Ecosystem Experiments Site. The Glacier Lakes Ecosystem Experiments Site. Gen. Tech. Rep. GTR-RM-249. Fort Collins, The Glacier Lakes Ecosystem Experiments Site.

CO: Rocky Mountain Experiment Station, USDA Forest Service; 94 p.

Musselman, Robert C.; McCool, Patrick M.; Lefohn, Allen S. 1994. Ozone descriptors for an air quality standard to protect

vegetation. Journal of Air and Waste Management Association 44: 1383-1390. vegetation

Nikolov, Nedialko T. 1995. Modeling transient response of forest ecosystems to climatic change. In: Tinus, R.W, ed. Interior Modeling transient response of forest ecosystems to climatic change

west global change workshop proceedings; 1995 April 25-27; Gen. Tech. Rep. GTR-RM-262. Fort Collins, CO: Rocky

Mountain Experiment Station, USDA Forest Service.

Nikolov, Nedialko T.; Massman, William J.; Schoettle, Anna W. 1995. Coupling biochemical and biophysical processes at the

leaf level: an equilibrium photosynthesis model for leaves of C3 plants. plants. Ecological Modeling 80: 205-235. plants. 

Stocker, David W.; Zeller, Karl F.; Stedman, Donald H. 1995. O3

 and NO2 fluxes over snow measured by eddy correlation. fluxes over snow measured by eddy correlation.

Atmospheric Environment 29: 1299-1305.

Wooldridge, Gene; Zeller, Karl; Musselman, Robert [In press]. Ozone concentration characteristics in and over a high-eleva- Ozone concentration characteristics in and over a high-elevation forest. tion forest. Theoretical and Applied Climatology. tion forest. 

Zeller, Karl; Hehn. Ted 1995. Ozone deposition in a snowcovered subalpine spruce-fir environment. Ozone deposition in a snowcovered subalpine spruce-fir environment. In: Biogeochemistry of Ozone deposition in a snowcovered subalpine spruce-fir environment.

seasonally snow-covered catchments. Proceedings of a symposium; July 1995, Boulder, CO.: IAHS Publ. No. 228; 17-22.

Zeller, Karl; Hehn. Ted 1996. Measurements of upward turbulent ozone fluxes above a subalpine spruce-fir forest. Measurements of upward turbulent ozone fluxes above a subalpine spruce-fir forest. Geophysi- Measurements of upward turbulent ozone fluxes above a subalpine spruce-fir forest. 

cal Research Letters 23(8): 841-844.

Session I1 Ozone and Modeled Stomatal Conductance at a High Elevation Subalpine Site Musselman, Zeller, Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 79

Abstract

Bulgaria’s forest ecosystems (31 percent of the country’s area) are considered vulnerable to dry and wet pollution

deposition. Coniferous forests that cover one-third of the total forest land are particularly sensitive to pollution loads.

The USDA Forest Service, Sofia University, and the Bulgarian Forest Research Institute (FRI) established a cooperative

program to study a Rila Mountain forest ecosystem site, its climatology, air quality, surrounding watershed, hydrology,

and soil characteristics. The cooperative program was initiated in September 1991 by upgrading an existing climatology

site: Ovnarsko no.3 in the Govedartsi Valley on the north slope of the Rila Mountains. The dominant tree species

surrounding the site include Scotch pine (Pinus sylvestris L.), black pine (Pinus nigra Arn.), Norway spruce (Picea

abies Karst.), white fir (Abies alba Mill.), beech (Fagus sylvatica L.), and several oak species (i.e., Quercus petraea

Liebl, Q. pubescens Willd., Q. conferta Kit., Q. cerris L., and Q. robur). Weekly concentrations of nitrate, sulfate, sulfur

dioxide, calcium, sodium, and potassium were measured using a filter-pack technique. Data for 18 weeks between October

1991 and November 1993 were collected. Annual acidic aerosol deposition values for 1992 and 1993 were estimated for SO4

2

,

which was slightly above 3 kg ha -1 yr-1, and for NO3

-

, which decreased from 1.3 to 0.3 kg ha-1 yr-1.

Introduction

The Bulgarian Higher Institute of Forestry (HIF), Govedartsi Valley forest

meteorological monitoring site, Ovnarsko no. 3, was upgraded in September 1991

(Zeller 1991). The manual recording protocol was enhanced to include continuous

hourly measurements of standard meteorology and a filterpack system to determine

pollutant dry deposition by direct measurements (Zeller and others 1992). A

memorandum of understanding, MU RM-93-164, entitled “Atmospheric Interaction

with Natural Ecosystems in Bulgaria and the U.S.A.” was executed in July 1993

between the Bulgarian Forest Research Institute (FRI), Sofia University Department

of Meteorology (SUDM), and the USDA Forest Service’s Rocky Mountain Forest

and Range Experiment Station (RMFRES) to formalize the cooperative effort.

This paper describes the Rila Mountain forest ecosystem site in Bulgaria and

its surrounding watershed, location, forest, hydrology, geology, soils, and climate

by using land manager’s records and the European International Food and

Agriculture Organization (FAO) system (Bojinov 1981, FAO 1989); and it presents

sulfate and nitrate measurements resulting from the cooperative effort.

Location and Methods

Bulgaria is located in the southeastern part of the Balkan peninsula south of

Romania, east of the Black Sea, north of Greece and west of Serbia. Forests cover

about 33,300 km2, 31 percent of the country’s area. About 37 percent of the forested

area is occupied by coniferous forests, 19 percent by seed-originated broad-leaved

stands, and 44 percent by deciduous coppices of low productivity. The dominant

tree species include Scotch pine (Pinus sylvestris L.), black pine (Pinus nigra Arn.),

Norway spruce (Picea abies Karst.), white fir (Abies alba Mill.), beech (Fagus sylvatica

L.), and several oak species (i.e. Quercus petraea Liebl, Q. pubescens Willd., Q. conferta

Kit., Q. cerris Lq., and Q. robur).

Ovnarsko no. 3 (elevation 1,600 m) is located 4 km west of Govedartsi, Bulgaria,

situated on the north slope of the Rila Mountains (see site map in Donev and

others, this volume). Govedartsi (cattle place) is a small, agrarian village that sports

Bulgarian Rila Mountain Forest Ecosystems Study Site:

Site Description and SO4

2-, NO3-

 Deposition1

1 An abbreviated version of this paper

was presented at the USDA Forest Service International symposium on Air

Pollution and Climate Effects on Forest

Ecosystems, February 5-9, 1996, Riverside, California.

2 Research Scientist, Rocky Mountain

Forest and Range Experiment Station,

USDA Forest Service, 240 W. Prospect,

Fort Collins CO 80526 USA.

3 Deputy Director, Forest Research Institute, Bulgarian Academy of Science 132

Kliment Ochridsky Str., 1080 Sofia,

Bulgaria.

4 Professor, Faculty of Physics, Department of Meteorology, Sofia University,

5 James Baucher Blvd., 1126 Sofia,

Bulgaria.

5 Visiting Scientist, Rocky Mountain Forest and Range Experiment Station,

USDA Forest Service, 240 W. Prospect,

Fort Collins CO 80526 USA.

Karl Zeller,2 Christo Bojinov,3 Evgeny Donev,4 Nedialko Nikolov5
80 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

a small commercial ski lift, Maljovica, 13 km southwest. Govedartsi is about 70

km south of Sofia, the capital city of Bulgaria, and 14 km southwest of Samokov, a

small city. The climate discussion herein is specific to Ovnarsko no. 3. The forest,

hydrology, geology and soils discussions are for the general area on the north

slope of the Rila Mountains and Govedartsi Valley.

The northern slopes of the Rila Mountains are located in the south of Sofia

County. The forested areas are managed by the Samokov and Borovets forest

districts and are typical Bulgarian forest regions. In addition to Govedartsi are the

small villages Madgare, Mala Tsarkva, Borovets, and Beli Iskar. The Borovets

meteorological station location is also a famous all-season national resort of the

same name. The general study area described here includes three parts of four

north slope Rila Mountain morphographic spurs: Malyovishki (northwestern);

Skakavishki (central); and Moussalevsni (eastern). The area has the shape of a leaf

with its long fingers about 26 km long and about 12 km wide covering latitudes:

42°06' - 42°17' N; and longitudes 23°12' - 23°40' E. The terrain is very rugged, with

clearly outlined hillsides and deeply incised rivers, streams, and ravines. Elevation

of the area described is 1,050 - 2,200 m.

Ovnarsko Site Instrumentation Upgrade

Wind speed and direction, air temperature, and humidity sensors are installed at

5 m height; total solar radiation, wetness (actually psuedo-wetness: an electronic

plate that simulates a leaf surface and records the value 1 when wet and 0 when

dry), precipitation, soil temperature, and soil heat flux measurements are also

collocated. Hourly means and variances of the measured quantities were recorded

with a Campbell Scientific 21x data logger6 at 0.1 hertz. Ozone was sampled during

the spring and summer months of 1994 and 1995 (Donev and others[this volume])

at 2 m height (table 1).

6 Mention of trade names or products is

for information only and does not imply endorsements by the U.S. Department of Agriculture.

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov

Table 1 – Parameters measured at the Ovnarsko site. Table 1

Sensor Parameter Period Units

Filter Pack SO4

2- week μg m-3

Filter Pack NO3

- week μg m-3

Filter Pack Mg2+ week μg m-3

Filter Pack Ca2+ week μg m-3

Filter Pack Na+ week μg m-3

Filter Pack K+ week μg m-3

Dasibi 1000H Ozone hour ppb

Pump flow Sample volume hour liter pm

MetOne, Inc Wind direction hour Degree, Deg2

MetOne, Inc Wind speed hour m s-1, m2s-2

Eppley Solar radiation hour watt m-2

MRI Inc. Precipitation hour mm

MetOne, Inc. Temperature hour ° C

MetOne, Inc. Relative humidity hour pct

Rad Energy Soil heat flux hour watt m-2

thermocouple Soil temperature hour ° C

thermocouple Soil °C gradient hour ° C

Campbell Sc. Pseudo wetness hour wet:1;dry:0
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 81

Particulate and Gaseous Ambient Concentration

Pollutant concentration measurements were made by drawing ambient air at the

rate 2.0 ± 0.2 liters per minute (24 hr day-1, multi-day period-1) through a three-stage

filter pack mounted on the meteorology mast at 4 m above ground level. The filter

pack system was designed and supplied by the U.S. National Biological Service/

Environmental Science and Technology Center (ESTC). The filter pack inlet is 0.95

cm in diameter, which allows for the collection of aerosols smaller than about

10 mμ diameter when oriented downward. The filter pack itself was mounted under

a 20.3 cm diameter polyvinyl chloride (PVC) rain shield. Air flow is volumetrically

controlled with a diaphragm pump, pelton-wheel flow meter. Filter packs were

exchanged periodically as indicated in the record.

Filter packs provide a convenient, low-cost method for measuring acidic air

pollutants (Anlauf and others 1985). Various acid gases and particles are retained

on the stacked filters. Wet chemistry extraction of the filtered material yield

pollutant masses. The filter pack chemistry used at the Ovnarsko site is the same

used for the U.S. Environmental Protection Agency National Dry Deposition

Network (NDDN) (Edgerton and others 1989). Three filters are placed in series

within an inert Teflon holder: first, teflon “Zeflour” 2μm filter traps suspended

particulate aerosols; next, a nylon “Nylasorb” 1μm filter absorbs acidic gases such

as nitric acid and sulfur dioxide; and last, two Whatman 41 ashless cellulose filters

infuse a 25 percent K2

CO3 solution and glycerol SO2 is oxidized to SO4

2- when passed

over K2CO3, and retained as SO4

2- on the Whatman filters (Zeller and others 1996).

Filter pack preparation, extraction, and analysis was accomplished by a

Bulgarian Laboratory under the supervision of the FRI as outlined in the “NDDN

Laboratory Operations Manual” (Hunter/ESE 1989). Filter specific laboratory

species weights are divided by the measured air volume that passed through the

filter pack to provide average concentrations: SO4

2- (teflon filter SO42- mass) and

NO3

-

(teflon filter NO3

-

 mass).

Record Data

Forest

Rila Mountain northern slope forests are mostly indigenous conifers and are

managed for timber production. Human occupations that have influenced forest

health and use in the Rila Mountain area include iron and gold mining, range

management, logging, agriculture, and in the past charcoal production associated

with mining. The study area can be examined by forest activity, forest type, and

density (table 2) (Bojinov 1981). Total timber production area is 16,845 hectares.

Table 2 Table 2 — Forest type, density, and area of the Rila Mountain site. Table 2 1

 Type Coniferous Broadleaf Broadleaf Total Percent Percent

Density 0.4-1.0

 Indigenous 13,641 556 14,197 73.2

 Crop 590 50 640 3.3

Density 0.1-0.4

 Indigenous 1,696 30 1,726 8.9

 Crop 51 0 51 0.3

Density 0.0

 Seeded 230 0 230 1.2

 Dwarf pine 97 97 0.5

 Bare lands 2,442 2,442 12.6

1 ha

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
82 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The majority of timber production occurs on slopes between 10 to 30 percent

steepness (table 3). Bulgarian forests are also classified by slope orientation exposure:

sunny (SE, S, SW, and W) 4,498 hectares (ha); and shady (NW, N, NE and E) 12,348

ha. Table 4 gives a breakdown by elevation.

The main species composition of the study area include: Scots pine (Pinus

sylvestris), Norway spruce (Picea abies), silver fir (Abies alba) and Macedonian pine

(Pinus peuce). There are, however, 23 different coniferous species, including 10 Pinus

species, and 54 different broad-leaved species including 8 Acer (Delkov 1988,

Stefanov and Ganchev 1958). Sensitivity of these Bulgarian tree species to human

activities has been expressed as the resistance of each tree species to air pollutants

(Prokopiev 1978). Table 5 (Bojinov 1981) gives the coverage area by main species

and age.

Silver fir is most sensitive to environmental factors. It mostly inhabits cool

sites where winter temperatures are moderate. It is particularly sensitive to extreme

temperatures, especially spring frosts. Silver fir only occupies a small part of the

study region: shady and semi-shady sites with mean annual temperatures between

4.5 and 7.5 °C.

Scots pine is a species with almost no sensitivity to particular environmental

factors. It tolerates low tundra-like temperatures to high steppe-like temperatures.

It withstands extreme temperatures and quickly colonizes bare areas.

Macedonian pine expresses itself as a native species. It demands high and

permanent moisture, particularly high humidity. Macedonian pine trees usually

grow in a narrow strip above 1,800 m where environmental conditions are favorable

to their development or in areas with mean annual temperatures around 4 °C.

The range of Norway spruce coincides with that of silver fir. In the lower

portions of the mountains, it begins at regions with mean annual temperatures of

6.5 °C and grows to elevations with mean annual temperatures of about 2.5 °C.

Table 5 — Table 5 — Rila Mountain forest by age and coniferous type. Table 5 — 

Type / Age (yrs.) Type / Age (yrs.) 0 - 40 40 - 80 80 - 120 120 - 160 Percent Percent

Scots pine (ha) 922 3,792 1,112 50 37

Norway spruce 593 1,629 2,796 329 31

Silver fir 25 297 824 21 7

Macedonian pine 185 669 1,117 99 13

Mixed coniferous 124 651 989 93 12

 Table 3 — Rila Mountain forest by area and slope. — 

 Slope

(percent) 0-4 5-10 11-20 21-30 >30

Area (ha) 152 977 6,081 7,901 1,735

 Table 4 — Table 4 — Rila Mountain forest by area and elevation. Table 4 — 

 Evevation

Elevation(m) <1,300 1,301-1,500 1,501-1,700 1,701-1,900 >1,900

Area (ha) 4,354 5,294 3,439 2,717 1,042

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 83

Hydrology, Geology and Soils

The headwaters of the Iskar and the Marica, Bulgaria’s two longest rivers, are located

within the study area. The Iskar flows north to the Duna (Danube) and the Marica

flows southeast to Turkey and eventually the Aegean Sea. Numerous mountain

lakes are situated deeply within the Rila alpine at elevations above the study site.

The Rila Mountains are composed of intrusive and metamorphic rocks. South

Bulgarian granite is a representative for the first group and is the major component

of the mountain core. This granite is found mostly in the south half of the study

area just beneath the soil surface. Metamorphic rocks are found in the north half of

the study area and are represented mainly by gneiss and schists. Sandstones and

conglomerates are found to a lesser extent and lime rocks as well as mechanical

sediments represented by gravels and sands also occur. The Rila Mountains were

formed during the Paleozoic era (the Hercynian folding) and were strongly

denuded in the Mesozoic era. In the beginning of the Tertiary, the study area

experienced vertical movements that caused the formation of the surrounding

kettle, such as the inner Govedarska Kettle Hole (Bojinov 1981).

The soils in the study area are humic, entric and distric cambisols (table 6).

Humic cambisols, which are the deepest and most stony, are distributed in the

form of a belt in the south portion of the study area above the granite. These soils

occur mainly under stands of Macedonian pine and Norway spruce, and pure and

mixed Norway spruce stands that comprise 5,027 ha or 30 percent of the study

area timber. Entric cambisols, the next deepest and moderate to slightly stony,

occupy 9,815 ha (58 percent of the timber area) and are located in the north and

lower elevations of the study area above the gneiss. These soils are dominated by

Norway spruce and silver fir with some Macedonian and Scots pines. Distric

cambisols, which are only slightly stony, are situated mainly on south-facing slopes

in the NW portion of the study area and total 2,004 ha (12 percent) of the timber

with mostly Scots pine stands.

Moisture reserves in all north slope soils are favorable for tree vegetation

and growth (table 6). These soils are also well-supplied with nutrients (humus

and total nitrogen), especially the humic and entric cambisols (table 6). The

acidity of distric cambisols varies within the narrowest range (5.2 to 5.4); humic

cambisols (3.6 to 5.2) are similar but slightly lower compared to the entric

cambisols (4.2 to 5.7).

Climate

The climatology of Ovnarsko was recorded between 1961 and 1970 (Serafimov 1978)

before the site was upgraded (table 7). Other climate stations in the study area

include: Dolna Bania (42° 18’ N, 23° 45’ E; 635m); Samokov (42° 19’ N, 23° 34’ E;

1,030m); Borovets (42 ° 15’ N, 23 ° 37’ E; 1346m); Sitniakovo (42° 14’ N, 23° 37’ E;

1,742m); Yazavor Beli Iskar (42° 09’ N, 23° 16’ E; 2390m); Hija Musala (42° 12’ N, 23°

26’ E; 2,392m); and Vrah Musala (42° 11’ N, 23° 35’ E; 2,925m).

Table 6 — Average characteristics of soil types on the north Rila Mountain slopes (Bojinov 1981). Table 6

 Depth/humus Water capacity Nutrient Nutrient

 Type Horizon (cm) Horizon (cm) Total/active Humus/total N pH

 (mm) (t/ ha) (t/ ha)

Humic cambisol 90/35 433/382 442/17 4.4

Eutric cambisol >80/25 393/359 426/20 5.0

Dystric cambisol <80/<20 146/139 83/3 5.3

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
84 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Precipitation chemistry was sampled in 1989 and pH was below 4.5 (table 8).

Because there were no nearby sources causing low pH precipitation and an exact

record does not exist, it was expected that the Govedartsi valley should be clean of

air pollutants. The low pH measurements (table 8) were a surprise at the time

(Serafimov 1978) and remain unexplained. The concentrations of nitrates are low.

Concentrations of chlorides and sulfates and concentrations of heavy metals are

indicative of those naturally occurring.

Sulfate (SO4

2-) and Nitrate (NO3-

) Concentrations and Deposition

A total of 15, multi-day filter pack samples exposed during summer and autumn

periods between May 6, 1992 to November 6, 1993 were recorded (table 9). The

three initial samples taken between the period September 11, 1991 to October 27,

1991, as reported by Zeller and others (1992) were atypical and not repeated here.

Deposition in kilograms per hectare per year (kg ha-1 yr-1) was calculated by assuming

0.4 cm s-1 deposition velocity (Bytnerowicz and others 1987). If the 1992 and 1993

data are extrapolated for an entire year, NO3

-

 deposition would be 1.3 and 0.3

kg ha-1 yr-1 respectively. Deposition for SO4

2- would be 3.6 and 3.3 kg ha-1 yr-1

respectively. These last deposition estimates give an indication of annual values

only and should be used with caution because measurements were never taken for

the entire year, nor were any measurements taken during winter and spring seasons.

Sulfate concentrations are similar to those found in other parts of Europe

(Ruijrok and others 1995) but below those in more polluted regions (Zeller and

Table 8 Table 8 — Precipitation chemistry composition of 1989 samples with < 4.5 pH. Table 8 

Ion NOx Cl SO x Pb Cu Mn Ni Cd

mg L-1 (FAO 1989) 0.58 0.44 1.4 11.5 1.6 12.5 2.0 0.8

Table 7 — Table 7 — Ovnarsko no. 3 Climatology, 1961-1970. Table 7 — 

 Parameters Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

2m temperatures - 5.4 - 3.4 - 1.4 3.1 8.2 11.7 13.5 13.1 9.9 5.6 2.7 - 2.2

within stand ( °C )

2m temperature - 4.9 - 2.8 - 0.6 4.4 9.5 13.0 14.8 14.1 10.4 6.1 3.2 - 2.0

in open ( °C )

2m max temperature - 1.7 0.4 1.8 6.9 12.2 15.3 17.4 17.3 14.1 9.6 5.6 0.8

within stand ( °C )

2m max temperatures - 0.2 2.7 3.6 9.1 13.9 16.8 18.9 19.3 16.2 11.8 7.4 2.5

in open ( °C )

2m min temperature - 8.0 - 6.3 - 4.2 - 0.1 4.1 7.2 8.7 8.6 6.4 2.6 -0.5 - 6.0

within stand( °C )

2m min temperature - 8.7 - 7.5 - 4.6 - 0.5 3.9 6.9 8.2 8.2 5.7 2.0 - 0.9 - 5.6

in open ( °C )

2m wind speed 0.25 0.31 0.25 0.28 0.17 0.16 0.16 0.16 0.20 0.26 0.25 0.28

stand (m s-1)

2m wind speed 0.73 0.85 0.77 0.81 0.59 0.53 0.53 0.55 0.69 0.79 0.90 0.91

open (m s-1)

Precipitation (mm) 71 61 66 82 106 120 90 63 57 69 72 70

2m RH in 86 83 82 78 77 79 76 75 76 75 79 85

stand (pct)

2m RH in open 84 81 79 72 72 73 70 71 73 72 76 84

(pct)

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 85

others 1996). The projected annual SO4

2- aerosol deposition is similar to results for

higher polluted regions (Zeller and others 1996), where deposition velocities were

individually modelled and typically below 0.4 cm s-1, which might explain the

comparison difference between concentration and deposition results. Nitrate

concentration and deposition at Ovnarsko are lower than those found in more

polluted regions.

Conclusion

The cooperative Bulgarian Rila Mountain forest ecosystems study site, Ovnarsko

no. 3, has been described with initial dry deposition results. Although data were

not collected for the winter and spring seasons, results indicate SO4

2- and NO3-

 aerosol

concentrations are similar to other less polluted regions of Europe, while deposition

of NO3

-

 is also low and deposition of SO4

2- may be similar to higher polluted areas.

This result is dependent on modelled deposition velocity and needs further study.

Acknowledgments

We are grateful for the following support: Al Reibau, Lucenda Smith and Michael

Sestak NBS/ESTC, Fort Collins, Colorado for providing the filter pack pump system;

the USDA Office of International Cooperation and Development for initial site

survey funds; Mr. Spas Ashkov and staff, FRI Ovnarsko station, Govedartsi for

local site support, and Danko, Ltd. for electronic support. We also thank Laurie

Dunn for technical editing of this manuscript.

References

Anlauf, K.G.; Fellin, P.; Wiebe, H.A.; Schiff, H.I.; MacKay, G.I.; Braman, R.S.; Gilbert, R. 1985. A comparison of three meth- A comparison of three methods for measurement of atmospheric nitric acid and aerosol nitrate and ammonium. ods for measurement of atmospheric nitric acid and aerosol nitrate and ammonium. Atmospheric Environment ods for measurement of atmospheric nitric acid and aerosol nitrate and ammonium.

19: 325-333.

Bojinov, Christo. 1981. Determining and mapping of the types of the forest sites, and optimizing the composition of tree

species on the northern slopes of the Rila Mountains. Bulgarian Higher Institute of Forestry. 150 p. Ph.D. dissertation. species on the northern slopes of the Rila Mountains

Table 9 Table 9 — Filter pack concentrations and calculated deposition. Table 9 1

Concentration Deposition

Days (μg m-3) kg ha-1 yr-1

exposed Date On NO3

- SO42- NO3- SO42-

32 05/06/92 0.17 4.30 0.21 5.42

 8 06/06/92 0.26 2.03 0.33 2.56

 7 06/14/92 0.38 2.17 0.48 2.74

 7 06/21/92 0.27 1.48 0.34 1.87

 7 06/28/92 0.18 1.80 0.23 2.27

 7 07/05/92 4.89 5.20 6.17 6.56

16 06/13/93 0.39 4.19 0.49 5.29

17 06/29/93 0.59 5.25 0.74 6.62

20 07/16/93 0.14 2.13 0.17 2.69

17 08/05/93 0.20 1.31 0.25 1.65

17 08/22/93 0.17 1.17 0.21 1.48

17 09/08/93 0.22 3.09 0.27 3.90

15 09/25/93 0.14 1.78 0.17 2.25

13 10/10/93 0.16 2.38 0.20 3.00

14 10/23/93 0.22 2.29 0.27 2.89

1

 0.4 cm s-1 deposition velocity.

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
86 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Bytnerowicz, Andrzej.; Miller, Paul, R.; Olszyk, D.M. 1987. Dry deposition of nitrate, ammonium and sulfate to a Dry deposition of nitrate, ammonium and sulfate to a Ceanothus

crassifolius canopy and surrogate surfaces. Atmospheric Environment 21(8): 1749-1757. canopy and surrogate surfaces

Delkov, N. 1988. Dendrology. Sofia: Zemizdat; 306 p. Dendrology

Donev, Evgeny; Zeller, Karl; Bojinov, Chistro 1998. Ozone concentrations at the Bulgarian Govedartsi ecosystem site in early

summer of 1994 and 1995. In: Bytnerowicz, Andrzej, technical coordinator. Proceedings of the international symposium summer of 1994 and 1995

on air pollution and climate change effects on forest ecosystems; 1996 February 5 - 9; Riverside, CA: Gen. Tech. Rep. PSWGTR-166. Albany, CA: Pacific Southwest Research Station, USDA Forest Service; [this volume].

Edgerton, Eric; Lavery, Thomas; Prentice, Horace. 1989. National dry deposition network: third annual progress report. National dry deposition network: third annual progress report. U.S. National dry deposition network: third annual progress report.

EPA Atmospheric Exposure and Assessment Laboratory, EPA/600/3-91/018,. Research Triangle Park, NC: U.S. EPA; 89 p. ,

FAO. 1989. Manual for methodology and criteria for harmonized sampling, assessment monitoring and analysis of the effects

of air pollution on forests. Geneva: European Forestry Commission Programme Task Force; 203 p. of air pollution on forests

Hunter/ESE. 1989 National dry deposition network: laboratory operations manual. National dry deposition network: laboratory operations manual. prepared for U.S. EPA by Hunter/ESE, National dry deposition network: laboratory operations manual.

EPA Contract Doc # 86-612-0212-3170, Research Triangle Park, NC.: Hunter/ESE 183 p.

Prokopiev, E. 1978. Afforestation of industrious region. Sofia: Zemizdat; 216 p. Afforestation of industrious region

Ruijgrok, W.; Davidson, C.I.; Nicholoson, K.W. 1995. Dry deposition of particles: implications and recommendations for

mapping of deposition over Europe. mapping of deposition over Europe. Tellus 47B: 587-601. mapping of deposition over Europe.

Stefanov B.; Gantchev, A. 1958. Dendrology. Sofia: Zemizdat; 423 p. Dendrology

Zeller, Karl. 1991. US Forest Service - Bulgarian forest pollution project. Regional Environmental Center for Central and East- US Forest Service - Bulgarian forest pollution project

ern Europe, Information Bulletin 1(4): 16.

Zeller, Karl; Kichukov, Emil; Mirtchev, Stefan; Nikolov, Nedalko; Bojinov, Christo 1992. Status of air pollution impact on

forest ecosystems in Bulgaria. forest ecosystems in Bulgaria. AWMA, 92-71.01, 85th annual meeting; 1992 June 21-26; Kansas City, MO: Air and Waste forest ecosystems in Bulgaria.

Management Association; 10 p.

Zeller, Karl; Cerny, Martin; Smith, Lucenda; Sestak, Michael; Michalic, Miroslav; Perneg, Vindsor; Kucera, Jura 1996. Weekly

dry deposition of NO3

-

 and SO4

2- at Brdy Mountains, Czech Republic. In: Proceedings of NATO-advanced research work- at Brdy Mountains, Czech Republic

shop: atmospheric deposition and forest management. 1996 April 23-26; Spindleruv Mlyn, Krkonose Mtn., Czech Republic: NATO.

Session I1 Bulgarian Rila Mountain Forest Ecosystems Study Site Zeller, Bojinov, Donev, Nikolov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 87

Abstract

Boreal forests and woodlands comprise about 29 percent of the world’s forest cover. About 70 percent of this forest is in

Eurasia, mostly in the Russian Federation. Boreal forests contain about 45 percent of the world’s growing stock and

are an increasingly important part of global timber production. Fire impacts large areas of boreal forest annually in

Russia, and in excess of 60 percent of these fires are believed to be caused by human activity. Because of the large extent

of boreal forest, the large amount of fire activity, and the expected sensitivity of carbon cycling in these systems to fire

and climate patterns, there is justifiable concern about the potential effects of boreal fire regimes on atmospheric

chemistry, global carbon cycling, and global climate, as well as potential feedback effects of global climate change on

fire regimes and carbon cycles in these systems. Accurate estimates of these impacts are hampered by a lack of good

data on the area burned and on fire severity. We estimate that the average area burned annually in Russian boreal

forests is around 7.3 million ha, an order of magnitude greater than indicated by official fire statistics. This discrepancy

has major implications for estimates of emissions of CO2 and other greenhouse gases. In severe fire seasons, there are

also frequent long-term episodes of air pollution caused by wildfire smoke. Because of the interactions between fire,

landscape patterns, air quality, and climate in boreal forests, fire regimes and fire management in Siberia have

potentially large impacts on regional air quality and on the global environment.

Introduction

Boreal forests and woodlands cover a large area of the earth’s surface between 45

and 70 ̊ north latitude. Boreal forest and woodland cover an estimated 1.2 billion

ha, about 900 million ha of this in closed boreal forest (FIRESCAN 1994). This

represents about 30 percent of the world’s forested area. More than 50 percent of the

world’s forests are in North America and Eurasia (Nyejlukto 1994). Russia alone

contains about 620 million ha of boreal forest (Tchebakova and others 1994), about 520

million ha of which is in closed (stocked) boreal forest (Alexeyev and others 1995).

Because of their extent, and the large carbon reserves they contain, boreal forests

have long been recognized as critical to the earth’s carbon balance (e.g., Bonan

1991, Crutzen and others 1979, Levine 1991). Estimates vary, but suggest carbon

storage in boreal forest worldwide in the range of 66-127 Pg C in plant material,

and 135-247 Pg C in soils and downed woody material and litter (Apps and others

1993, Bolin 1986, Kauppi and others 1992, Olson and others 1993, Tans and others

1990.) This represents perhaps 10 to 17 percent of the total global terrestrial carbon

reserves in soils and plant materials (2,100-2,200 Pg C; Bolin 1986). A recent

comprehensive analysis of carbon in Russian ecosystems (Alexeyev and Birdsey

1994) estimates that the boreal forests in Russia contain 99.8 Pg C, with 25 percent

of this in vegetation and the remainder in soils and forest floor material.

This paper discusses the potential effects on carbon cycling, global climate,

and air quality of fires in the boreal forests of Russia, which comprise about twothirds of the world’s boreal forest.

Extent of Fire in Russia’s Boreal Forests

Fire is one of nature’s primary carbon-cycling mechanisms. It is a critical disturbance

factor in Russia’s boreal forests, where many fire scars are visible across the landscape

and vegetation patterns are often dominated by evidence of past fire history. During

the summer fire season, one can hardly fly across central Siberia without observing

smoke plumes from the often numerous wildfires. Although data are somewhat

Fire in Siberian Boreal Forests — Implications for Global

Climate and Air Quality1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Formerly Project Leader, USDA Forest

Service PSW Research Station, Forest

Fire Laboratory, 4955 Canyon Crest

Drive, Riverside, CA 92507, and now

Program Leader, Forest Vegetation

Management and Protection Research,

USDA Forest Service, P.O. Box 96090,

Washington, D.C. 20090-6090.

3 Chair, Department of Science and Technology, Russian Aerial Forest Protection Service (Avialesookhrana), 20

Gorkova Street, Pushkino, Moscow

District 141200, Russia.

Susan G. Conard2 and Eduard P. Davidenko3
88 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

sparse, dendrochronological studies have estimated typical fire return intervals of

25 to 50 years for Pinus sylvestris forest types in central Siberia (Furyaev 1996,

Swetnam 1996). Longer fire return intervals (90-130 years) appear more typical for

forests dominated by Larix species (Furyaev 1996, Valendik [In press]) and by Abies

siberica and Picea obovata. Fire return intervals also decrease from north to south

(they may be as short as 10 years on some sites), and are longer on the most wellwatered sites along rivers or in areas where forest stands are isolated by wet bogs

(FIRESCAN 1997). As a reflection of the difference in dominance and in fire return

intervals of the various forest types, about 70 percent of the fires in Russia occur in

light coniferous pine and larch-dominated stands (Pinus sylvestris or Larix), and

only about 15 percent in dark coniferous boreal forest (primarily Abies, Picea; based

on Korovin 1996), although the latter cover about 31 percent of the boreal forest

area (Tchebakova and others 1994). About 80 percent of the area typically burns in

surface fires (Furyaev 1996, Korovin 1996), but patchy crown fires are common, and in

severe fire seasons, crown fires may dominate. The Russian Forest Service (Nyejluktov

1994) reports a range of 12,000 to 34,000 wildfires a year in Russia during the period

from 1974-1993, with a long-term average of about 17,000 fires per year. Most of these

fires occur in a 4-month fire season that extends from May through August (fig.1).

Estimates of the extent of fires in Russia’s boreal forest vary over an order of

magnitude (table 1); those made by Russian government agencies are among of

the smallest. Some of the variation in these estimates undoubtedly results from

differences in areas covered. The official Russian fire statistics cover only areas

that receive fire protection. The fire protection area includes most of European

Russia, but is more limited in Siberia and the Far East. Overall, about 66 percent of

the forested area of Russia was monitored for fires in the early 1990’s (Dixon and

Krankina 1993), most of this by airborne pilot-observers. With recent budget

decreases, this area is now considerably less. While the monitored area in the past

included most of the closed boreal forest, some boreal forest fires, especially in the

more northern regions, were not reported. Other estimates have been based on

extrapolations from North American fire regimes (Kasischke and others 1993, Stocks

1991). On the basis of satellite data for three areas of Siberia in 1992, Stocks and

others (1996a) estimated close to 1 million acres had burned in the study areas,

and suggested this might represent 30 to 40 percent of the area burned in Siberia

that year. This would lead to an estimate of 2.5 to 3.3 million acres of burned area

for all of Siberia in 1992. On the other hand, analysis of satellite data at the time of

the 1987 China fires led to an estimate of about 14 million ha burned in southern

Siberia and China (mostly in Siberia) during May and June of 1987 (Cahoon and

others 1994). Emissions from this amount of burned area were estimated to

represent about 4 percent of the annual global emissions from biomass burning.

Although progress is being made in analysis of current satellite data, it is likely to

Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko

Figure 1 — Monthly distribution

of fire frequency during the fire

season in Russia. Source:

Unpublished fire statistics on file

at Russian Aerial Forest

Protection Service, 20 Gorkova

Street, Pushkino, Moscow Region

141200, Russia.

APR MAY JUN JUL AUG SEP OCT

Number of Files

160

140

120

100

80

60

40

20

0
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 89

be a number of years before adequate information will be available from satellite

data to allow better long-term estimates. At this point the uncertainty in the area

burned seems one of the greatest barriers to accurate estimates of the effects of

boreal forest fires in Russia on global biogeochemistry.

In the light of this wide range in estimates of fire area, we decided to take

another approach to obtaining a reasonable independent estimate of area burned

in Russian boreal forests based on the relationship between fire-return interval

and area burned per year. By using a relatively conservative estimate of 500 million

ha of closed boreal forest in Russia, we can predict (fig. 2) the average burned area

per year for different fire return intervals. For a fire return interval of 60-80 years,

which seems reasonable based on the literature, we would estimate between 6.25

and 8.3 million ha of burned area per year. This is similar to a recent estimate of 7.4

million ha/yr for the 1988-1991 period by Dixon and Krankina (1993), but

considerably higher than their longer-term estimate of 2.9 million ha/yr (table 1),

which would imply a fire return interval of 172 years (267 years if you use their estimate

of 774 million ha of total forest area). Some of the lower estimates of burned areas

(table 1) would yield fire return intervals in excess of 1000 years. This has clearly not

been the pattern over the last several hundred years, or in the 1900’s, as revealed by

landscape patterns and dendrochronology data (FIRESCAN 1994, 1997).

We recognize the possibility of some error in this approach, as it does not

consider possible recent impacts on fire regime of fire suppression activities and

changes in ignition patterns because of human activities. Although evidence from

Ontario, Canada, suggests an 89 percent decrease in area burned annually (increase

in fire return interval from 65 to 580 years) in areas under fire suppression (Ward

and Tithecott 1993), we find no compelling evidence for a similar trend in Russia

Table 1 — Average area (ha) of boreal forest burned annually in Russia and worldwide. Table 1

 Russia Worldwide Period Source

—— 1.3 x 106 —— Crutzen and others 1979

3.0 x 106 5.6 x 106 1980-1989 Stocks 1991

4.5 x 106 8.5 x 106 1940-1991 Kasischke and others 1993

1.9 x 106 —— 1971-1986 Dixon and Krankina 1993

7.4 x 106 —— 1987-1991 Dixon and Krankina 1993

2.9 x 106 —— 1971-1991 Dixon and Krankina 1993

0.3 x 106 —— 1979-1988 Nyejlukto 1994

0.8 x 106 —— 1989-1994 Nyejlukto 1994

Figure 2 — Relationship

between annual area burned in

forest fires and the average fire

return interval. This graph uses a

conservative estimate of 500 ha

of closed boreal forest in Russia

and assumes no variability in fire

return intervals.

Time Betwen Fires (years)

0 20 40 60 80 100 120 140 160

Area Burned (million ha/yr)

30

25

20

15

10

5

0

Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko
90 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

(Antonovski and others 1992, Korovin 1996, FIRESCAN 1997). Many fires in Russia,

especially large fires and those in remote areas, are little affected by fire suppression.

In fact, Russian Forest Service statistics suggest a possible trend toward increasing

areas burned over the past 10-15 years (Nyejlukto 1994), which may be associated

with increased human ignitions. Despite the great interannual variability of area

burned and the unreliability of fire size data for Siberia, it would certainly be

possible to detect a major trend of decreased burned areas or increased fire return

intervals. In the absence of such evidence, we believe that our estimate is a

reasonable one until more accurate ones are available.

Emissions from Boreal Forest Fires

Obviously, wide variations in estimates of area burned will lead to similar variations

in estimates of the contribution of emissions from boreal forest fires to the global

carbon balance, and this is reflected in the literature. We can assume, for example,

that the estimates by Crutzen and others (1979) of 40 to 60 Tg dry matter (18-27 Tg

C) consumed per year are probably about an order of magnitude too low, as they

are based on an estimate of 1.3 million ha burned annually worldwide (including

North America and Eurasia). A better combined global estimate might be about 10

million ha per year (0.2 million ha per year for Alaska and Scandinavia, and 2.4

million ha per year for Canada [Stocks 1991]; 7.3 million ha per year for Russia (our

estimate)). On the basis of various sources of Russian fire statistics for the period

1971 to 1991 (see table 1), Dixon and Krankina (1993) estimated an average of about

47 Tg C emissions per year after fires. If we adjust this upward based on our estimate

of an average burned area of 7.3 million ha per year, we can project annual average

C emissions from fire in Russia of about 118 Tg C per year, and 292 to 702 Tg C/yr

from biogenic sources. Adding in a rough estimate for North American forests would

increase these values by about 30 percent (to about 153 Tg carbon/yr from fire

emissions, and about 380 to 913 Tg C/yr from postfire biogenic emissions). This

direct amount from fire emissions represents about 4 percent of the estimated 3,900

Tg C/yr released globally from biomass burning (Andreae 1991). If our estimates

are reasonable (and there is every reason to believe they are conservative), then

annual contributions of boreal fire to global carbon release may have been vastly

underestimated by many authors.

Even if we have a good estimate of the annual extent of boreal forest fires in

Russia, extrapolation to impacts on emissions or on carbon cycling is complicated

by the wide variability in fire intensity across the landscape. Perhaps only 20 percent

of the burned area in Russia burns in crown fires (Furyaev 1996, Korovin 1996). In

the other 80 percent of the area, low levels of combustion of needles and other fine

aerial fuels will result in lower emissions of carbon compounds in smoke per unit

area. On the basis of data on a Pinus sylvestris/lichen forest in central Siberia

(FIRESCAN 1997) we can estimate that fuel consumption in a surface fire in this

vegetation would be about 80 percent of that in a crown fire (assuming 4.059 kg/m2

consumed in crown fire; 3.252 kg/m2 consumed in a surface fire). So estimates of

emissions based on assumption of crown fires will be about 20 percent too high.

Dixon and Krankina (1993) recognized this concern and assumed that an average

of 10 percent of the biomass was consumed in boreal forest fires in Russia. In other

studies it is not always clear how this issue was dealt with.

An even greater potential source of error is the amount of combustion of the

litter/forest floor and soil layers, which contain about 75 percent of the carbon

stored in Russia’s boreal forests (Alexeyev and Birdsey 1994). Depending on the

seasonality and intensity of fire and on the vegetation type, combustion of these

layers in North American boreal forests can range from about 16 to 90 percent,

with an average of perhaps 40 to 60 percent (Kasischke and others 1995).

Furthermore, fire characteristics also have generally poorly understood, but

substantial, effects on the magnitude of postfire increases in biogenic emissions,

Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 91

which have been estimated to be 70 to 85 percent of the total fire-related emissions

(Dixon and Krankina 1993). Still another source of error is that tree mortality in

areas of surface fire can vary widely, and may occur over several years, affecting

photosynthesis and respiration, as well as temporal and spatial patterns of postfire

decomposition and biogenic emissions. Accurate estimates of the extent and

severity of fire in boreal Russia and their effects on combustion patterns and

biogenic emissions are desperately needed.

Fire and Global Climate Change

Several authors suggest that, for boreal forest areas in North America, there will be

increased fire weather severity (Flanigan and Van Wagner 1991, Stocks 1993),

increased length of fire season (Street 1989, Wotton and Flannigan 1993), and

increased lightning (Fosberg and others 1990) with CO2-induced global climate

change. Changes in temperature are expected to be considerably higher in boreal

regions than the global averages. Stocks (1993) discussed the possible impacts of a

doubled CO2 scenario on fire danger for Canada, based on predictions of the

Canadian Climate Center global climate model and the Canadian fire danger rating

system. This model predicts a 4-5 °C increase in summer temperatures and a 8-10 °C

increase in winter temperatures, along with decreased soil moisture availability

throughout central Canada. Wotton and Flannigan (1993) have predicted an increase

in the length of the fire season by 30 days, and Flannigan and Van Wagner (1991)

predicted a 46 percent increase in seasonal fire severity across Canada under a double

CO2 scenario. Stocks and Lynham (1996) used the Monthly Severity Rating from

the Canadian Forest Fire Danger Rating System and the Russian Nesterov Index to

compare fire weather severity between Canada and Russia. Both of these indices

represent the fire danger if an ignition occurs, based on weather patterns and fuel

moisture conditions. Their results suggest that Russia may be even more at risk of

increased fire activity in response to climate change than Canada, as Siberia currently

has about three times as much area in extreme fire danger categories as Canada

(based on Monthly Severity Rating for July). Increases in fire occurrence can also be

expected to cause positive feedbacks with atmospheric CO2, thereby exacerbating

effects of global warming (Kurz and others 1994).

Pollution

Direct impacts of fire on air pollution are also cause for concern. It is not uncommon

in severe fire seasons for multiple fires to burn over long periods, casting a pall of

smoke over large areas. This is by no means a recent problem. Shostakovich (1925)

reported on extensive fires in Siberia in 1915. These fires burned for 50 days, and

burned about 14 million ha, but the smoke pall from the fires covered 680 million

ha, and severely reduced visibility at ground level over a large percent of the area.

Extensive smoke plumes were visible on Advanced Very High Resolution

Radiometer (AVHRR) imagery in 1987, during the period of the great China fires

(Cahoon and others 1994). Given the large number of fires that burn in Russia every

summer (an average of perhaps 140 starts per day over the 4-month fire season,

and many more at the peak of the season), and the large areas burned, it is not

surprising that smoke intrusion over extensive areas occurs.

In addition to the obvious effects of such extensive smoke on visibility and

other aesthetic values, prolonged wildfire smoke episodes in western North

America in recent years (particularly northern California in 1987, Yellowstone in

1988) have raised concerns over potential negative impacts of wildfire smoke

exposure on human health, as well as short-term toxicity from elevated carbon

monoxide levels (Reinhardt and others 1995). The main components of concern

for human health in wildfire smoke include carbon monoxide, aldehydes, benzene,

and particulate matter (Reinhardt and others 1994). Carbon monoxide has been

Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko
92 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

recognized as a potential hazard to wildland firefighters for a number of years

(Jackson and Tietz 1979), but specific information is scant. Studies on prescribed

fires in the Pacific Northwest have observed smoke levels exceeding established

exposure limits (Reinhardt and others 1994). In a recent study on levels of smoke

exposure for firefighters on wildfires in northern California grass and light brush

fuels (Reinhardt and others 1995), however, peak exposures never reached

recommended ceiling limits of CO of 200 ppm, and only 2 of 23 peak exposures to

respiratory irritants exceeded exposure limits. Reinhardt and others (1995) point

out that these fires were all in relatively light fuels, and experienced firefighters

involved in the incidents rated the smoke levels as low to average. Furthermore,

exposures in these fires were typically of short duration. Other research (Materna

and others 1992) has found high CO exposure among engine crews on wildland

fires in California. Reinhardt and others (1995) concluded that extended exposure

to forest fire smoke could exceed exposure guidelines for both CO and respiratory

irritants, and recommended that more extensive studies be conducted under a

wide range of fuel and burning conditions. A recent review (Reinhardt and Ottmar

1997) of published and unpublished research on exposure of wildland firefighters

to smoke and potential health hazards reported mostly on studies conducted in

the western U.S. The authors stated that few data on smoke exposure of wildland

firefighters are available from other parts of the world. Considering the periodic

exposure of rural and urban populations in Siberia to wildfire smoke over

prolonged periods, further research on firefighter smoke exposure and on the effects

of long-duration smoke exposure from wildfires on human health is vitally needed.

Human Influences on Fire Regimes

Statistics over a recent 3-year period for Russia show that more than 65 percent of

wildfires are human-caused, and about 17 percent are confirmed to be caused by

lightning (fig. 3). Newer information, however, suggests that the importance of lightning

ignitions may have been underestimated. Of the 17 percent of fires where cause is

unknown, evidence from recently-installed lightning detection systems suggests that

many of these may be lightning fires. And the importance of lightning as an ignition

source varies greatly geographically (table 2), with the highest percent of lightning

ignitions in relatively unpopulated areas such as the Krasnoyarsk and Yakutia regions.

Nonetheless, even in these regions, a large percentage of fires is human-caused.

Furthermore, it appears possible, based on tendencies visible in the fire statistics

of the Russian Forest Service (Nyejlukto 1994) and those reported by Korovin (1996)

that there has been an increase in fire numbers and area burned in the past 10

years. After some decrease in the number of fires and areas burned between the

early 1970’s and about 1983, the trend shows a gradual increase through 1992-93.

Average burned area for 5-year periods reported by the Russian Forest Service

(Nyejlukto 1994) increased from 0.23 million ha in the 1979-1983 period to 1.03

million ha in 1989-1993, although levels still are not as high as they were in the

early 1950’s (as high as 2.5 million ha in 1953 [Korovin 1996]). The large influence

of human populations on fire occurrence is well-illustrated by Korovin (1996),

who shows that about 68 percent of fires occur within 5 km of a road, and 60

percent of fires occur within 10 km of a populated area. Human-caused ignitions

may be having a short-term net effect of increased fire in many parts of Russia,

despite a strong fire suppression program. Changes in area burned, whether

resulting from ignition patterns, weather cycles, or fire management strategies,

have particularly important implications for global climate change, as alterations

in fire return interval can have significant impacts on carbon storage (which

increases with stand age). Model estimates for stored carbon in boreal forest

(Kasischke and others 1995) show strong sensitivity to fire return interval in the

range of 50 to 150 years, with projected increases in carbon storage of about 10 to

17 percent as fire return interval increases from 50 to 100 years, and increases of

about 25 to 40 percent as fire return interval increases from 50 to 150 years. This

Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 93

sensitivity in the range of current fire return intervals suggests the great potential

for human impacts on the fire regime to significantly affect carbon storage, as well

as the importance of fire management in managing fire regimes and carbon cycling.

An active fire suppression program will be a key element in maintaining desirable

fire return intervals in the face of increased human ignitions.

Conclusions

Fire is an important factor in boreal forests of Russia. Despite the paucity of accurate

data, it is clear that boreal forest fires in Russia contribute significantly to global

CO2 emissions. Increased CO2 is likely to substantially increase fire danger and

occurrence in boreal forests, leading to possibilities of positive feedbacks to global

CO2 and temperatures (Kurz and others 1994) and significant ecosystem impacts

if fire suppression and other fire protection and prevention approaches are not

able to compensate. Extensive fires can cause smoke pollution over large areas,

which may result in hazards to human health and other negative impacts. The

large percent of human-caused fires in Russia’s boreal ecosystems suggests the

critical importance of fire management and fire suppression for maintaining

desirable fire regimes and preventing decreases in fire return interval and loss of

carbon storage in these ecosystems.

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Table 2 Table 2 — Percent of fires caused by lightning in Russia’s major aerial forest protection regions over the 3-year period Table 2 

1993-1995.1

Protection Region 1993 1994 1995 Average

——————Percent of total fires————-

Krasnoyarsk 42 35 43 40

Irkutsk 16 12 23 17

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1 Source: Unpublished fire statistics on file at Russian Aerial Forest Protection Service, 20 Gorkova Street, Pushkino,

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Session I1 Fire in Siberian Boreal Forests – Implications for Global Climate and Air Quality Conard, Davidenko
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 95

Abstract

Preliminary results of studies on nutrient fluxes in forests of the eastern Sierra Nevada were compared to those from

more humid and polluted ecosystems. Snowmelt, soil solution, soil, and streamwater were collected from Jeffrey and

lodgepole pine (Pinus jeffreyii [Grev. and Balf.] and Pinus contorta Dougl.) stands in Little Valley, Nevada, and

from California red fir (Abies magnifica A. murr.) and Jeffrey pine/white fir (Pinus jeffreyii/Abies concolor [Gord

and Glend.] Lindl.) stands at Sagehen, California. Snowmelt, soil solutions, and streamwaters from both sites were

circumneutral and dominated by base cations and bicarbonate. The red fir stand at Sagehen had high NO3

-

concentrations (approximately 30-100 μmol /L) in both snowmelt and soil solution during the relatively dry 1993-4

water year. The Little Valley sites had substantially lower NO3

-

 concentrations in both snowmelt (5-20 μmol /L) and

soil solution (0.5-3 μmol /L) in both wet and dry years. At both sites, a pulse of streamwater NO3

-

 (from 0.5 to 20-40

μmol /L) was detected during dry years but not wet years. The Andic soils at the Sagehen site have trace levels of

available P in soils, whereas the Entisols and Inceptisols in Little Valley have 10 to 100 times greater levels. The results

suggest a hypothesis that the greater mobility of NO3

-

 in the Sagehen red fir site was caused by the amount and timing

of NO3

-

 release from snowmelt (e.g., too concentrated and early for dormant biota to respond to), and that P may also

be limiting at Sagehen. Compared to other forest ecosystems in more humid and polluted environments, N, S and H+

deposition and fluxes at the Little Valley site are extremely low, but HCO3

- and base cation fluxes are high. This reveals

that the Little Valley site is relatively pristine, and that ionic fluxes at this site are dominated by natural carbonic acid

leaching and weathering reactions.

Introduction

In comparison to more humid regions, information about nutrient cycling in forests

of arid and semi-arid regions is scant. Some nutrient cycling work has been done

on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) (Hart and Firestone 1989,

Klemmedson 1975) and lodgepole pine (Pinus contorta Dougl.) ecosystems (Fahey

1983, Fahey and Knight 1986, Schimel and Firestone 1989), but these studies are

relatively fewer than those in more humid regions (Cole and Rapp 1981, Johnson

and Lindberg 1991). Except for Stark’s (1973) study on Jeffrey pine (Pinus jeffreyi

Grev. and Balf.), little is known about the cycling of nutrients in forests of the eastern

Sierra Nevada.

Scientific knowledge of nutrient cycles in forests of the eastern Sierra Nevada

should be increased for several reasons. First, this knowledge would allow

intelligent assessments of forest management practices such as harvesting, burning,

and site preparation as well as the effects of exogenous influences such as fire, air

pollution, and climate change on these forests. Second, these forests act as filters

for nutrients, especially nitrogen, which might otherwise enter the sensitive surface

waters of the region. The long-term deterioration of water suality of Lake Tahoe

has been clearly documented (Goldman and others 1983). This deterioration is

thought to be caused by increasing nutrient inputs from land development,

atmospheric deposition, and N fixation by riparian mountain alder (Alnus tenuifolia

Nutt.) (Byron and Goldman 1989, Coats and others 1976, Leonard and others 1980).

With N and H+ inputs increasing in the Tahoe Basin (Byron and others 1991), it is

important to gain some knowledge of the capacity of forest ecosystems in the basin

to filter these inputs. Galloway and others (1995) forecast that anthropogenic N

fixation (by a combination of energy production, fertilizer production, and

cultivation of N-fixing crops) will increase by about 60 percent by the year 2020.

They predict that about two-thirds of this increase will take place in Asia, which

Nutrient Fluxes in Forests of the Eastern Sierra Nevada:

Comparisons with Humid Forest Systems1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Forest Soils and Graduate

Student, respectively, Biological Sciences Center, Desert Research Institute,

PO Box 60220, Reno, NV and Environmental and Resource Sciences, University of Nevada, Reno, NV 89512.

3 Professor of Soils, Land, Air, and Water Resources, University of California,

Davis, Davis, CA 95616.

4 Professional Researcher, Marine Sciences Institute, University of California, Santa Barbara, PO Box 605 Los

Olivos, CA 93411.

5 Ecologist, Pacific Southwest Research

Station, USDA Forest Service, 4955

Canyon Crest Drive, Riverside, CA

92507.

Dale W. Johnson,2 Richard B. Susfalk,2 Randy A. Dahlgren,3 Virgina Boucher,4 and

Andrzej Bytnerowicz5
96 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

by 2020 will account for over half of the global anthropogenic N fixation. Thus,

there is the potential for increased N deposition in western North America from

both local and long-range sources. At the same time, the capacity of forests to

retain N has been reduced because of increased insect- and drought-induced

mortality and from fires.

This paper reports progress to date on studies of nutrient cycling in forests of

the eastern Sierra Nevada, and compares these results with those from a variety of

forest ecosystems in more humid climates by using the Integrated Forest Study

database on forest nutrient cycling (Johnson and Lindberg 1991).

Sites and Methods

The Little Valley site is located about 30 km southwest of Reno, Nevada in the

eastern Sierra Nevada (fig. 1). It ranges from 2,010 to 2,380 m elevation. Vegetation

is dominated by lodgepole pine in riparian areas, Jeffrey pine at lower elevations

and white fir (Abies concolor [Gord and Glend.] Lindl.) at higher elevations. Mean

annual temperature (MAT) near the valley floor is 5 °C and mean annual

precipitation (MAP) is 550 mm, about 50 percent of which falls as snow. Site 1 in the

northern end of the Valley is located on the Corbett series soil, a typic frigid

Xeropsamment derived from outwash of decomposed granite. Overstory vegetation

consists of Jeffrey pine with an occasional white fir. Understory vegetation is absent.

Site 2 is located about 5 km south of site 1 on Marta series soil, a sandy, mixed

Aquic Cryumbrept derived from colluvium of decomposed granite. Overstory

vegetation in site 2 consists of lodgepole pine with occasional Jeffrey pine.

Understory vegetation consists of sage (Artemesia tridentata Nutt.) and various

grasses and forbs.

The Sagehen Creek watershed is located about 10 km north of Truckee,

California (fig. 1). Elevation ranges from 1,830 to 2,500 m elevation. MAP and MAT

at the Sagehen field station (elevation = 1,830 m) are 870 mm and 4.8 °C, respectively.

More than half of total precipitation falls as snow. Site 1 is located on Waca series

soil, a loamy skeletal, mixed, frigid Andic Xerochrept derived from andesitic lahar

and tuff. Overstory vegetation consists of 80-160-year-old California red fir (Abies

magnifica A. murr.) with occasional white fir and an understory of pinemat

manzanita (Arctostaphylos nevadensis Gray) with Ribes and Wyethia spp. Elevation

at the site is 2,100 m. Site 2 is located on Fugawee series soil, a fine-loamy, mixed

frigid Ultic Haploxeralf derived from andesitic lahar and tuff. Overstory vegetation

consists of 80-160-year-old Jeffrey pine and white fir. Understory consists primarily

of occasional Ribes and squaw carpet (Ceanothus prostratus Benth.).

Over a 6-year period (1989 - 1995), we collected snowmelt, soil solutions, and

streamwater from various sites. Because of a combination of fiscal and logistical

limitations, we could not collect solutions from all sites simultaneously. Thus,

solutions were collected from Site 1 in Little Valley during the relatively droughty

1989-90 water year, from Site 1 at Sagehen site during the relatively droughty 1993-

94 water year, and from Site 2 at Little Valley during the relatively wet 1994-95

water year. We attempted to collect from Site 1 at Sagehen during 1994-95, also,

but the deep snowpack (as much as 5 m) there during that year precluded consistent

water collections.

Snowmelt collectors and tube-type tension lysimeters were located in six

randomly located positions within a 0.05-ha plot at each site. Snowmelt collectors

consist of buried collection bottles to which the open collectors are attached. The

open collectors sit on the ground surface partially buried and consist of the cut-off

tops of polypropylene bottles fitted with a drainage tube that is attached to the

buried collection bottle. These devices collect rain or throughfall (depending upon

their location) during the snow-free seasons and snowmelt during the snow season.

The collection bottle is fitted with tubing that is run-up poles so that collections of

snowmelt solutions are possible from beneath snowpack without disturbance.

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 97

Tube-type lysimeters are fitted with an extra tube which, along with a vent tube,

allows removal of soil solutions from under snowpack without disturbance

(Johnson 1995). All solutions were analyzed for conductivity, pH and alkalinity

(titration to pH 5.0), cations (Ca2+, Mg2+, K+, Na+, NH4

+) by Dionex ion

chromatography,6 anions (Cl-, NO3

-

, ortho phosphate, SO4

2-) by Dionex ion

chromatography, and Si by ICP spectroscopy.

We used the chloride balance method (Lindberg and Johnson 1989) to estimate

nutrient fluxes at Little Valley. (Because of an insufficient number of reliable

snowmelt water volume collections at the other sites, these calculations were not

made). Chloride fluxes were measured directly from water volumes and concentrations

in the snowmelt collectors beneath the forest canopy. We used these rather than the

open collectors in order to better incorporate dry deposition of Cl- into the calculations.

It was then assumed that Cl- fluxes through the soil were equal to amounts from

snowmelt (i.e., that Cl- was a conservative tracer), and soil water fluxes were calculated

from this. These water fluxes were then used with soil solution concentrations to

estimate soil leaching rates for the various ions.

Figure 1 — Research sites were

located at Little Valley, Nevada,

and Sagehen, California.

6 Mention of trade names or products is

for information only and does not

imply endorsement by the U.S.

Department of Agriculture.

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz

California

Nevada

SAN FRANCISCO

STOCKTON

SACRAMENTO

LOS ANGELES

RENO

CARSON CITY

SPARKS

80

80

50

LAS VEGAS

uLittle Valley Sagehen u
98 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Results and Discussion

In previous studies in Little Valley (Site 1), we followed up the studies of Coats and

others (1976) in the Tahoe Basin and investigated the effects of N fixation by both

mountain alder (Alnus tenuifolia Nutt.) and snowbush (Ceanothus velutinus Dougl.)

on soils and soil solutions in comparison to Jeffrey pine. (We know of no previous

studies have been conducted on soil solution chemistry beneath snowbush.) In

contrast to the results of Coats and others (1976), we found no evidence of

excess NO3

-

 beneath either mountain alder or snowbush during the 1989-90

water year. Soil solution NO3

-

 concentrations were near trace levels under all

three species (table 1; Johnson 1995). We found a sizable NO3

-

 peak in streamwaters

from nearby Franktown Creek during snowmelt in 1990 (a dry year) (fig. 2) but not

in 1995 (a wet year). Sulfate, HCO3

-

, total cation and total anion concentrations in

Site 1 in Little Valley were quite high. Sulfate and Cl- concentrations in soil solutions

were two orders of magnitude greater than in nearby streamwaters (Johnson 1995).

More recent studies (1994-95 water year) in a lodgepole pine stand growing on more

weathered soil lower in the valley (Site 2) showed similarly low soil solution NO3

-

concentrations, but also much lower SO4

2-, Cl-

, total cation and anion concentrations

than in the original site. At this stage, we do not know the reasons for the differences

in soil solutions between these two sites, but possibilities include the age of the parent

material and the amount of water flux. The parent material in Site 1 consisted of

alluvium from nearby Ophir Creek and is noticeably less oxidized than that in Site 2.

Research is underway to determine the reasons for these differences.

Snowmelt and soil solutions from the red fir stand (Site 1) at Sagehen were

similar to those from Site 2 in Little Valley in terms of pH, HCO3

-

 , SO4

2- , total

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz

Table 1 Table 1 — Concentrations of selected ions in soil solutions and streams from Little Valley, Nevada, and Sagehen, Table 1 

California.1

pH HCO3

- NH4+ SO42- Cation Anions

μmolc

 L-1

Little Valley: Little Valley

Site 1 2

Soil solution

 Ceanothus 6.7 ± 0.1 306 ± 47 3 ± 3 107 ± 15 1107 ± 129 614 ± 113

 Pine 7.1 ± 0.1 417 ± 94 <0.5 273 ± 46 1098 ± 200 842 ± 75

 Mt. Alder 7.0 ± 0.2 688 ± 192 <0.5 181 ± 59 1084 ± 315 1017 ± 296

Stream 7.1 ± 0.2 373 ± 15 3 ± 2 3 ± 0.2 373 ± 12 396 ± 15

Site 2

Snowmelt 6.0 ± 0.3 102 ± 88 4 ± 6 15 ± 18 337 ± 203 143 ± 104

Soil solution

 15 cm 6.4 ± 0.4 260 ± 257 0.2 ± 0.6 10 ± 25 514 ± 328 292 ± 255

 30 cm 6.6 ± 0.4 281 ± 138 0.2 ± 0.5 7 ± 6 501 ± 109 328 ± 125

Stream 6.8 ± 0.4 389 ± 30 0.5 ± 1.0 11 ± 12 579 ± 77 419 ± 48

 Sagehen Creek: Sagehen Creek:

Site 1

Snowmelt 6.8 ± 0.2 125 ± 75 64 ± 60 13 ± 8 340 ± 120 231 ± 93

Soil solution

 15 cm 7.2 ± 0.7 300 ± 111 64 ± 48 19 ± 12 693 ± 297 408 ± 129

 30 cm 7.3 ± 0.3 407 ± 283 48 ± 33 20 ± 16 702 ± 329 505 ± 301

Stream 7.4 ± 0.4 977 ± 440 19 ± 21 2 ± 1 1280 ± 219 1006 ± 429

1 Standard errors are given.

2 Johnson (1995).
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 99

Figure 2 — Temporal patterns

in NO3

- concentration in

Franktown Creek, Little Valley,

Nevada.

Figure 3 — Temporal patterns

in NO3

- concentration in Sagehen

Creek, California.

Franktown Creek

8/17/899/12/899/27/89

10/13/8910/18/8910/27/89

11/8/89

1/9/90

3/21/903/30/904/13/904/26/905/15/906/20/90

8/8/90

8/29/90

12

10

8

6

4

2

0

12

10

8

6

4

2

–––––––––––––––

0

μmol L-1μmol L-1

Sagehen Creek

50

45

40

35

30

25

20

15

10

5

0 –

–––––––––––––––––––––––––––––––––––––––––––

μmol L-1μmol L-1

10/6/93

10/25/9311/21/93

12/6/93

12/20/93

1/3/94

1/17/941/31/942/14/942/28/943/25/943/30/94

4/9/94

4/13/944/28/94

5/5/94

6/17/94

7/6/94

7/25/948/11/94

10/11/9410/19/94

11/4/9412/2/94

12/26/94

1/16/95

2/1/95

2/24/953/17/954/14/95

5/2/945/8/95

5/14/95

6/1/956/6/95

6/13/956/16/956/26/95

7/3/95

7/10/957/24/957/31/95

8/7/95

8/22/95

–

-

-

-

-

–

-

-

-

-

–

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

50

45

40

35

30

25

20

15

10

5

0

–

-

-

-

-

–

-

-

-

-

–

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz

cations and total anions, but were considerably higher in both NO3

-

 and NH4

+ (table 1).

(Snowmelt and soil solution data from Site 2 at Sagehen are being collected in the

1995-96 season and are not yet available.) Sagehen Creek showed a peak in NO3

-

during snowmelt in a drought year (1993-94) but not in a wet year (1994-95) (fig.

3). At both Little Valley Site 2 and Sagehen Site 1, snowmelt NO3

-

 peaked during

the spring of 1994 and 1995 (fig. 4). But the two sites differed: the peak NO3

-

concentrations at the Sagehen site were an order of magnitude greater than at

Little Valley Site 2, and the pulse of NO3

-

 at Sagehen penetrated the soil whereas

this was not the case in Little Valley Site 2. The same patterns were apparent in

NH4

+ release from snowpack and mobility at these two sites (not shown).

Differences in snowpack amounts, snowmelt NO3

-

 and NH4

+ concentrations, the

timing of NO3

-

 and NH4

+ release from snowpack NH4+ , and site conditions could

contribute to these differences in NO3

-

 and NH4

+ mobility at the two sites. At

Sagehen Site 1, peak snowmelt NO3

-

 was much higher and occurred earlier (MarchMay) than at Little Valley Site 2 (May-June). Only in June, when snowmelt NO3

-

concentrations decreased and biological N uptake presumably commenced, did

soil solution NO3

-

 at Sagehen Site 1 drop to levels typical of N deficient ecosystems.

The same patterns were apparent for NH4

+ (not shown).

Currently, we are entertaining two alternative hypotheses to explain the

differences in N-

 mobility in the Little Valley and Sagehen sites. The first hypothesis

is that there is a temporal discoupling of N release from melting snowpack and
100 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

the activation of biological uptake in these systems. In Mediterranean ecosystems

like these, when soil moisture conditions are most favorable for biological activity

(winter/spring), the soil and air temperatures may be limiting; when soil and air

temperatures are most favorable (summer) for biological activity, soil moisture

conditions are limiting. Thus, only during brief periods (e.g., after the snow pack

melts and in the fall period before snow pack accumulates) is biological activity

not severely inhibited by temperature and moisture conditions. We hypothesize

that amount and timing of N release during snowmelt is critical to the mobility of

N in the system. If snowmelt N release is concentrated and early, as apparently

was the case at Sagehen in the droughty spring 1994, biota may not have broken

dormancy sufficiently to take up N as it passed by. If snowmelt N release is more

diluted and later, such as was the case in the wet spring 1995, there is a better

chance for biotic uptake of N. We are currently testing this hypothesis by making

simultaneous collections in the low-elevation pine-dominated stands (Site 2) at

both Little Valley and Sagehen, where we are comparing and contrasting N mobility

during wet and dry years.

The second hypothesis to explain the difference in N mobility in these two

watersheds relates to the soil parent material and its effects on P status. The available

P status of the Sagehen soils, which nearly fall into the Andisol order (known to

have high phosphate adsorption capacity), is low compared to the soils in Little

Valley. Soils from Site 1 in Little Valley have extractable levels that are an order of

magnitude greater than those typical of more humid region forest soils (Johnson

and Lindberg 1991) (table 2). In the more weathered and oxidized soils in Site 2 in

Figure 4 — Snowmelt and soil

solution NO3

-

 in Sagehen Site 1

(top) and Little Valley Site 2

(bottom).

Sagehen Red Fir Site

Dec 16.93

Jan 12, 94

Mar 7, 94

Mar 16, 94Mar 31, 94

Apr 11, 94Apr 21, 94

May 9, 94

Jun 3. 94

250

200

150

100

50

0

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–_

_________

Snowmelt

Soil Solution 15cm

Soil Solution 30cm

Little Valley Lodgepole Pine Site

20

15

10

5

0

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–_

________

Snowmelt

Soil Solution 15cm

Soil Solution 30cm

Mar 15, 95

Apr 25, 95

May 3, 95

May 16, 95May 30, 95

Jun 21, 95

Jul 17, 95Jul 27, 95

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 101

Little Valley, available P levels are in the range typically encountered in more humid

forest soils. In contrast, extractable P in the Sagehen soils from two different sites

are below detection limits (table 2). We hypothesize that high P fixation capacity

on non-crystalline materials (e.g., ferrihydrite, allophane, imogolite, Al and Fe

humus complexes) in the Andic soils at Sagehen has caused a potential P limitation.

In contrast, we hypothesize that P levels in Little Valley Entisols and Inceptisols

are high because of low contents of non crystalline materials and low P fixation

capacity. We hypothesize that P limitation in the Sagehen sites limits the capability

of these forests to take up N released during snowmelt. We are currently testing

this hypothesis in part by performing single-tree N and P fertilization studies at

both Little Valley and Sagehen. These fertilization trials and their associated vector

analysis of foliage response have proven to be extremely reliable indicators of

potential nutrient limitation in other coniferous forests (Timmer and Stone 1978).

We do not believe that the differences in N in the two sites are related to air

pollution inputs. Chemical analysis of air (Bytnerowicz and Riechers 1995) was

performed in August 1994 at the Sagehen and Little Valley sites (table 3).

Concentrations of ammonia (NH3), nitric acid vapor (HNO3), and particulate nitrate

(NO3

-

) were about three times higher at the Little Valley site compared with the

Sagehen site. The concentrations at the Little Valley site were similar to the values

at the mixed coniferous forest locations near Kings Canyon, western Sierra Nevada

(Bytnerowicz and Riechers 1995). HNO3

 concentration at Sagehen was about two

times higher than the average summer concentration at the clean site of the eastern

Sierra Nevada (Miller and Walsh 1991).

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz

1 Standard errors are given. Nitrogenous compounds were measured with annular denuder system (Bytnerowicz and

Reichers 1995).

Table 3 Table 3 — Concentrations of selected nitrogenous air pollutants at two Sierra Nevada sites ( Table 3 μg m-3).1

 Location NH3 HNO3 NO3

-

particulate

Sagehen, California 0.347 0.636 0.149

(0.044) (0.060) (0.060)

Little Valley, Nevada 1.142 2.019 0.506

(0.099) (0.212) (0.039)

1 Standard errors are given.

2 Johnson (1995)

Table 2 — Table 2 — Soil extractable P (NH Table 2 — 4 F/HCL) in Little Valley, Nevada, and Sagehen, California soils.1

 Little Valley Little Valley 2 Sagehen

 Horizon and extr. P Horizon and extr. P

 depth (cm) depth (cm) (μg g-1) depth (cm) depth (cm) (μg g-1)

Site 1 Site 1

A (0-20) 101 ± 21 A (0-10) <0.5

BA (20-40) 106 ± 36 B (10-18) <0.5

BC (40-60) 69 ± 12 BC (18-45) <0.5

Site 2 Site 2

A (0-5) 10 ± 3 A (0-10) 0.3 ± 0.3

BA (5-20) 8 ± 3 B (10-18) 0.1 ± 0.1

BC (20-50) 4 ± 2 BC (18-45) 0.1 ± 0.1
102 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Comparison of Little Valley to Other Forest

Ecosystems

Deposition and soil leaching fluxes of N, S, H+, and Ca2+, Mg2+, and HCO3

-

 in Site 2

at Little Valley were compared to fluxes in a variety of forest ecosystems from the

Integrated Forest Study (IFS) (figs. 5, 6). (Because of logistical problems and funding

limitations, we have not been able to collect sufficient data to calculate fluxes for

Site 1 in Little Valley or the Sagehen site as yet.)

Although the pulses of NO3

-

 in the Little Valley and Sagehen sites are of concern

in terms of water quality in the sensitive surface waters of this region, comparisons

with the IFS sites clearly reveal that the Little Valley (LV) site is relatively pristine.

The fluxes of H+, N, and S in LV are lower than those in the other systems, including

the relatively unpolluted systems in the Pacific Northwest (DF and RA). To some

extent, this comparison may reflect the lack of inclusion of dry deposition to the

LV site, since deposition fluxes include only ionic fluxes via rain and snowmelt.

However, it seems quite clear that deposition at LV will remain at the very low

end of the scale even if annual dry deposition increases inputs estimates by severalfold. If flux data were available for Sagehen Site 1, the N outputs would be

substantially greater, judging by the tenfold higher soil solution NO3

-

concentrations.

Figure 5 — Atmospheric

deposition and leaching of H+, S,

and N in the Integrated Forest

Study sites (Johnson and

Lindberg 1991) and Site 2 in Little

Valley, Nevada (LV). Stands: CP

(Pinus strobus, Coweeta, NC); DL

(Pinus taeda stand at Duke, NC);

GS (Pinus taeda, B.F. Grant Forest,

GA); LP (Pinus taeda, Oak Ridge,

TN); FS (Pinus eliottii, Bradford

Forest, FL); DF (Pseudotsuga

menziesii, Thomspson, WA); RA

(Alnus rubra, Thompson, WA); NS

(Picea abies, Nordmoen, Norway);

HF (northern hardwood,

Huntington Forest, NY); MS

(Picea rubens, Howland, ME); WF

(Picea rubens, Whiteface, NY); ST

(Picea rubens, Clingman’s Dome,

NC); LV (Pinus contorta/P. jeffreyii,

Little Valley, NV). Deposition

fluxes at LV include bulk

precipitation during the snowfree season and snowmelt inputs.

Hydrogen Ion

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

2000

1500

1000

500

0

- 500

- 1000

- 1500

- 2000

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

––––––––––––

molc ha-1yr-1

Nitrogen

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

3000

2000

1000

0

- 1000

- 2000

- 3000

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

––––––––––––

molc ha-1yr-1

Sulfur

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

2500

2000

1500

1000

500

0

- 500

- 1000

- 1500

- 2000

-2500

–-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

-

–

––––––––––––

molc ha-1yr-1

Warmer Sites Colder Sites

Session I1 Nutrient Fluxes in Forests of the Eastern... Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 103

In contrast, the LV site had was in the high range for HCO3

-

 fluxes and in the

moderate to high range for Ca2+ and Mg2+ fluxes compared to the other IFS sites.

The high inputs no doubt reflect high inputs of dust to the LV site from nearby

desert systems, and the high outputs reflect the relatively high base saturation of

the soils, high weathering rates of primary minerals, and high rates of HCO3

-

leaching in these systems. The LV site seems to be experiencing net losses of Ca2+,

Mg2+, and K+ during the 1994-95 water year, but it should be noted that this was a

very wet year (>200 percent of normal snowpack), and leaching fluxes in drier

years are almost certainly lower. We will continue to monitor this site in both dry

and wet years to develop long-term average flux values.

Conclusions

Studies to date reveal a wide range in N and P status in the four sites investigated

in the eastern Sierra Nevada, with no clear patterns yet emerging. In one low-P

high-elevation red fir site at Sagehen, NO3

-

 released during snowmelt readily leached

through the soil profile, suggesting the possibility of a temporal disconnection

between snowmelt N release and biological uptake. In lower-elevation pine sites in

Little Valley, snowmelt NO3

-

 release was later and less concentrated, and did not

penetrate through the soil profile. Streamwater NO3

-

 pulses occurred during dry

years but not wet years in both sites, however, suggesting that snowmelt NO3

-

 shunts

Figure 6 — Atmospheric

deposition and leaching of HCO3

-

, K,

and Mg in the Integrated Forest

Study sites (Johnson and Lindberg

1991) and Site 2 in Little Valley,

Nevada (LV). See figure 5 for

legend. Deposition fluxes at LV

include bulk precipitation during

the snow-free season and

snowmelt inputs.

Bicarbonate

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

500

0

- 500

- 1000

 -1500

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

––––––––––––

molc ha-1yr-1

Calcium

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

2000

1500

1000

500

0

- 500

- 1000

-1500

- 2000

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

––––––––––––

molc ha-1yr-1

Magnesium

Deposition

Leaching

CP DL GS LP FS DF RA NS HF MS WF ST LV

600

400

200

0

- 200

- 400

- 600

- 800

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

––––––––––––

molc ha-1yr-1

Warmer Sites Colder Sites

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz
104 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

around the biological system in at least part of each watershed during dry years.

The differences in NO3

-

 mobility among these sites may also be related to soil parent

material. A large range in available soil P was evidenced among the different soil

parent materials, and the degree of weathering varied. The possibility of P deficiency

in the Andic soils at Sagehen is being explored.

Compared to other forest ecosystems in more humid and polluted

environments, N, S, and H+ deposition and soil leaching fluxes at the Little Valley

site are extremely low, but HCO3

- and base cation fluxes are high. These differences

reveal that the Little Valley site is relatively pristine, and that ionic fluxes at this

site are dominated by natural carbonic acid leaching and weathering reactions.

Acknowledgments

The research reported was supported by the Nevada Agricultural Experiment

Station, University of Nevada, Reno. Hal Klieforth provided field assistance and

Valerie Yturiaga provided laboratory assistance. We would also like to thank Laurie

Dunn for technical editing of this manuscript.

References

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and Soil Pollution 61: 139 168.

Byron, E.R.; Axler, R.P.; Goldman, D.R. 1991. Increased precipitation acidity in the central Sierra Nevada. Increased precipitation acidity in the central Sierra Nevada. Atmospheric Increased precipitation acidity in the central Sierra Nevada.

Environment 25A: 271-275.

Byron, E.R.; Goldman, C.R. 1989. Land-use and water quality in tributary streams of Lake Tahoe, California-Nevada. Journal Land-use and water quality in tributary streams of Lake Tahoe, California-Nevada

of Environmental Quality 18: 84-88

Bytnerowicz, A.; Riechers, G. 1995. Nitrogenous air pollutants in a mixed conifer stand of the western Sierra Nevada, Nitrogenous air pollutants in a mixed conifer stand of the western Sierra Nevada,

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Coats, R.N.; Leonard, R.L.; Goldman, C.R. 1976. Nitrogen uptake and release in a forested watershed, Lake Tahoe Basin, Nitrogen uptake and release in a forested watershed, Lake Tahoe Basin,

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Fahey, T.J. 1983. Nutrient dynamics of aboveground detritus in lodgepole pine (Pinus contorta ssp. Latifolia) ecosystems, ssp. Latifolia) ecosystems,

southeastern Wyoming. southeastern Wyoming. Ecological Monographs 53: 51-72. southeastern Wyoming.

Fahey, T.J.; Knight, D.H. 1986. Lodgepole pine ecosystems. Bioscience 36: 610 617. Lodgepole pine ecosystems

Galloway, J.N.; Schlesinger, W.H.; Levy II, H.; Michaels, A.; Schnoor, J.L. 1995. Nitrogen fixation: Anthropogenic

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Goldman, D.R.; Jassby, A.D.; Hackley, S.H. 1993. Decadal, interannual, and seasonal variability in enrichment bioassays at

Lake Tahoe, California Nevada, USA, Canadian Journal of Fisheries and Aquatic Science 50: 1459-1495. Lake Tahoe, California Nevada, USA

Hart, S.C.; Firestone, M.K. 1989. Forest floor-mineral soil interactions in the internal nitrogen cycle of an old-growth forest. Forest floor-mineral soil interactions in the internal nitrogen cycle of an old-growth forest.

Biogeochemistry 12: 103-127.

Johnson, D.W. 1995. Soil properties beneath Ceanothus and pine stands in the Eastern Sierra Nevada. Soil properties beneath Ceanothus and pine stands in the Eastern Sierra Nevada. Soil Science Society of Soil properties beneath Ceanothus and pine stands in the Eastern Sierra Nevada.

America, Journal 59: 918 924.

Johnson, D.W., Lindberg, S.E., eds. 1991. Atmospheric deposition and forest nutrient cycling: A synthesis of the Integrated

Forest Study. Forest Study. Ecological Series 91, New York: Springer-Verlag; 707 p. Forest Study.

Johnson, D.W.; Van Miegroet, H.; Lindberg, S.E.; Harrison, R.B.; Todd, D.E. 1991. Nutrient cycling in red spruce forests of the

Great Smoky Mountains. Great Smoky Mountains. Canadian Journal of Forest Research 21: 769-787. Great Smoky Mountains. 

Klemmedson, J.O. 1975. Nitrogen and carbon regimes in an ecosystem of young dense ponderosa pine in Arizona. Nitrogen and carbon regimes in an ecosystem of young dense ponderosa pine in Arizona. Forest Nitrogen and carbon regimes in an ecosystem of young dense ponderosa pine in Arizona.

Science 21: 163-168.

Leonard, R.L., Kaplan, L.A.; Elder, J.F.; Coats, R.N.; Goldman, C.R. 1980. Nutrient transport in surface runoff from a

subalpine watershed, Lake Tahoe Basin, California. subalpine watershed, Lake Tahoe Basin, California. Ecological Monographs 49: 281-310. subalpine watershed, Lake Tahoe Basin, California. 

Lindberg, S.E.; Johnson, D.W. 1989. Sampling and analysis protocols and project description for the Integrated Forest Study. Sampling and analysis protocols and project description for the Integrated Forest Study.

ORNL/TM-11241. Oak Ridge, TN: Oak Ridge National Laboratory.

Miller, D. F.; Walsh, P.A. 1991. Air quality and acidic deposition in the southwestern Sierra Nevada. Air quality and acidic deposition in the southwestern Sierra Nevada. Presented at the 84th Air quality and acidic deposition in the southwestern Sierra Nevada.

annual meeting and exhibition of the Air and Waste Management Association, Vancouver, British Columbia, Canada;

1991, June 16-21; 22 p.

Olsen, S.R.; Sommers, L.E. 1982. Phosphorus. Phosphorus. In: Phosphorus. Page, A.L.; R.H. Miller, R.H.; Keeney, D.R., eds. Methods of soil analysis.

Part 2. Chemical and microbiological properties. 2nd ed. Number 9 in the Series, Agronomy. Madison, WI: American

Society of Agronomy; 403-430.

Schimel, J.P.; Firestone, M.K. 1989. Nitrogen incorporation and flow through a coniferous forest soil profile. Nitrogen incorporation and flow through a coniferous forest soil profile. Soil Sci. Soc. Nitrogen incorporation and flow through a coniferous forest soil profile.

Amer. J. 53: 779-784.

Stark, N.M. 1973. Nutrient cycling in a Jeffrey pine ecosystem. Nutrient cycling in a Jeffrey pine ecosystem. Missoula, MT: University of Montana. Nutrient cycling in a Jeffrey pine ecosystem.

Timmer, V.R.; Stone, E.L. 1978. Comparative foliar analysis of young balsam fir fertilized with nitrogen, phosphorus, Comparative foliar analysis of young balsam fir fertilized with nitrogen, phosphorus,

potassium, and lime. Soil Science Society of America, Proceedings 42: 135-130. potassium, and lime

Session I1 Nutrient Fluxes in Forests of the Eastern Johnson, Susfalk, Dahlgren, Boucher, Bytnerowicz
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 105

Abstract

Nitrogenous (N) air pollutant concentrations and surface deposition of nitrate (NO3

- ) and ammonium (NH4+) to

branches of ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) seedlings were measured on a vertical transect in a

mature ponderosa/Jeffrey (Pinus jeffreyi Grev. & Balf.) pine canopy in the mixed conifer forest stand in the San

Bernardino Mountains of southern California during the 1992 to 1994 summer seasons (mid-May to mid-October

periods). In general, concentrations of nitric acid vapor (HNO3 ), nitrous acid vapor (HNO2 ), ammonia (NH3 ),

particulate NO3

-

 and NH4

+ were elevated and not affected by the position within the canopy. In contrast, surface

deposition of NO3

-

 and NH4

+ to ponderosa pine seedling branches strongly depended on position in the canopy. The

highest deposition of the two ions occurred at the canopy top, while at the canopy bottom deposition was often three to

four times lower. The range of deposition fluxes of NO3

-

 and NH4

+ were similar in each of the three seasons. A comparison

of the results of this study with other sites in California indicated that deposition fluxes of the studied ions were

elevated. In addition to the deposition of NO3

-

 and NH4

+, conductance (K1 ) of total NO3-

, HNO3 vapor, and particulate

NH4

+ to pine branch surfaces depended on the position in the canopy. The highest values were always measured at the

canopy top.

Introduction

Little is known about concentrations of nitrogenous (N) pollutants in forest locations

in California (Bytnerowicz and Fenn 1996). Annular denuder systems (Possanzini

and others 1983) allow for effective separation and precise determination of gaseous

and particulate N pollutants. These systems have been successfully used in chaparral

stands of the San Gabriel Mountains (Grosjean and Bytnerowicz 1993), mixed conifer

forests in the central Sierra Nevada (Bytnerowicz and Riechers 1995) and coastal

sage scrub communities in southern California (Allen and others, this volume).

Short-term, preliminary measurements of N pollutants in mixed conifer forests in

the San Bernardino Mountains have been reported (Fenn and Bytnerowicz 1993),

but detailed information on their temporal and spatial distribution is lacking.

In the arid climates of the California mountains, dry deposition is one of the

major mechanisms of ion transfer from the atmosphere to forests (Rundell and

Parsons 1977). Foliage rinsing techniques have been successfully used for

determinations of atmospheric deposition of various ions to trees (Bytnerowicz

and Fenn 1996, Bytnerowicz and others 1987, Lindberg and others 1986). This

technique is especially valuable in the dry climates of California where precipitation

(which would allow for throughfall collection) is scarce in the summer season.

This paper presents results of a study that monitored concentrations of gaseous

and particulate N air pollutants; measured deposition of nitrate (NO3

-

) and

ammonium (NH4

+) to branches of ponderosa pine seedlings; and calculated

conductance to branch surfaces (K1) of total nitrate, nitric acid vapor, and particulate

ammonium on a vertical transect in a canopy of mature ponderosa/Jeffrey pines

within the mixed conifer forest in the San Bernardino Mountains in southern

California during the mid-May to mid-October periods of 1992 to 1994.

Methods

The study was performed in Barton Flats in the San Bernardino Mountains at an

elevation of 2,100 m (fig. 1). A 30 m tall tower was located in the mixed conifer

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Ecologist, Plant Pathologist, Statistician, respectively, Pacific Southwest Research Station, USDA Forest Service,

4955 Canyon Crest Drive, Riverside, CA

92507.

Concentrations and Deposition of Nitrogenous Air

Pollutants in a Ponderosa/Jeffrey Pine Canopy 1

Andrzej Bytnerowicz,2 Mark E. Fenn,2 Michael J. Arbaugh2
106 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

forest stand of mature ponderosa/Jeffrey pines (Arbaugh and others, this volume).

The forest stand was open with about 67 percent of its ground area under tree

canopy (Fenn and Bytnerowicz 1996). Single annular denuder systems were placed

at each of four different levels of the tower: 29 m (canopy top), 24 m and 16 m

(canopy middle), and 12 m (canopy bottom) (fig. 2). Average 24-hour concentrations

of nitric acid (HNO3) vapor, nitrous acid (HNO2) vapor, ammonia (NH3), particulate

nitrate (NO3

-

), and particulate ammonium (NH4

+) were determined with an annular

denuder system (Peake and Legge 1987, Possanzini and others 1983). Constant air

flow of 17 L min-1 through the systems was provided by steady flow pumps. This

allowed for quantitative determinations of gaseous compounds deposited inside

annular denuder tubes and the fine particle fraction (< 2.2 ␣ m in diameter) collected

on teflon and nylon filters. Concentrations of NO3

 - and NH4+ in the annular tube

extracts and filter extracts were determined with ion chromatography (Dionex 4000i

chromatograph).3␣ Concentrations of N pollutants on the tower were measured six

times during the 1992 summer season, eight times during the 1993 summer season,

and four times during the 1994 summer season.

Deposition fluxes of NO3

-

 and NH4

+ to branches of ponderosa pine seedlings

were determined on each of the four levels of the tower in the vicinity of the annular

denuder systems. Four 2-year old seedlings were located at each of the tower levels.

Seedlings were about 40-50 cm tall and were grown in 6 L pulp pots filled with

2:1:1 mixture of Oakley sand, peat, and fir bark. The seedlings were watered with

tap water as needed (typically once a week). At the beginning of each collection

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Figure 1 — The study site,

Barton Flats, is located in the San

Bernardino Mountains in

southern California (a), south of

State Highway 38 (b).

Study Site

Silverwood

Lake Gregory Lake

Arrowhead

Holcomb Greek

Big Bear Lake

Baldwin

Lake (dry)

Bear 

River

1212 m

Los Angeles

Santa Ana

SantaRiver

Ana

Creek

Lake

1212 m 1212 m

San Bernardino 1212 m

Mountains

California

Kilometers

Study Area

Lake

National Forest boundary

1212 m Contour

(a)

(b) Barton Flats Area

San Bernardino Mountains

Barton Flats Area
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 107

period, seedling branches about 10 cm long were thoroughly rinsed with double

deionized water. The branches were re-rinsed at the end of 1-week long collection

periods with 100 mL of double deionized water and the rinses were collected in

250 mL Nalgene bottles. Bottles were placed on ice, immediately transferred to the

laboratory and placed at -18 oC. Concentrations of NO3

-

 were determined with ion

chromatography (Dionex 4000i ion chromatograph) and concentrations of NH4

+

by colorimetry (TRAACS, Model 800 instrument) (Bytnerowicz and others 1987).

Deposition was measured four times during the 1992 season, six times during the

1993 season, and five times during the 1994 photochemical smog season.

Conductances (K1) of total NO3

-

 (KNO3), HNO3 vapor (KHNO3) and particulate

NH4

+ (KNH4) to surfaces of pine branches have been calculated (Bytnerowicz and

others 1996). Conductance of these compounds is analogous to the deposition

velocity at a canopy level (vd) and is also expressed as cm s-1 (Hanson and Lindberg

1991). A general equation for conductance calculations is as follows:

K1 = F / c

in which:

F - deposition flux to branch surface (μg m-2 h-1)

c - pollutant concentration (μg m-3)

KNO3 was derived by dividing the NO3

-

 deposition flux to branch surfaces by a

sum of ambient concentrations of particulate NO3

-

 and gaseous HNO3 . KHNO3 was

calculated by dividing the deposition flux of NO3

-

 to branches that could be attributed

to HNO3 by the ambient concentrations of HNO3. Deposition fluxes of NO3

-

 attributed

to HNO3 were obtained by subtracting the portion of particulate NO3

-

 deposition

from the total NO3

-

 deposition (calculated by multiplying ambient concentrations of

particulate NO3

-

 by the literature value of particulate NO3

-

 deposition velocity to

conifer foliage) (Davidson and Wu 1990). KNH4 was calculated by dividing NH4

+

deposition to branches by the ambient concentrations of particulate NH4

+.

Figure 2 — Schematic of the

tower in the mature ponderosa/

Jeffrey pine canopy indicating the

four levels at which concentrations of N pollutants and

their deposition to branches were

measured.

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh

Meteorology
108 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Significance of the effects of position on the tower and a period of measurement

on concentrations, deposition fluxes, and K1 values of N pollutants during the

1992 to 1994 summer seasons were evaluated with a two-way analysis of variance

and a Bonferroni t-test (SigmaStat 1994).

Results

Concentrations of N pollutants

Seasonal concentrations of HNO3 vapor, NH3, particulate NO3

-

 and NH4

+ were

averaged at different heights in the canopy for the 1992 to 1994 seasons (table 1). In

the 1993 season the highest HNO3 vapor concentrations occurred at 24 m and were

significantly higher than the concentrations at 12 m. In 1992 and 1994, position

within the canopy had no significant effect on HNO3 concentrations. Position

within the canopy did not affect the concentrations of NH3, NO3

-

, or NH4

+ during

the entire study.

During the 3 years of study, the highest concentration of HNO3 vapor

(9.7 ␣μg m-3) was measured in June 1994 and the lowest (1.3 ␣μg m-3) in May 1993.

The highest concentration of NH3 (2.6 ␣μg m-3) was recorded in July 1992, while the

highest concentrations of particulate NO3

-

 (9.5 ␣μg m-3) and particulate NH4

+

(2.0 ␣μg m-3) were determined in June 1992. The lowest concentrations of

NH3, particulate NO3

-

 and particulate NH4

+ were determined in September

1992 (1.0, 0.35, and 0.2 ␣μg m-3, respectively). Concentrations of HNO2 vapor

during the 1992 season ranged from 0.10 to 0.42 ␣μg m-3, from 0.00 to 0.35 ␣μg m-3

in the 1993 season, and from 0.00 to 0.12 ␣μg m-3 in the 1994 season. In general, the

season significantly affected the concentrations of all studied N species, but no

consistent trends were evident (Miller and others 1996).

Table 1 — Average 24-hour concentrations of N pollutants in the ponderosa/Jeffrey pine canopy during three able 1 

summer seasons (1992-1994) of investigations (μg m3) Different letters assigned to the concentration numbers

indicate statistically significant differences between the canopy levels.

Canopy level Canopy level 1992 1993 1994

HNO3

29 m 3.65 3.24 ab 6.48

24 m 3.33 3.69 a 6.74

16 m 3.05 3.32 ab 6.78

12 m 3.00 3.14 b 5.86

P value 0.084 0.039 0.517

NH3

29 m 1.43 1.51 1.6

24 m 1.43 1.54 1.56

16 m 1.49 1.41 1.56

12 m 1.49 1.35 1.56

P value 0.841 0.119 0.963

NO3

-

29 m 3.86 2.37 2.33

24 m 3.3 2.44 2.33

16 m 3.32 2.72 2.37

12 m 2.97 2.92 1.91

P value 0.250 0.661 0.187

NH4

+

29 m 1.08 0.75 0.69

24 m 0.99 0.64 0.69

16 m 0.98 0.63 0.7

12 m 0.71 0.68 0.5

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 109

Surface Deposition of Washable Nitrate and Ammonium

Deposition of NO3

-

 to branches of the ponderosa pine seedlings was significantly

affected by position within the canopy (table 2). The highest deposition flux values

were always determined at the canopy top (29 m) and approached about 200 ␣μg m-2 h-1

during individual 1-week long determination periods (Bytnerowicz and others

1996). Deposition at 24 m was consistently lower than at the top but nearly always

higher than at 16 m and the canopy bottom (12 m). The highest deposition fluxes

occurred in July and August in each of the three years. Nitrate deposition values

were similar during the three years (Bytnerowicz and others 1996).

Ammonium deposition was also strongly affected by position within the

canopy (table 2). The highest deposition flux values occurred at the canopy top

and were sometimes as high as 20 ␣μg m-2 h-1 during individual determination

periods (Bytnerowicz and others 1996). Deposition at 24 m was higher than at

16 m and at 12 m. Although significant differences in deposition fluxes between

the individual collection periods occurred, no clear seasonal trend was

observed. The values of NH4

+ deposition flux were similar in all three seasons

(Bytnerowicz and others 1996).

NO3

-

, HNO3 and NH4

+ Conductance at a Branch Level

The highest conductance (K1) values for total NO3

-

, HNO3 vapor, and particulate

NH4

+ (table 3) were recorded at the canopy top and the lowest at the bottom of the

canopy. For total NO3

-

 and HNO3 vapor these differences were strongly significant

during three seasons. For particulate NH4

+ a significant effect of the position within

the canopy was seen only in 1993. Large differences were found between individual

periods of measurements within each season, i.e., during the 1993 season the range

of calculated KHNO3 values for the canopy top was between 0.4 cm s-1 and 3.1 cm s-1

(Bytnerowicz and others 1996). Significant differences between the KNO3 total and KHNO3

values during individual collection periods occurred in 1993 and 1994, and between

the KNH4 values only in 1993.

Discussion

Position within the canopy did not affect concentrations of the studied pollutants

probably because of the openness of the canopy upwind from the scaffold (with

Table 2 — Average seasonal deposition fluxes of NO able 2 3

-

 and NH4

+

 (μg m-2h-1 ) to seedling branches within the ponderosa

/Jeffrey pine canopy during three summer seasons (1992-1994). Different letters assigned to the deposition numbers

indicate statistically significant differences between the canopy levels.

Canopy level Canopy level 1992 1993 1994

NO3

-

29 m 126.2 a 118.6 a 132.6 a

24 m 67.6 b 79.8 b 84.0 b

16 m 40.7 c 51.9 c 62.9 bc

12 m 35.4 c 52.4 c 54.7 c

P value <0.0001 <0.0001 <0.0001

NH4

+

29 m 13.29 a 14.57 a 12.00 a

24 m 9.77 b 10.01 b 9.02 b

16 m 6.61 c 8.13 c 8.28 b

12 m 6.33 c 7.02 c 5.74 c

P value <0.0001 <0.0001 <0.0001

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh
110 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

the exception of significantly higher concentrations of HNO3 at 24 m than at 12 m

during the 1993 season). In the westerly direction, from which pollutants were

transported, no nearby canopy inhibited vertical mixing. In more dense forest

canopies, such as in the eastern United States, a vertical gradient of pollutants

with diminishing concentrations from canopy top to bottom is typical (Meyers

and others 1989).

Concentrations of HNO3 vapor at Barton Flats were moderately high (seasonal

means 3.0 to 6.8 ␣μg m-3) compared to the summer average concentrations in forest

locations in Tennessee (3.0 ␣μg m-3) and near Gottingen, Germany (5.8 ␣μg m-3)

(Lindberg and others 1990). At Tanbark Flat in the San Gabriel Mountains of

southern California, a mountain site which is considered one of the most polluted

in the United States (Bytnerowicz and others 1987), the average 24-hour

concentrations during the 1988 to 1991 summer seasons ranged from 6.7 to 11.7

␣μg m-3 (Grosjean and Bytnerowicz 1993). At various locations in the Sierra Nevada

the average 24-hour summer concentrations were below 1 ␣μg m-3 (Ashbaugh and

others 1991).

Nitrous acid vapor concentrations (seasonal means 0.00 to 0.42 ␣μg m-3) were

lower than the summer average concentrations at the Tanbark Flat site (0.55 to

0.65 ␣μg m-3) (Grosjean and Bytnerowicz 1993). However, they were higher than

the annual mean concentration of 0.05 ␣μg m-3 reported at the remote location in

the Canadian Rockies (Legge and Krupa 1989).

Ammonia concentrations (seasonal means 1.35 to 1.6 ␣μg m-3) were elevated

and similar to the seasonal means 1.1 to 1.85 ␣μg m-3 determined at Tanbark Flat

(Grosjean and Bytnerowicz 1993) and lower than the estimated seasonal mean

3.0 ␣μg m-3 determined near Gottingen, Germany (Lindberg and others 1990).

The measured concentrations were several times higher than the annual mean

0.26 ␣μg m-3 value measured in the Canadian Rockies (Legge and Krupa 1989)

Table 3 Table 3 — Average conductance values (K Table 3 1 ) of N species to single ponderosa pine branches during 3 years (1992-1994)

of investigations (cm s-1 ). Different letters assigned to the K1 values indicate statistically significant differences between

canopy levels.

Canopy level Canopy level 1992 1993 1994

NO3

-

 total

29 m 0.50 a 0.61 a 0.42 a

24 m 0.31 ab 0.45 ab 0.27 ab

16 m 0.16 b 0.29 bc 0.21 b

12 m 0.16 b 0.26 c 0.19 b

P value <0.002 <0.000 <0.002

HNO3

29 m 0.83 a 1.26 a 0.57 a

24 m 0.48 ab 0.74 ab 0.36 ab

16 m 0.30 b 0.55 b 0.26 b

12 m 0.25 b 0.52 b 0.26 b

P value <0.010 <0.003 <0.011

NH4

+

29 m 0.28 0.66a 0.66

24 m 0.22 0.50 ab 0.29

16 m 0.16 0.40 b 0.25

12 m 0.18 0.34 b 0.29

P value 0.099 0.012 0.160

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 111

and the estimated seasonal mean 0.2 ␣μg m-3 in Tennessee (Lindberg and others

1990). However, concentrations of NH3 in the Netherlands and Great Britain may

be several times higher than the reported values reaching 15 ␣μg m-3 (Draaijers and

others 1989, Keuken and others 1988).

Concentrations of particulate NO3

-

 (seasonal means 1.9 to 3.9 ␣μg m-3) were

high and similar to the values determined at Tanbark Flat (seasonal means 1.75 to

2.9 ␣μg m-3) (Grosjean and Bytnerowicz 1993). These values greatly exceeded the

annual mean 0.13 ␣μg m-3 reported for the Canadian Rockies (Legge and Krupa

1989), and seasonal mean 0.19 ␣μg m-3 determined in Tennessee (Lindberg and others

1990). The reported values were similar to 2.1 to 5.1 ␣μg m-3 measured in the San

Bernardino Mountains (Fenn and Bytnerowicz 1993) and 3.3 ␣μg m-3 measured in

the Los Angeles Basin, California (Sickles and others 1988). Mean NO3

-

concentration

in Claremont, Los Angeles Basin (7.9 ␣μg m-3) (Pierson and Brachaczek 1988) was

higher than the concentrations determined in this study.

Concentrations of particulate NH4

+ (seasonal means 0.5 to 1.1 ␣μg m-3) were

similar to the values determined at Tanbark Flat (seasonal means 0.65 to 0.85 ␣μg

m-3) (Grosjean and Bytnerowicz 1993) and other locations in the San Bernardino

Mountains (0.1 to 0.9 ␣μg m-3) (Fenn and Bytnerowicz 1993). Ammonium

concentrations were also similar to the mean concentration of 1.0 ␣μg m-3 determined

in the urban Los Angeles area (Sickles and others 1988). Ammonium

concentrations reported for the forest locations in Tennessee and Germany

(seasonal means 2.4 and 5.5 ␣μg m-3, respectively) (Lindberg and others 1990)

were higher than in this study.

Nitrate fluxes to branches were elevated; the highest values during individual

collection periods (approaching 200 ␣μg m-2 h-1) were similar to the value of 180 ␣μg

m-2 h-1 determined for mature ponderosa pine in Camp Paivika of the San

Bernardino National Forest (Fenn and Bytnerowicz 1993). Higher NO3

-

 deposition

values were determined at Tanbark Flat; deposition to hoaryleaf ceanothus

(Ceanothus crassifolius) ranged from 88 to 455 ␣μg m-2 h-1 (Bytnerowicz and others

1987). Deposition of nitrate to native pines in other parts of the state were much

lower: about 4 to 6 ␣μg m-2 h-1 for Eastern Brook Lake in the eastern Sierra Nevada

(Bytnerowicz and others 1992); 27 to 30 ␣μg m-2 h-1 for Emerald Lake in the western

Sierra Nevada (Bytnerowicz and others 1991); and 6 to 31 ␣μg m-2 h-1 for Whitaker

Forest in the western Sierra Nevada (Bytnerowicz and Riechers 1995).

Deposition fluxes of NH4

+ were also elevated at Barton Flats; the highest values

determined during individual collection periods approached 20␣μg m-2 h-1

(Bytnerowicz and others 1996). These values were similar to deposition of 21␣μg

NH4

+ m-2 h-1 determined at Camp Paivika, a high pollution site in the western San

Bernardino Mountains (Fenn and Bytnerowicz 1993). Deposition of NH4

+ to

hoaryleaf ceanothus at Tanbark Flat (17 to 46 ␣μg m-2 h-1) was higher than at Barton

Flats (Bytnerowicz and others 1987). Deposition to native pines at some other forest

sites in California were lower than in the present study: at Eastern Brook Lake

ranged from 0.8 to 1.6 ␣μg m-2 h-1 (Bytnerowicz and others 1992); Emerald Lake, 6.0

to 9.0 ␣μg m-2 h-1 (Bytnerowicz and others 1991); and Whitaker Forest, 1.0 to 1.5 ␣μg

m-2 h-1 (Bytnerowicz and Riechers 1995). The results of this study have clearly

indicated that NO3

-

 and NH4

+ deposition fluxes in a mixed coniferous forest strongly

depend on the position of the branches within the canopy. Ion deposition increased

with increasing branch height. Because concentrations of gaseous and particulate

pollutants did not depend on canopy position, the observed distribution of

deposition fluxes within the forest canopy could not be related to the air pollution

concentration differences. Difference in deposition fluxes could be attributed to

varying wind speed occurring along the vertical transect within the canopy because

the highest deposition values occurred at the canopy top where wind speeds were

highest (Arbaugh and others, this volume). The HNO3 conductance (kHNO3) values

at the canopy top reached 3.1 cm s-1 (Bytnerowicz and others 1996) and were similar

to the kHNO3 values of 0.6 to 3.4 cm s-1 determined for loblolly pine in Tennessee

(Hanson and Garten 1992). In general, the conductance of a pollutant to a branch

Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh
112 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

multiplied by the ambient pollutant concentration provides an estimate of

deposition at the branch level. Branch level deposition multiplied by the canopy

leaf area index (LAI) provides an estimate of canopy level deposition. Therefore,

theoretically, the KHNO3 and ambient HNO3 concentration values can be used for an

estimate of surface NO3

-

 deposition to ponderosa pine canopies from atmospheric

HNO3. Similarly, information on conductance to branches of total NO3

-

 and

particulate NH4

+ can be used for estimates of surface dry deposition of total NO3-

and NH4

+ if information on ambient concentrations of HNO3

 vapor + particulate

NO3

-

 or particulate NH4

+ are provided. On the basis of the results of this study, and

other related companion studies, empirical models for estimating N atmospheric

deposition to the mixed conifer forest stands in California are being developed

(Arbaugh and others, this volume).

Total N deposition to the forest stand in Barton Flats was estimated based on

the presented results, information on deposition to other dominant tree species,

and LAI for the overstory species (Bytnerowicz and others 1996). The estimated

forest stand deposition for the 1993 season was 5 to 9 kg ha-1 yr-1, and occurred

mostly as dry deposition to surfaces of plants and soil (Bytnerowicz and others

1996). Judging from the results of this and other studies (Fenn and Bytnerowicz

1993, Grosjean and Bytnerowicz 1993), Barton Flats can be characterized as a site

with elevated N deposition located in the middle to low end of the west to east air

pollution gradient in the San Bernardino Mountains. Chronic N deposition in the

more polluted regions of the San Bernardino and San Gabriel Mountains is

associated with symptoms of N excess as described by Aber and others (1989),

which are especially evident as high NO3

- concentrations in soil solution and

streamwater (Fenn and others 1996, Riggan and others 1985). Long-term effects of

N deposition on plant community dynamics in these forests are as yet unknown.

In summary, concentrations of the measured N species at the Barton Flats site

were elevated compared to the pristine mountain areas in California and elsewhere.

No differences in concentrations of the measured N species were found on the

vertical transect in the mature pine canopy. However, deposition fluxes of NO3

-

and NH4

+ as well as conductance (K1

) of gaseous HNO3, total NO3

-

, and particulate

NH4

+ were always the highest at the canopy top.

Acknowledgments

This study was partially sponsored by the California Air Resources Board, Contract

No. A032-180. The support of Brent Takemoto is greatly appreciated, and we thank

Robert Glaubig and the staff of the chemical laboratory of the Pacific Southwest

Research Station, USDA Forest Service, for their excellent technical assistance. We

thank Laurie Dunn for technical editing of this manuscript.

References

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Session I1 Concentrations and Deposition of Nitrogenous Air Pollutants Bytnerowicz, Fenn, Arbaugh

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 115

Abstract

A 3-year study of nitrogenous (N) air pollution deposition to ponderosa pine (Pinus ponderosa Dougl. ex. Laws.)

seedlings along a mature tree vertical canopy gradient was conducted in the mixed conifer forest of the San Bernardino

Mountains of southern California. Concentrations of nitric acid vapor (HNO3 ), particulate nitrate (NO3

-

 ), and

ammonium (NH4

+

) were measured, as well as dry surface deposition of NO3

-

 and NH4

+

. By using this data, along with

meteorological information, a series of simple models were developed that predict the vertical gradient of foliage-rinse

surface deposition for NO3 

-

, NH4

+

, and total N. Individual models for NO3

-

 and NH4

+

 were calculated by using

deposition data, air concentrations at the top of the canopy, and wind speed. These models explained 80 percent of the

variation between deposition values at the lower canopy positions. Examination of model coefficients indicated that

the two models were not significantly different, and a single model was developed to estimate total N deposition. This

model explained less variation than the individual models (R2

N = 0.69) but is a simpler description of the system. All

models have two parameters (a0 and a1) that are estimated by nonlinear regression. Independent data indicates that

the a1 parameter depends only on the rate of decline in wind speed vertically down the canopy. The a0 parameter

reflects the difference between foliage rinse surface deposition at the top of the canopy and that at lower canopy positions.

It indicates that about 20 percent less dry surface deposition occurs at lower canopy positions than expected from

wind speed alone for all nitrogenous compounds, most likely because decreased turbulence within the canopy results in

increased quasi-laminar boundary layer resistance at lower canopy positions.

Introduction

Dry deposition is one of the major mechanisms of nitrogenous (N) transfer from

the atmosphere to plant surfaces in arid California mountains (Bytnerowicz and

Fenn 1996, Rundell and Parsons 1977). High dry deposition fluxes of N pollutants

have been reported in exposed areas of the San Gabriel and San Bernardino

Mountains (Bytnerowicz and others 1987, Fenn and Bytnerowicz 1993), and in lesser

amounts in the southern and western portions of the Sierra Nevada Mountains

(Bytnerowicz and others 1991).

Foliage rinsing techniques (Bytnerowicz and others 1987, Lindberg and Lovett

1985) have been successfully used for determinations of atmospheric deposition

of various ions to trees. The technique is especially valuable in dry climates of

southern California where in summer and fall periods precipitation is scarce, which

limits collecting throughfall measures of deposition.

Extensive foliage rinse sampling requires considerable time and expense to

analyze. Seedlings must be obtained and stored in greenhouses before field

exposure. A scaffold tower must be erected at the remote site, and periodic watering

and rinsing must be done for groups of seedlings at different tower heights to

obtain sufficient information to quantify the vertical deposition profile. This

extensive sampling reduces the number of sample locations and times that can be

directly measured using this approach.

Expanding limited foliage rinse measurements to larger areas and longer time

periods has been accomplished in some systems by using inferential models that

have been developed over the last decade (Hicks and Meyers 1988, Meyers and

Baldocchi 1988). These models estimate deposition flux (Fd) from air concentration

measurements (C) and deposition conductance (K1) by using air concentrations

and micrometeorological information. To use inferential models, aerodynamic

transport, boundary layer transport, and physiochemical surface interactions must

be estimated for model input. This model approach works best when ecosystems

Predicting Nitrogen Flux Along a Vertical Canopy Gradient

in a Mixed Conifer Forest Stand of the San Bernardino

Mountains in California1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Statistician, Ecologist, and Plant

Pathologist, respectively, Pacific

Southwest Research Station, Forest

Service, 4955 Canyon Crest Dr.,

Riverside, CA 92507.

Michael J. Arbaugh,2 Andrzej Bytnerowicz,2 and Mark E. Fenn2
116 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

have uniform canopy and vegetation and when atmospheric conditions are stable

and predictable.

The mixed conifer forest of the San Bernardino Mountains is quite different

(fig. 1) from these systems. This forest type is comprised of several species including,

ponderosa pine (Pinus ponderosa Dougl. ex. Laws) or Jeffrey pine (Pinus jeffreyi

Grev & Balf.) (in transition zones both species are found), sugar pine (Pinus

lambertiana), white fir (Abies concolor), incense cedar (Calocedrus decurrens), California

black oak (Quercus kelloggii), and a herbaceous or shrubby understory. The forest

often has a discontinuous canopy that results in increased atmospheric mixing

and edge effects that seldom result in a significant vertical air pollutant

concentration gradient (Bytnerowicz and others, this volume). However, equal

deposition does not occur over the canopy. These conditions reduce the ability to

apply inferential models to this system, and can result in excessive estimates of N

by inferential models (Taylor and others 1996) compared with empirical estimates.

In this study we measured atmospheric concentrations and foliar deposition

in the San Bernardino Mountains to determine relationships between atmospheric

concentrations, meteorological factors, and deposition. The primary data consists

of annular denuder and foliage-rinse data gathered at several heights on a

scaffolding tower erected between the canopies of three trees (Miller and others

1996). Filter pack data (Chow and Watson 1996) that continuously monitored air

pollution and meteorology at a location about 5 kilometers from the tower location

was used to corroborate the results of the annular denuder data.

Our goal was to develop a predictive model for foliage rinse flux along a vertical

canopy profile that minimizes the number of flux measurements needed to estimate

N deposition flux to mature pine canopies. The model developed is intended to

reduce the amount of time and expense to determine deposition vertically along

the canopy, and enable estimates of total N dry deposition to the forest.

Methods

Modeling and Analysis Data

As part of a multi-disciplinary study (Miller and others 1996), foliar rinse data were

gathered to determine the effect of canopy position on deposition of nitrogenous

and sulfurous pollutants, to compare deposition to branches of mature trees and

seedlings, and to compare deposition to three tree species (ponderosa/Jeffrey pine,

white fir, and California black oak). Measurement of nitrate, ammonium, and sulfate

deposition to branches of seedlings and mature trees of ponderosa pine at various

levels of the canopy were performed during three photochemical smog seasons

(1992, 1993 and 1994). Fluxes of NO3

-

 and NH4

+

 to branches of ponderosa pine

seedlings were determined on a tower located in a stand of mature ponderosa/

Jeffrey pines. Four seedlings were located at each of four different levels of the

tower (29 m at the top of the canopy; 24 m and 16 m in the middle of the canopy;

and 12 m at the canopy bottom).

In the beginning of each collection period branches about 10 cm long were

thoroughly rinsed with double deionized water. The branches were re-rinsed at

the end of the collection period with about 100 mL of double deionized water and

the rinses were collected in 250 mL Nalgene bottles. Bottles were placed on ice,

immediately transferred to the laboratory and placed at -18 °C. Concentrations of

nitrate were determined with ion chromatography (Dionex 4000i ion

chromatograph)3␣ and concentrations of ammonium colorimetrically with a

Technicon TRAACS Autoanalyzer.

The filter pack dry deposition sampling system (Chow and others 1993) was

equipped with an acid-washed PFA (perfluoralkoxy) sampling surface with PFA

Teflon-coated Bendix-Sensidyne Model 240 cyclone. Particles and gases were drawn

through the Teflon-coated PM2.5 inlet at a constant flow rate of 113 l/min. A total of

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn

Figure 1 — The Barton Flats

Tower site in a mixed conifer

forest in the San Bernardino

Moutnains in southern Califronia

photographed aerially to illustrate

the diverse canopy structure of

the study site. Canopy diameters

vary and are interspersed with

large open areas. The arrow

points to the location of the

scaffold tower where data was

gathered for this study.

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 117

62 l/min was drawn through four filter packs simultaneously. In routine operation,

air was drawn through the filter pack for NO2 sampling at a flow rate of 2 l/min

and through the remaining three filter packs at a flow rate of 20 l/min apiece.

Four additional ports were provided for field blanks that were used to evaluate

filter loadings during passive sampling periods and during filter handling.

Daytime (0600 to 1800 PST) and nighttime (1800 to next day 0600 PST)

measurements were taken every sixth day between 11/02/91 and 09/28/93 and

between 06/01/94 and 08/30/94 for ammonia (NH3) and nitric acid (HNO3) gases

on absorbing substrates, and ammonium (NH4+) and nitrate (NO3- ) particulates

on filter substrates.

Model Development

Empirical relationships are subject to spurious correlations and relationships,

especially when limited numbers of locations and sampling times are available. To

reduce this effect, we based our model upon simplified theoretical relationships

rather than observed ones. We chose the equilibrium relationship between aerosol

and gaseous concentrations as the basis for model formulation:

NH4NO3 <=> HNO3 + NH3

If it is assumed that all particulates are deposited as NH4NO3, then it can be

assumed that:

KNO3p= KNH4p= KNH4NO3

in which:

K = conductance of N to the plant surface (cm s-1)

NO3p = N from particulate NO3-

NH4

+p = N from particulate NH4

+

Expressing atmospheric concentrations (C) as ␣μg m-3 N, then

C NO3p = CNH4p

These relationships implicitly assume that NH4+p deposition to plant surfaces is

caused by NH4+ rather than NH3. Several studies that have linked diurnal patterns

of NH3 deposition to stomatal aperture (Aneja and others 1986, Hutchinson and

others 1972, Rogers and Aneja 1980) found that little NH4+ is absorbed cuticularly

(Van Hove and others 1987), and that NH3 deposition contributes little to surface

deposition (Van Hove and others 1987).

The equilibrium relationship is also only valid if the majority of particulates

are in the range of 0.1 μm to 2␣μm diameter. Wall and others (1988) found nitrate

and ammonium particulate sizes vary between 0.52 μm and about 2.0 μm in a

study conducted in Claremont, California. Similar results were found by Lundgren

and others (1979) in a study conducted in Riverside, California, which is about 50

km from our study site.

Both the annular denuder and filter pack data were used to examine model

assumptions. High significant correlations were found by both systems between

particulate NO3

-

 and particulate NH4

+

 concentrations ( r=0.92 and r=0.93,

respectively) during foliage wash periods, indicating that the majority of

particulates were in the form of NH4NO3. Average NO3

-

 concentrations were 1.80

mg N/m3± 0.98 mg N/m3 compared to 1.98 mg N/m3± 0.74 mg N/m3 estimated

from NH4

+

. The two estimates are not significantly different (paired t-test, p = 0.05).

By using these relationships, flux (F) and atmospheric concentration (C) data

can be expressed as:

F NO3p = FNH4p

and

FHNO3 = FNO3t - FNH4p

in which F is flux (μg N m-2 s-1).

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn
118 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Fluxes at lower canopy positions can be estimated for nitrate deposition as:

FNO3h = a0e[a1(1- Hh/Ht)](FHNO3t + FNO3t) + eh

for ammonium deposition as:

FNO3h = a0e[a1(1- Hh/Ht)]( FNH4t) + eh

or for either type of deposition as:

FNh = a0e[a1(1- Hh/Ht)]( FNt) + eh

in which:

h = vertical height along canopy (H)

t = max(h) = top of canopy

FNt = flux to leaves from either nitrate and nitric acid or ammonium

a0, a1 = constant terms estimated from nonlinear regression

eh = independently and identically Gaussian distributed errors.

The model was fitted with a nonlinear regression procedure, NLIN (SAS

1990), by using the derivative-free multivariate secant method. The starting value

of wind speed parameter a1 = - 1.2 was estimated as:

Wind speed = e[a1(1- Ht/Hh)],

from wind speed information gathered during 1994 at the tower location (Miller

and others 1996). A grid search was used to identify a starting value of a0. Graphical

and analytical analyses of residuals were conducted to determine if any model

assumptions were violated.

Results

Model results indicate that deposition estimated at the top of the canopy is strongly

related to deposition at lower canopy positions for individual ion models (table 1).

Nonlinear R2 (analogous but not equal to linear R2) was high for both pollutants,

indicating that conductance for both compounds at lower levels of the canopy is

also strongly linked to conductance at the top of the canopy (figs. 2a, 3a).

The observed decline in flux is dominated by declines in wind speed from the

top to the bottom of the canopy. Neither NO3

-

 or NH4

+

 model a1 parameters were

significantly different from a1 developed from wind speed information. The a1

parameters from both models are lower than those reported for other systems

(Davidson and Wu 1990), indicating that wind speed declines more linearly in this

open forest relative to closed canopy forests.

Table 1 Table 1 — Model parameters and summary statistics for individual ion and combined ion deposition models. Table 1 1

 Model n Mean Std 2 a0 (SE) 3 a1 (SE) RMSE 4 R2

N

NO3- 33 6.79 0.98 0.761 (0.05) -1.39 (0.17) 1.03 0.81

NH4

+ 33 6.25 0.77 0.853 (0.05) -1.04 (0.15) 1.28 0.80 +

N

either 66 6.52 0.88 0.791 (0.04) -1.26 (0.14) 1.27 0.69

1 All models use the equation: Fh=a0e[a1(1-Ht

/Hh)](Ft) + ehj .

All ion values are expressed as mg N/m2/hr.

2 Standard deviation.

3 Standard error.

4 Root mean square error.

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 119

Coefficients were not significantly different (at p = 0.05) for the NO3

-

 and NH4

+

models, indicating that it might be possible to develop a combined model. A single

model was parameterized using NO3

- and NH4+

 expressed as mg N. Model results

indicate that the single model, although marginally poorer for explaining variation,

may be a simpler representation of the system. The combined ion model a1

parameter estimate was nearly identical to that estimated using wind speed

information (a1 = -1.2). Individual model differences (table 1) may be due to smaller

data sets used in the analysis.

Interpretation of a0 is more difficult for these models. The parameter reflects a

proportional reduction in the estimated foliar deposition at lower canopy positions

relative to the canopy top. This constant reduction does not depend on canopy

position and is similar to the proportion of total deposition retained on foliage

after rinsing or throughfall occurs (Friedland and others 1991, Lovett and Lindberg

1993). Model differences are not, however, the result of canopy retained N; rather,

the parameter may reflect an additional resistance to deposition in the canopy not

present at the more open areas of the canopy top. Although data is sparse, this

resistance may result from reduced turbulence occurring in the canopy interior

that results in increased quasi-laminar layer depth and stability, which increases

diffusive and interception resistance. Detailed measurements are needed, however,

to confirm this interpretation.

Residual distribution was best for the combined model (figs. 2b, 3b, 4b). No

significant heteroscadisticity or serial correlation was found for the models, nor

was there any apparent relationship between canopy position and residual

distribution. Analysis of residuals did suggest that individual ion models may

overestimate deposition at low air concentrations (figs. 2b, 3b). This may be

due to reductions in the ion concentrations near the ground caused by reduced

atmospheric mixing. This overestimation was lessened in the combined model

(fig. 4b).

Figure 2 — Model results for

predicting NO3

-

 deposition at 24

m, 16 m, and 12 m on the canopy.

Predicted deposition is compared

with observed; the line represents

a perfect model fit (1:1) and

corresponds to deposition

calculated at 29 m (A). Model

residuals lack any pattern that

would indicate poor model fit (B).

120

100

80

60

40

20

Predicted Deposition (

0

μg/m2/hr)

–

–

–

–

–

A

R2 = 0.81

1:1

NO3-

0 20 40 60 80 100 120 140

Observed deposition (μg/m2/hr)

––––––––

Observed deposition (μg/m2/hr)

Residual

30

20

10

0

- 10

- 20

0 20 40 60 80 100 120 140

B

–

–

–

–

––––––__

–

–

–

–

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn
120 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 3 — Model results for

predicting NH4

+

 deposition at 24

m, 16 m, and 12 m on the canopy.

Predicted deposition is compared

with observed; the line represents

a perfect model fit (1:1) and

corresponds to deposition

calculated at 29 m (A). Model

residuals lack any pattern that

would indicate poor model fit (B).

Figure 4 — Single model results

for N deposition from ammonium and nitrate. Predicted

deposition is compared with

observed; the line represents a

perfect model fit (1:1) and

corresponds to deposition

calculated at 29 m (A). Model

residuals lack any pattern that

would indicate poor model fit (B).

20

15

10

5

0

Predicted Deposition (

μg/m2/hr)

–

–

–

A

R2 = 0.80

1:1

NO4+

0 5 10 15 20

Observed deposition (μg/m2/hr)

–––––

Observed deposition (μg/m2/hr)

Residual

4

2

0

- 2

- 4

0 5 10 15 20

B

–

–

–

–––––

–

–

–

–

15

10

5

Predicted Deposition (

0

μg/m2/hr)

–

–

A

R2 = 0.69 1:1

N

0 2 4 6 8 10 12 14

Observed deposition (μg/m2/hr)

–––––––

Observed deposition (μg/m2/hr)

Residual

4

2

0

- 2

- 4

B

–

–

–

–

0 2 4 6 8 10 12 14

––––––––

–

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 121

Conclusions

The estimators developed in this study have potential for reducing the necessary

work in future studies to obtain estimates of whole canopy flux. The physically

based models used in this study have high R2, favorable residual properties, and

interpretable parameters. Study results indicate that the models are applicable over

a range of concentration values and pollution exposure seasons. The models need

to be tested at other locations and for other tree species to verify their predictive

ability. The scaffold tower data used for the analysis represents only a single point,

and the foliage extraction information is most relevant for only two species,

ponderosa and Jeffrey pine. Additional locations, especially in the western San

Bernardino Mountains and the southern Sierra Nevada, need to be examined.

A single combined model may have application throughout the Transverse

Ranges in southern California and the southern Sierra Nevada, since the

chemical composition of N pollutants and forest types are similar for these

areas. It may also have application to other open montane systems that lack

vertical concentration gradients of pollutants and where the dominate form of

aerosol N pollution is NH4NO3. Such systems may include coastal sage scrub,

oak-grasslands, and chapparal.

References

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species. species. Journal of the Air Pollution Control Association 36:1338-1341. species. 

Bytnerowicz, Andrzej; Dawson, P. J.; Morrison, C. L.; Poe, M. P. 1991. Deposition of atmospheric ions to pine branches and

surrogate surfaces in the vicinity of Emerald Lake watershed, Sequoia National Park. surrogate surfaces in the vicinity of Emerald Lake watershed, Sequoia National Park. Atmospheric Environment 25A: surrogate surfaces in the vicinity of Emerald Lake watershed, Sequoia National Park. 

2203-2210.

Bytnerowicz, Andrzej; Fenn, Mark E. 1996. Nitrogen deposition in California forests: A review. Nitrogen deposition in California forests: A review. Environmental Pollution, Nitrogen deposition in California forests: A review.

92:127-146.

Bytnerowicz, Andrzej; Fenn, Mark E.; Arbaugh, Michael J. 1998. Concentrations and deposition of N air Pollutants in a

ponderosa/Jeffrey pine canopy. ponderosa/Jeffrey pine canopy. In: Bytnerowicz, Andrzej; Arbaugh, Michael, J.; Schilling, Susan, technical coordinators. ponderosa/Jeffrey pine canopy.

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Service [this volume].

Bytnerowicz, Andrzej; Fenn Mark E.; Glaubig, Robert. 1996. Dry deposition of nitrogen and sulfur to forest canopies at three

plots. In: Assessment of acidic deposition and ozone effects on conifer forests in the San Bernardino Mountains. Final plots.

Report to California air Resources Board, Contract No. A032-180; 4-1 to 4-48 p.

Bytnerowicz, Andrzej; Miller, Paul R.; Olszyk, D. M.; Dawson, P. J.; Fox, Carl A. 1987. Gaseous and particulate air pollution in

the San Gabriel Mountains of southern California. the San Gabriel Mountains of southern California. Atmospheric Environment (21):1749-1757. the San Gabriel Mountains of southern California.

Chow, Judith; Watson, John G. 1996. Characterization of the aerometric data at the Barton Flats station. In: Assessment of Characterization of the aerometric data at the Barton Flats station

acidic deposition and ozone effects on conifer forests in the San Bernardino Mountains. Final Report to California Air

Resources Board, Contract No. A032-180; 2-1 to 2-34 p.

Chow, Judith C.; Watson, John G.; Bowen, J. L.; Gertler, A.W.; Frazier, C. A.; Fung, K.K.; Ashbaugh, L. 1993. A sampling

system for reactive species in the western U.W. system for reactive species in the western U.W. In: E. Winegar, ed. ACS Symposium Series, Washington D.C.: American system for reactive species in the western U.W. 

Chemical Society; 209-228.

Davidson, C. I.; Wu, Y-L. 1990. Dry deposition of particles and vapors. Dry deposition of particles and vapors. In: Lindberg, A. S.; Page, E. L; Norton, S. A., eds. Acidic Dry deposition of particles and vapors. 

precipitation, Vol. 3. Sources, deposition and canopy interactions. New York: Springer-Verlag; 103-216.

Fenn, Mark E.; Bytnerowicz, Andrzej. 1993. Dry deposition of nitrogen and sulfur to ponderosa and Jeffrey pine in the San

Bernardino National Forest in southern California. Bernardino National Forest in southern California. Environmental Pollution 81: 277-285. Bernardino National Forest in southern California. 

Friedland, Arthur J.; Miller, E. K.; Battles, J. J.; Thorne, J. F. 1991. Nitrogen deposition, distribution and cycling in a subalpine

spruce-fir forest in the Adirondacks, New York, USA. spruce-fir forest in the Adirondacks, New York, USA. Biogeochemistry 14:31-55. spruce-fir forest in the Adirondacks, New York, USA.

Hicks, B. B.; Meyers T.P. 1988. Measuring and modeling dry deposition in mountainous areas. Measuring and modeling dry deposition in mountainous areas. In: Unsworth M. H.; Fowler B., Measuring and modeling dry deposition in mountainous areas.

eds. Acid deposition at high elevation sites. Kluwer, Amsterdam; 541-552.

Hutchinson G. L.; Millington R. J.; Peters D. B. 1972. Atmospheric ammonia: absorption by plant leaves. Atmospheric ammonia: absorption by plant leaves. Science 175:771-772. Atmospheric ammonia: absorption by plant leaves.

Lindberg, Steven E.; Lovett, G. M. 1985. Field measurements of particle dry deposition rates to foliage and inert surfaces in a

forest canopy. forest canopy. Environmental Science and Technology 19:238-244. forest canopy. 

Lovett, G.M.; Lindberg, Steve E. 1993. Atmospheric deposition and canopy interactions of nitrogen in forests. Atmospheric deposition and canopy interactions of nitrogen in forests. Canadian Atmospheric deposition and canopy interactions of nitrogen in forests.

Journal of Forest Research 23:1603-1616.

Lundgren, D.A.; Harris, F. S. Marlow, W. H.; Lippmann, M.; Clark, W. E.; Durham, M. D. 1979. Aerosol measurement. Aerosol measurement.

Gainesville, Florida: University Presses of Florida.

Meyers, T. P.; Baldocchi, D. D. 1988. A comparison of models for deriving dry deposition fluxes of O3

 and SO2 to a forest

canopy. canopy. Tellus 40B:270-284. canopy. 

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Miller, Paul R.; Bytnerowicz, Andrzej; Jones, David; Schilling, Susan. 1996. Characterization of ozone, particle chemistry and

meteorology measurements in support of canopy deposition estimates. meteorology measurements in support of canopy deposition estimates. In: Assessment of acidic deposition and ozone meteorology measurements in support of canopy deposition estimates.

effects on conifer forests in the San Bernardino Mountains. Final Report to California Air Resources Board, Contract No.

A032-180; 3-1 to 3-9 p.

Rogers, H. H.; Aneja, V. P. 1980. Uptake of atmospheric ammonia by selected plant species. Uptake of atmospheric ammonia by selected plant species. Atmospheric Environ- Uptake of atmospheric ammonia by selected plant species.

ment 20:251-257.

Rundell, Philip W.; Parsons, David J. 1977. Montane and subalpine vegetation in the Sierra Nevada and Cascade ranges. Montane and subalpine vegetation in the Sierra Nevada and Cascade ranges. In: Montane and subalpine vegetation in the Sierra Nevada and Cascade ranges.

Barbour, M. J.; Major, J., eds. Terrestrial Vegetation of California, New York:Wiley: 559-599.

SAS Inc. 1990. SAS/STAT Users Guide, Version 6, Volume 2. SAS/STAT Users Guide, Version 6, Volume 2. 4th ed. Cary, NC: SAS Institute. SAS/STAT Users Guide, Version 6, Volume 2. 

Taylor, George Jr.; Beck, K.; Beaulieu, N.; Gustin, M. 1996. Modelling dry deposition of nitrogen, sulfur and ozone to the

forest stand at Barton flats. forest stand at Barton flats. In: Assessment of acidic deposition and ozone effects on conifer forests in the San Bernardino forest stand at Barton flats. 

Mountains. Final Report to California Air Resources Board, Contract No. A032-180; 8-1 to 8-31 p.

Van Hove, L. W. A.; Koops, A. J.; Adema, E.H.; Vrendenberg, W. J.; Pieters, G. A. 1987. Analysis of the uptake of atmospheric

ammonia by leaves of ammonia by leaves of Phasiolus vulgaris L. Atmospheric Environment 21:1759-1763. L.

Van Hove, L. W. A.; Koops, A. J.; Adema, E.H.; Vrendenberg, W. J.; Pieters, G. A. 1989. Physiological effects of long- Physiological effects of longterm exposure to low and moderate concentrations of atmospheric NH3

 on poplar trees. Plant Cell and Environ- on poplar trees

ment 12:899-908.

Wall, S. M.; John, W.; Ondo, J. L. 1988. Measurement of aerosol size distributions for nitrate and major ionic species. Measurement of aerosol size distributions for nitrate and major ionic species. Atmo- Measurement of aerosol size distributions for nitrate and major ionic species.

spheric Environment 22:1649-1656.

Session I1 Predicting Nitrogen Flux Along a Vertical Canopy Gradient Arbaugh, Bytnerowicz, Fenn
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 123

Abstract

Indicators of ecosystem nitrogen (N) status are needed for monitoring and for identifying ecosystems that are at risk of

becoming N saturated. The N chemistry of a number of plant, soil and hydrologic components were analyzed to assess

the N status of mixed conifer forests across an N deposition gradient in the San Bernardino Mountains east of Los

Angeles, California. All of the measured parameters at Camp Paivika on the western high-deposition end of the gradient

indicated that the forest is N saturated, while forest stands on the eastern end of the gradient are N-limited. On the

basis of these and other studies, the following parameters are recommended as indicators for monitoring forest N

status: foliar nitrogen:phosphorus (N:P), foliar nitrate (NO3

-

), foliar growth response to N fertilization,

soil carbon:nitrogen (C:N) ratio, NO3

-

 in soil extracts or soil solution, and streamwater NO3

-

. However, foliar NO3

-

levels are more temporally sensitive than the other indicators mentioned, and foliar NO3

-

 levels vary widely among

plant species. Nitrate levels measured in KCl extracts or in soil saturation extracts were as effective as soil solution

NO3

-

 measurements for comparing soil NO3

-

 concentrations among sites across the N deposition gradient. Extractable

NO3

-

 may be the preferred indicator of soil NO3

-

 because it can be measured when soils are wet or dry, whereas soil

solution cannot easily be obtained from soils with low water content. Elevated N trace gas emissions from soil (nitric

oxide [NO] is dominant under the arid conditions of California) seem to be diagnostic of excess soil N, but field

measurement of N trace gases (particularly NO) is not routine for most laboratories. Streamwater NO3

-

 concentration

is a highly useful integrative indicator of the net outcome of N cycling processes within a watershed. Temporal trends

in streamwater NO3

-

 are indicative of the N retentiveness of the watershed. Studying a combination of suitable indicators

is recommended when evaluating forest N status.

Introduction

Indicators of ecosystem nitrogen (N) status are needed for monitoring and for

identifying ecosystems that are at risk of becoming N saturated. The need for

indicators is evidenced by the expanding geographic scope of forested areas over

which elevated atmospheric N inputs are expected (Galloway and others 1994).

Nitrogen deficiency or limitation is common in northern temperate forests (Vitousek

and Howarth 1991), and undisturbed forests are generally considered to be highly

conservative of N. However, many recent studies have shown that forests exposed

to chronic N deposition often result in nutrient cycling processes that have been

significantly altered by excessive inputs of atmospheric N. In many forests the levels

of available soil N are in excess of biotic demand, and N losses from these forests

can be high (Dise and Wright 1995, Riggan and others 1985). Such forests with

available N over and above the biotic and abiotic retention capacities are considered

N saturated (Aber and others 1989). Many current and recent studies have focused

on the consequences of N deposition and enrichment on natural ecosystems that

are adapted to N-limiting conditions. Water quality has deteriorated in many

watersheds exposed to N air pollution as a result of high NO3

-

concentrations in

streamwater. Although low-to-moderate N deposition may increase productivity,

considerable evidence suggests that detrimental environmental effects will ensue

in the long-term in forests with high levels of N inputs (Aber 1992).

Nitrogen deposition and ozone concentrations in the South Coast Air Basin

(SCAB; Los Angeles, California, and surrounding counties) are among the highest

in North America. Montane forests encompassing the SCAB are exposed to high

concentrations of ozone and N pollutants (Bytnerowicz and Fenn 1996). Nitrogen

and ozone air pollution are also on the rise in the western and southern portions

of the extensive Sierra Nevada range, as a result of large urban and agricultural

emission sources throughout California (Blanchard and others 1996). The effects

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Plant Pathologist and Soil Microbiologist, respectively, Pacific Southwest Research Station, USDA Forest Service,

4955 Canyon Crest Drive, Riverside,

CA 92507, U.S.A.

Indicators of Nitrogen Status in California Forests1

Mark E. Fenn and Mark A. Poth2
124 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

of N deposition on forests within the SCAB may provide a preview of future

scenarios in other western forests as urbanization and fossil fuel consumption

continues to increase in western North America. The more extreme cases of air

pollution exposure increase the likelihood that signals of air pollution effects will

be more clearly discernible above the background variation inherent in natural

ecosystems (Lang and others 1982).

In the San Bernardino Mountains (SBM) east of Los Angeles, California, a steep

air pollution gradient spanning 55 km has been identified, with ozone and N

deposition decreasing from west to east (Fenn and Bytnerowicz 1993, Miller and

others 1986). Ozone injury to sensitive species has been observed for many years,

while more recently the effects of chronic N deposition have also been documented

(Fenn and others 1996). A number of nutrient pools and processes have been

monitored at sites with high and low N deposition in the SBM to determine the

impact of atmospheric N inputs on N cycling.

This paper reviews which nutrient cycling parameters may be useful as

indicators of forest N status under the environmental conditions prevalent in California.

Selection Criteria for Indicators

of Forest Nitrogen Status

Evaluating forest N status is challenging because N is a major constituent of all

life forms and is cycled through a complex web of processes involving many biotic

and abiotic mechanisms. Monitoring toxic pollutants or xenobiotics such as heavy

metals or pesticides is in many ways a much simpler task. To evaluate the N status

of a forest ecosystem, we recommend the use of a small set of reliable indicators

that are easy to implement. Although numerous components of the ecosystem

could be measured to provide information on forest N status in intensive research

studies, the selection of indicators may be limited by practical considerations in

monitoring programs. Suggested criteria for selecting suitable indicators for

monitoring include the following:

● Accurate measure of site N status

● Lack redundancy

● Lack sensitivity to time of sampling or other environmental factors

● Sufficiently high signal-to-noise ratio

● Relatively standard sampling and analytical methods

● Cost effective (low cost/high information value)

● Definable range of “normal” values

● Clearly identifiable N excess and N deficiency.

Although selected indicators must accurately indicate site N status, evaluations

of forest N status may be difficult to establish based on any one indicator. Ecosystem

N status should be evaluated with a complementary set of nonredundant indicators

so that the relative degree of N limitation, sufficiency, or saturation (excess) may

be more firmly established. Vegetation-based indicators of N status are particularly

useful because they function as biomonitors and allow site N status to be assessed

by plant-environment interactions. Vegetation-based indicators help ensure that

interpretations of site N status are biologically relevant. For some indicators,

seasonal trends should be monitored to interpret indicator responses.

For sites at risk of N saturation, indicators should be included that can identify

when available N exceeds biotic demand. Such indicators typically monitor

parameters that are normally at background or low levels in N-limited systems.

Sustained high activity levels or concentrations suggest that available N is in excess

of biotic demand. Examples of indicators from the literature and from our studies

(Fenn and others 1996) that seem to indicate excess plant-available N include foliar

accumulation of nitrate (NO3

-

) (Stams and Schipholt 1990) or amino acids (Ohlson

Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 125

and others 1995), elevated emissions of N trace gases from soil (Castro and others

1994, Sitaula and others 1995), high soil solution NO3

-

 accompanied by high leachate

losses of NO3

-

 (Aber and others 1989), and above normal NO3

-

 concentrations and

fluxes in streamwater (Riggan and others 1985, Stoddard 1994). High levels of

nitrate reductase activity (Norby and others 1989) may also be indicative of high

NO3

-

 availability or elevated atmospheric concentrations of nitric acid vapor

(HNO3). However, in the SBM, concomitant exposure to high ozone concentrations

may interfere with enzyme activity, confounding the use of nitrate reductase activity

as an indicator of excess N availability (Krywult and others [In press]).

However, the cardinal indicator or manifestation of N saturation in all

ecosystem types, including California forests and chaparral, is increased and

prolonged NO3

-

 loss below the main rooting zone and in streamwater (Dise and

Wright 1995). Changes in solution concentrations and fluxes as water passes

vertically through forest ecosystems in precipitation, throughfall and stemflow,

forest floor percolate, soil solution, surface runoff, and in streamflow have been

referred to as solution or water chemistry profiles (Binkley and others 1982,

Chorover and others 1994).

The N solution profile in a forest is highly indicative of the N status or degree

of N saturation. In N-limited forests, nearly all of the solution N is retained within

the vegetation or upper soil layers of the solution profile. However, in N-saturated

forests, excess N is transported through the canopy and soil layers and exported

mainly as leached NO3

-

 and nitrogenous trace gases (Aber 1992).

Recommended Indicators in Arid Western

Forests

Indicator selection in arid western forests is based on current understanding of N

cycling in temperate forest ecosystems and on studies from air pollution gradients

in the SBM (Fenn and others 1996) and San Gabriel Mountains (SGM) (Anderson

and others 1988, Kiefer 1995, Riggan and others 1985). Valuable indicators of forest

N status in the summer-dry conditions that predominate in much of western North

America include:

● Soil NO3

-

 (in soil solution or in soil extracts)

● Nitrate:ammonium (NO3

-

:NH4

+

) ratios in soil

● Soil carbon:nitrogen (C:N) ratios

● Nitric oxide (NO) emission levels from soil

● N-mineralization and nitrification rates

● Foliar nitrogen:phosphorus (N:P), C:N, and N:cation ratios

● Accumulation of NO3

-

 in foliage of understory and overstory species

● Plant response to N fertilization

● Streamwater NO3

-

 concentrations and fluxes

● Base saturation and soil pH.

Many of the indicators suggested for arid forests of the western United States

have also been demonstrated to be effective in more humid temperate forests as

well. However, indicator responses may differ depending on climate and ecosystem

characteristics. For example, emission levels of nitric oxide (NO) are good indicators

in western forests, while in more humid forests of the eastern U.S. or Europe, studies

have shown that nitrous oxide (N2O) emissions are much greater than NO emissions

(Aber and others 1989). Temporal dynamics of streamwater NO3

-

 fluxes may also

be different in Mediterranean ecosystems (Riggan and others 1985). Nitrate in the

soil solution can also build up to high concentrations during the dry season in

sites with low or moderate N deposition as soil drying results in low volumes of

soil solution with high ionic concentrations (Fenn and others 1996). As with all

indicators, data from low-deposition sites or other reference data should be

compared for interpreting indicator results.

Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
126 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Characteristics of Recommended Indicators

Soil Nitrate

In strongly N-limited systems, N cycling and plant uptake of N occurs mainly as

ammonium (NH4

+

) (Aber 1992). As N saturation develops, plant-available NO3

-

increases in soil. Thus, the NO3

-

:NH4

+

 ratio in soil may be a useful indicator of site

N status. Soil NO3

-

 but not soil NH4

+

 increased with greater N deposition in the

SGM (Kiefer 1995) and SBM (Fenn and others 1996). However, ponderosa pine

(Pinus ponderosa Dougl. ex Laws.) trees prefer NO3

-

 to NH4

+

 (Griffin and others

1995, Norton and Firestone 1996), contrary to the conventional belief that conifers

prefer NH4

+

 (Adams and Attiwill 1982, McFee and Stone 1968). Soil NO3

-

 and NH4

+

can be measured in soil solution or in soil extracts (e.g., saturation or KCl extracts).

Obtaining soil solution samples with the widely-used suction lysimeter method

(Fernandez and others 1995) from the coarse-textured soils typical for these

ecosystems can be very difficult during the dry summer growing season. We found

that the soil centrifugation method (Soon and Warren 1993) worked well for

obtaining soil solution samples from coarse-textured forest soils with water content

near field capacity. However, in our studies in the SBM, NO3

-

 levels measured in

soil saturation extracts and in KCl extracts were equally effective as soil solution

NO3

-

 measurements for comparing soil NO3

-

 concentrations at sites across the

deposition gradient. Soil extracts can be obtained from soils collected at any time,

regardless of soil water content or soil texture, and in that regard are probably

more practical as an indicator in a monitoring program. Another recent study

concluded that KCl extracts performed similarly to suction cup lysimeters in

quantifying soil NO3

-

 (Djurhuus and Jacobson 1995).

Streamwater Nitrate – a Watershed-Level Indicator

Streamwater ionic flux is a very useful integrator of overall ecosystem integrity

and serves as a watershed-level indicator of ecosystem nutrient loss or conservation.

Nitrogen saturated watersheds export above-normal amounts of NO3

-

 once the

capacity for N retention by the biota and soil chemical fixation mechanisms are

exceeded (Stoddard 1994). Temporal or seasonal trends in NO3

-

 export in

streamwater reveal the N retention capacity at the watershed scale and are a

consequence of the many nutrient cycling processes occurring within the

watershed. A brief and relatively small NO3

-

 loss from temperate forest watersheds

is expected during the spring water flow from the watershed brought on by

snowmelt. During this early spring season water flows are greatest and soil

temperatures are still low, and plant and microbial nutrient demand are insufficient

to retain all of the dissolved N. However, a state of N saturation is indicated when

NO3- concentrations begin to increase over this “normal” background level and

when losses are prolonged. Peak flows in N-enriched California ecosystems seem

to occur during major winter storms or during subsequent snowmelt periods when

more complete washout of the soil profile occurs (Riggan and others 1985).

Monitoring streamwater NO3

-

 dynamics throughout the hydrologic year will

provide the most information on watershed N processing and losses. However,

with an understanding of the temporal dynamics of streamwater chemistry within

a given set of environmental conditions and ecotypes, sampling windows may be

identified when NO3

-

 in streamwater is indicative of above-normal N losses or N

saturation. For example, high streamwater NO3

-

 concentrations during the summer

when the terrestrial and aquatic biota are normally capable of retaining available

N may suggest an N-saturated condition. When the objective is to compare annual

N inputs and outputs for a watershed, streamwater flow volumes and NO3

-

concentrations should be measured.

Soil C:N Ratio

The soil C:N ratio is a useful indicator of N enrichment in forests. Soil C:N ratio is

less temporally and environmentally sensitive than other indicators, such as NO3

-

Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 127

levels in soil and foliage, nitrate reductase activity, amino acid concentrations in

foliage, trace gas emissions from soil, and streamwater NO3

-

 concentrations. The

relative stability of soil C:N enhances the utility of this indicator and reduces the

odds of fluctuating environmental conditions or nutrient cycling processes

confounding interpretation of indicator results. However, the fire history of

monitored sites must be considered when implementing any N status indicator

(Chorover and others 1994), including C:N ratio. Percent C and N losses from

burned plant material are highly similar (Raison and others 1985), and soil C:N

ratios appear to be little changed after fire (Carreira and others 1994, Raison 1979).

However, total C and N pools can change dramatically in burned sites (Raison

1979), suggesting the need for caution when monitoring sites have burned recently.

Foliar Nitrate

Nitrate is stored in the vacuole (Granstedt and Huffaker 1982) but does not

accumulate in foliage until the growth requirements for N are satisfied (Zhen and

Leigh 1990) and the capacity for NO3

-

 assimilation is exceeded (Lee and others

1986). Nitrate accumulation in plants can be a very useful indicator of N excess

because NO3

-

 does not accumulate in foliage unless plant demand for N has been

met. Nitrate concentrations in foliage of pine, oak and fern were much higher at

an N-saturated site compared to an N-limited site in the SBM (Fenn and others

1996). Nitrate accumulation was especially high in bracken fern (Pteridium aquilinum

var. pubescens Underw.), a species that has been shown to accumulate NO3

-

 under

conditions of high N deposition (Stams and Schipholt 1990).

Amino acid levels in foliage may also be an effective indicator of N status or

storage of excess available N (Ericsson and others 1993, Ohlson and others 1995),

but this has not yet been investigated in western forests. However, amino acid

analysis is not likely to be routine for many plant and soil analysis laboratories.

Foliar Nutrient Ratios

Foliar nutrient ratios are often more reliable and informative indicators of plant

nutrient status than nutrient concentration alone (Ericsson and others 1993, Hüttl

1990, Mohren 1986, Zinke 1980). In three separate studies, foliar N:P ratio has been

found to increase with increasing N deposition in the mountains of southern

California (Fenn and others 1996, Poth and others 1991, Zinke 1980). Foliar C:N

ratio is also a useful indicator of site N status (Fenn and others 1996).

Nitrogen:cation ratios in foliage have not been sufficiently tested as N status

indicators in California but may also be useful indicators of N status or of nutrient

imbalances (Hüttl 1990).

Elaborate nutritional diagnosis systems have been developed based on a suite

of nutrient ratios. The Diagnosis and Recommendation Integrated System (DRIS)

is an example of a widely used system in agriculture (Walworth and Sumner 1987),

which has also been applied to a number of forest species (Binkley and others

1995, Riiters and others 1992, Shumway and Chappell 1995). DRIS uses nutrient

ratios in high-productivity sites as a guide to optimum ratios that could be achieved

through fertilization of less productive sites. Before DRIS can be used for a particular

plant species, however, DRIS “norms” have to be determined for that species based

on nutrient data from a large survey of the species in question (Needham and

others 1990). DRIS norms have not been developed for ponderosa pine or most

other western forest species; but if they are developed, DRIS could prove to be a

very effective tool for evaluating site N status and for identifying other nutrients

that may be limiting or in excess. This could be particularly useful in cases where

excess N may induce deficiencies of other nutrients (Houdijk and Roelofs 1993,

Mohren and others 1986, Schulze 1989).

Plant Response to Fertilization

Studies of plant response to N amendment treatments is an effective technique for

evaluating site N status, although this requires treatment application and

Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
128 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

subsequent comparisons of untreated control plants and fertilizer-treated plants.

Vector analysis is a simple, but elegant, graphical technique by which tree N status

can be determined from physiological responses to fertilization under field

conditions (Haase and Rose 1995, Timmer and Stone 1978, Weetman and Fournier

1982). Fertilization of individual trees with N before the growing season followed

by end-of-season analysis of tree response in terms of foliar biomass, N content,

and N concentration (vector analysis) can provide clear indications of the N status

of forest trees. The value of the fertilization/vector analysis method, however, is

that field grown trees can be used to biomonitor and interpret site N status — at

least for the selected tree species. Vector analysis studies in the SBM and SGM

suggest that ponderosa and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) trees in stands

with N deposition from ca. 6 to 16 kg ha-1 yr-1 are N deficient, while N was not

limiting at sites with more than 30 kg ha-1 yr-1, (Kiefer and Fenn [In press]).

Nitrogen Mineralization and Nitrification

In N-enriched sites, N-mineralization and nitrification rates are usually increased

(Aber and others 1989, Van Miegroet and others 1992). Under conditions of excess

available N, the need for N conservation is reduced, and canopy N turnover, net

N-mineralization and net nitrification increase. Nitrogen mineralization can be

measured in situ with the buried bag technique (Pastor and others 1984), or

potential N-mineralization and nitrification can be assayed under controlled

conditions. Both aerobic and anaerobic assays can be used, although nitrification

rates are not determined in anaerobic incubations (Binkley and Hart 1989). The

laboratory assays provide information on potential net N-mineralization and

nitrification rates, which are measures of the soil capacity to supply N, while

eliminating confounding environmental influences on soil N cycling. Assays

performed under standard conditions allow for comparisons of soil N status for

different sites and remove site environmental constraints (e.g., temperature and

moisture limitations) that are constantly changing. N mineralization rates were

higher in sites with greater N deposition in the SBM and SGM in southern California

(Fenn and others 1996, Kiefer 1995).

Fluxes of N Trace Gases

Fluxes of trace gases from soil often increase with greater soil N supply (Castro

and others 1994, Sitaula and others 1995). In forests in eastern North America and

in Europe, N2O emissions appear to be much greater than NO emissions (Aber

1992), although NO fluxes are frequently not measured. In California soils, NO

fluxes are typically much greater than N2O fluxes (Anderson and others 1988,

Davidson 1993). NO emissions were much higher in a high deposition site

compared to a low deposition site in the SBM (Fenn and others 1996). Nitric oxide

appears to be diagnostic of excess available soil N in western coarse-textured soils.

A major drawback of NO as an indicator of soil N status is the specialized

equipment required to measure NO. Under conditions where N2O emissions are

sufficiently high (finer textured soils with more than 60 percent water-filled pore

space) (Davidson 1993), simple flux chambers could be installed on the forest floor

and headspace samples collected, transported to the laboratory, and N2O quantified

via gas chromatography. Although N trace gas fluxes seem to be indicative of soil

N status, other indicators may be easier to implement for the general plant and

soil chemical analysis laboratory.

Soil pH and Base Saturation

Decreased base saturation and soil pH in forests with high N inputs does not prove

cause and effect because other stand development factors can also cause soil

acidification and decreased base cation levels in soil. However, if other indicators

provide evidence of N saturation, decreasing base saturation and soil pH may

provide supporting data of the effects of N enrichment. Decreased base saturation

Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 129

is a result of cation leaching as counter-balancing ions for leached NO 3 

-

. In the SBM

soils in sites with high N deposition had lower base saturation and pH than soil in

sites with low N deposition (Fenn and others 1996).

Summary

To assess the N status of a forest ecosystem several reliable and complementary

indicators should be selected. Under the arid conditions of much of the western

U.S., recommended indicators include the following:

● NO3

-

 concentrations in soil and foliage

● Foliar nutrient ratios such as N:P

● Soil C:N

● Streamwater NO3

-

 concentrations and fluxes

● Plant response to N fertilization or vector analysis.

The N status of a forest ecosystem can also be assessed in the context of solution

chemistry profiles that consider the sequential processing of N in precipitation,

throughfall and stemflow, forest floor percolate, soil solution, surface runoff, and

streamflow. Nitrogen mineralization and nitrification rates and trace gas fluxes

from soil (especially NO) are also potentially useful indicators of N status. With

further research, amino acid levels in foliage and the Diagnosis and

Recommendation Integrated System (DRIS) method could also be used in

evaluating site N status. However, DRIS norms would need to be developed for

the particular plant species.

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Session I1 Indicators of Nitrogen Status in California Forests Fenn, Poth
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 131

Abstract

The coastal sage scrub (CSS) vegetation of southern California has been declining in land area and in shrub density

over the past 60 years or more, and is being replaced by Mediterranean annual grasses in many areas. Although much

of this loss is attributable to agriculture, grazing, urbanization and frequent fire, even protected areas have experienced

a loss in native shrub cover. Nitrogen (N) deposition has not previously been examined as a contributor to CSS decline,

but up to 45 kg/ha/yr are deposited in the Los Angeles Air Basin. Several mechanisms were examined by which

atmospheric N deposition might affect the shrubs and promote growth of weeds. Field nitrogen fertilization studies at

sites of high and low deposition showed that most of the abundant native and introduced species had increased growth

after fertilization in the low deposition site, but in a high deposition site only one weedy species, small-podded mustard

(Brassica geniculata (Desf.) J. Ball), responded to N fertilization. Greenhouse studies showed that both shrubs and

weeds had high plasticity in their growth response to N fertilizer, an unexpected result for the shrubs. Preliminary

competition studies indicated there was no change in the relative competitive ability of the shrubs or grasses after

fertilization. However, negative effects of high N have been detected on the growth and survival of the shrubs. Greenhouse

grown California sagebrush (Artemisia californica Less.) began to senesce at 6 to 9 months when fertilized with 50

μg N/g soil. This soil N concentration corresponds to extractable N levels in polluted sites, while levels are typically

less than 10 μg/g in unpolluted sites. Another source of damage to plants can be cuticular lesions caused by nitric

acid, but how nitric acid affects CSS leaves is unknown. Fumigation of pine needles with high ambient levels of nitric

acid caused cuticular lesions and stomatal collapse, as well as modifications to nitrogen assimilation pathways. The

preliminary evidence suggests that CSS vegetation may decline due to elevated nitrate levels in the soil, and additional

studies are need to test effects of ambient nitric acid on CSS leaves.

Introduction

The coastal sage scrub (CSS) vegetation of southern California has been declining

in land area and in shrub density over the past 60 years or more, and is being

replaced by Mediterranean annual grasses (Davis 1994, Freudenberger and others

1987, Minnich and Dezzani [In press], O’Leary and others 1992, Zink and others

1995). Although much of this loss is attributable to agriculture, grazing, urbanization

and frequent fire, even protected areas have experienced a substantial loss in native

shrub cover (Minnich and Dezzani [In press]). Nitrogen (N) deposition has not

previously been examined as a contributor to CSS decline, but up to 45 kg/ha/yr

are deposited in the Los Angeles Air Basin (Bytnerowicz and Fenn 1996).

Nitrogenous compounds in polluted air affect even those tracts of land that have

been set aside as reserves, which are in many cases surrounded by urbanization in

southern California (O’Leary and others 1992). The coastal sage scrub is of particular

interest to conservationists because it supports some 200 sensitive plant species

and several federally listed animal species (DiSimone 1995, O’Leary 1989). We need

to understand how to manage lands that are influenced by urban air pollution, and

whether the biotic communities of these lands can be preserved.

Nitrogen deposition is known to cause vegetation type conversions in other

countries, notably the Netherlands (Bobbink and Willems 1987), which have up to

85 kg/ha/yr of N deposited, the highest measured in the world. Nitrogen

deposition to reserves is a conservation problem in the Netherlands, where

heathlands and species-rich pastures are turning into species-poor grasslands

(Asman and others 1989, Bobbink and Willems 1987). The high rates of N deposition

in southern California have caused increased soil fertility and surface litter

decomposition rates in mixed conifer forests (Fenn 1991), but less is known about

Nitrogen Deposition Effects on Coastal Sage Vegetation of

Southern California1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Natural Resources Extension Specialist

and Associate in the Agriculture

Experiment Station, Department of

Botany and Plant Sciences, University of

California, Riverside, CA 92521-0124.

E-mail: edith.allen@ucr.edu,ppadgett

3 Forest Ecologist, Pacific Southwest

Research Station, USDA Forest Service,

4955 Canyon Crest Ave., Riverside, CA

92507. E-mail: andrzej@deltanet.com

4 Associate Professor, Department of

Earth Sciences, University of California,

Riverside, CA 92521.E-mail:

richard.minnich@ucr.ed

Edith B. Allen,2 Pamela E. Padgett,2 Andrzej Bytnerowicz,3 Richard Minnich4
132 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

vegetation change with elevated N deposition in this region. The coniferous forests

of the San Bernardino Mountains exhibit symptoms of N saturation, such as high

N:P ratios in leaf tissue, high soil N, high rates of N loss from the ecosystem, and

others (Fenn and others [In press]). If the relatively productive forests of the Los

Angeles Air Basin are N-saturated, then less productive vegetation such as CSS

that is receiving similar levels of deposition would also surely be saturated. Less

productive vegetation may become N saturated more rapidly than highly

productive vegetation because the ratio of deposited N to plant biomass is greater,

as suggested by Aber and others (1989, 1992) concept of anthropogenic nitrogen

saturation. In addition, long-lived trees may respond less rapidly than shorterlived shrubs, so we may expect to see a more rapid response in vegetation change

in CSS than in nearby forests.

Here we examine the mechanisms by which deposited nitrogen might cause

CSS shrubs to be replaced by Mediterranean annual grasses. We present three

hypotheses with preliminary data to explain how nitrogen deposition may affect

CSS species. Stated in the null form they are 1) CSS shrubs are equally plastic in

their growth response to N as are introduced grasses, 2) CSS shrubs are equally

competitive as the grasses after N fertilization, and 3) CSS shrub growth and

mortality are not affected by high N levels.

Effects of N Deposition on

Coastal Sage Scrub Vegetation

We have done series of field and greenhouse experiments on the effects of N

deposition on CSS species. The field experiments were done along a gradient of

atmospheric N concentrations, using one site with relatively high and another with

relatively low N air concentrations for intensive measurements (fig. 1). The high

deposition site is at Box Springs Mountain near the University of California,

Riverside at an elevation of 670 m, and the low deposition site is some 60 km to the

south at the Lake Skinner Reserve, elevation 540 m. Longterm climate data are not

available at either site, but the city of Riverside, adjacent to Box Springs Mountain,

receives 280 mm precipitation annually. We are still working out the actual

deposition rates, which are likely lower than the estimated high values of 30 kg

N/ha and more recently 45 kg N/ha at Camp Paivika on the western end of the

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich

Figure 1 — Sample sites on the

Perris Plain referenced in the text.

The site of highest measured N

deposition is Camp Paivika (10)

and the site with lowest air

concentrations is (9) at Mission

Trails Park in San Diego. Arrows

show air flow patterns, with

polluted air flowing inland from

Los Angeles and cleaner air

flowing inland from other coastal

sites. A convergence zone of

polluted and clean air occurs in

the middle of the Perris Plain.

1 = Jurupa Hills,

2 = Mockingbird Reservoir,

3 = Box Springs Mountain,

4 = Lake Matthews,

5 = Motte Rimrock Reserve,

6 = Simpson Park in Hemet,

7 = Lake Skinner,

8 = Santa Margarita Ecological

Reserve,

9 = Mission Trails Park in San

Diego,

10 = Camp Paivika, San

Bernardino Mountains.

Mójave Desert
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 133

San Bernardino Mountains (Bytnerowicz and Fenn 1996, Fenn and Bytnerowicz

1993). Peak air concentrations were measured with an annular denuder system in

August, 1994 as 31.0 ␣μg/m3 NO3

-

 plus 8.6 ␣μg/m3 NH4

+ at Box Springs Mt., and 13.9

␣μg/m3 NO3

-

 and 3.9 ␣μg/m3 NH4

+ at the Lake Skinner Reserve (fig. 2). Sulfur was

relatively low across the gradient, as is the case in other western air pollution

measurements (Bytnerowicz and others 1987). The vegetation at both sites is CSS,

with a higher proportion of introduced grasses on Box Springs Mountain than at

Lake Skinner (Minnich and Dezzani [In press]). Both sites are on granitic soils.

Additional sites were used for less intensive measurements.

In addition to air measurements, soil samples were taken along the gradient

of pollution that also included sites to the northwest of Box Springs Mountain

(fig. 3). The top 2 cm of soils of the Jurupa Hills had as much as 86␣μg/g of

extractable N in the form of nitrate plus ammonium (fig. 3), with nitrate

predominating at the more polluted sites. Box Springs Mountain had 44␣μg/g,

and Lake Skinner had 8␣μg/g of extractable N. These soils were collected in

September during the dry season, when soil N accumulates because plants have

senesced and are no longer taking up N. The soil N measurements confirm the

gradient of N pollution by showing that the soils also accumulate N. We do not

yet know if the soil N is higher because of accumulated N deposition, or because

of increased mineralization that is induced by N deposition (Fenn and Dunn 1989).

However, such high concentrations of N in the soil are likely to affect the plant

community, which at each of these sites is CSS with an understory of annual

grasses. At some sites, such as Box Springs Mountain and the Jurupa Hills, the

annual grasses have become dominant with interspersed patches of shrubs.

We explore the mechanisms to explain how the vegetation may change from

shrubland to grassland after N deposition.

Plasticity Hypothesis

For one species to replace another in a high N soil, it must have a greater response

to N, or in other words be more plastic in response to changes in resources (Jennings

and Trewavas 1986). To test this hypothesis, we did N fertilization experiments in

the field and the greenhouse. In the field, we fertilized plots at the Box Springs

Mountain and the Lake Skinner sites. Both sites had burned in November 1993,

and the fertilizer treatments included both burned and unburned vegetation. Each

site had ten 5 by 5 m2

 plots and N was applied at a rate of 60 kg N/ha as NH4NO3

in two doses of 30 kg/ha each in February and March, 1994. Plant response was

Figure 2 — Nitrogen (NO3

- -N

and NH4

+ -N) and total S (SO42--

S) concentrations in the

atmosphere at five sites

representing a gradient of air

pollution from Box Springs

Mountain in Riverside and

southward to San Diego during

August 1994. Location of sites is

shown in figure 1.

45

40

35

30

25

20

15

10

5

0

Box

Springs

Mt.

Motte

Reserve

Hemet Lake

Skinner

San

Diego

County

Total N

Total S

August 1994 Intensive Study

Total (

μg/m3

)

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
134 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

evaluated in May using non-destructive percent cover data. The unburned plots

did not respond to fertilizer at either site with one exception described below, so

results of fertilization trials are shown only from the burned plots (table 1).

At Box Springs Mountain only one introduced forb, small-podded mustard

(Brassica geniculata [Desf.] J. Ball), responded significantly to N fertilizer on the

burned plots, and this was also the only species that responded on the unburned

plots (data not shown, but small-podded mustard increased from 17 to 29 percent

after N fertilization on unburned plots). None of the other introduced grasses and

none of the native species responded significantly at Box Springs Mountain in the

burned plots (table 1). The introduced grasses included species in the genera wild

oats (Avena), brome (Bromus), fescue (Vulpia) and split grass (Schismus), while the

native species included a diverse mixture of some 70 species.5

 By contrast, at Lake

Skinner almost all of the categories of species, both native and introduced,

responded to fertilizer, with the exception of the introduced forbs (table 1). Smallpodded mustard occurred infrequently at Lake Skinner, with < 5 percent cover.

The total percent vegetative cover was greater in fertilized plots at both sites, but

was higher overall at Box Springs Mountain. These results suggest that the plants

are N deficient at Lake Skinner, and N saturated at Box Springs Mountain, with

the exception of small-podded mustard, which continued to grow and take up N

after fertilization. Leaf tissue N of small-podded mustard increased from 3.2 percent

to 4.2 percent after fertilization. This suggests that small-podded mustard may be

one of the “winners” in the high N deposition zones, as it can take advantage of

additional N, even when soil N is already high. The results from the Lake Skinner

site suggest that both native and introduced species are plastic in their responses

to N, whereas our original hypothesis was that only the introduced species would

be highly responsive to N, or “nitrophilous.”

The responses in the field may have been influenced by a number of factors,

especially competition in a complex community. To understand the responses of

native and weedy plants under more controlled conditions, we performed

greenhouse N fertilizer trials of monocultures of three introduced annuals (wild

oats [Avena fatua L.], red brome [Bromus rubens L.], and small-podded mustard),

and three native shrubs (California sagebrush (Artemisia californica Less.), brittlebush (Encelia farinosa Gray.) and California buckwheat (Eriogonum fasciculatum

Benth.). The plants were grown in 3.5l pots in native soil amended with 0, 10, 50

and 100 ␣μg/g of N as NH4

NO3. The soil was collected from the Motte Reserve

after scraping off the top 5 cm of soil. After 4 month’s growth in the greenhouse,

the native shrubs were just as plastic in their responses to N as were the introduced

Figure 3 — Extractable soil

nitrogen (NO3

-

 -N and NH4

+

 -N)

from sites on a nitrogen gradient

in September 1995. Soil cores

were divided into the upper 2 cm

and lower 2 to 5 cm. Location of

sites is shown in figure 1.

5 Unpublished data on file, Department

of Botany and Plant Sciences, University of California, Riverside.

100

90

80

70

60

50

40

30

20

10

0

Jurupa

0 - 2 cm

2 - 5 cm

Soil Depth

Total Inorganic Nitrogen (

μg-N/g soil)

Mockingbird

Reservoir 

Box Springs Mt.

Lake Matthews

Motte Reserve

Hemet

Santa Margarita

Reserve 

Lake Skinner

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 135

annuals (figs. 4 and 5). In fact, the annuals tended to saturate at 50 μg/g, while the

shrubs had continued increased growth to 100 μg/g. This was an unexpected result,

as native wildland species are typically thought to have low responses to nutrient

additions compared to weeds and crop plants (Chapin 1980). Clearly the native

shrubs are adapted to rapid growth during the brief 4 to 6 month rainy period, at

which time they likely take up nutrients as rapidly as possible. The growth period

is brief in Mediterranean climates, which have a moist winter and spring but the

other seasons are dry. However, in an unpolluted situation, the loading of available

N in the soil would not be as great as after N deposition. Our field observations

showed that available N built up as high as 86 μg/g in the upper 2 cm of soil (fig.

3) during the dry season, an amount that would be available to newly growing

seedlings at the beginning of the next rainy season. Both the native shrubs and the

introduced grasses are apparently able to take advantage of this high soil N.

These results were unexpected and not only confound our notions of how

plants behave in the wild (the paradigm states they should have slow rates of

nutrient uptake and growth), the results also do not explain why N deposition

would shift the vegetation from a shrub-dominated to a grass-dominated type.

An alternative hypothesis is that competition for N occurs between the grasses

and shrubs that is skewed in favor of the grasses.

Figure 5 — Relative shoot dry weight of three introduced weed

species subjected to levels of 0, 10, 50, and 100 μg/g N as NH4

NO3

in the soil after 4 months of growth.

Figure 4 — Relative shoot dry weight of three native shrub species

subjected to levels of 0, 10, 50, and 100 μg/g N as NH4

NO3 in the

soil after 4 months of growth. In the case of brittle- bush (Encelia

farinosa), N was added either as NH4Cl or as Ca(NO3)2.

100

80

60

40

20

0

–

-

–

-

–

-

–

-

–

-

-–-–-–-–-–

–

-

–

-

–

-

–

-

–

-

-–-–-–-–-–

0 20 40 60 80 100

Introduced Species

 Brassica geniculata

 Bromus rubens

 Avena fatua

Relative Shoot Dry Weight

Soil Inorganic Nitrogen (ppm)

100

80

60

40

20

0

–

-

–

-

–

-

–

-

–

-

-–-–-–-–-–

–

-

–

-

–

-

–

-

–

-

-–-–-–-–-–

0 20 40 60 80 100

Native Species

 Artemisia californica

 Eriogonum fasculatum

 Encelia farinosa - NH4

 Encelia farinosa - NO3

Relative Shoot Dry Weight

Soil Inorganic Nitrogen (ppm)

1 Significantly different using a t-test at

P < 0.05.

2 n.s.=not significantly different.

Table 1 Table 1 — Percent cover of vegetation in nitrogen fertilized and unfertilized treatments in burned plots at the high N (Box Table 1 

Springs) and low N (Lake Skinner) deposition sites.

BOX SPRINGS LAKE SKINNER

 Species N Fert. No N N Fert. No N

Brassica geniculata 17.1 9.0 ( 1 ) 0.0 0.0 n.s.

Bromus rubens 15.6 13.5 n.s.2 4.5 2.0 P = 0.07

Other exotic grasses 20.1 14.7 n.s. 10.5 4.2 ( 1 )

Other exotic forbs 27.5 26.3 n.s. 19.8 18.9 n.s.

Native forbs 20.7 19.2 n.s. 37.5 22.7 ( 1 )

Native shrubs 0.0 0.1 n.s. 11.1 4.9 (1 )

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
136 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Competition Hypothesis

To determine whether N shifts the competitive balance between the grasses and

shrubs, we initiated a competition experiment in a patch of introduced grassland

that was once dominated by CSS species. This research was done at the Motte

Rimrock Reserve about 24 km south of Riverside and intermediate in nitrogen

deposition to the Box Springs and Lake Skinner sites. In a blocked experimental

design we weeded grasses from ten, 1.2 m2 plots and left an additional ten plots as

controls dominated by the introduced grasses red brome and foxtail fescue (Vulpia

megalura Rydb.) A few introduced forbs were also present, mainly species of

storksbill (Erodium), but few native species. One-half of the cleared plots and onehalf of the grassy plots were fertilized with two doses of NH4NO3 at the rate of 30

kg N/ha each time in spring 1995. Nine seedlings of California sagebrush were

planted in each of the plots in a 2 by 2 factorial design to test two levels of N (0 and

60 kg/ha) and two levels of competition (with and without grasses) on the growth

of California sagebrush. The seedlings were spaced 30 cm apart, so they did not

interfere in aboveground growth during the first growing season. Because we did

not wish to harvest seedlings during the first year, we did non-destructive

measurements of height and width to calculate shrub volume. Volume was

calculated by assuming that the shape of California sagebrush is spheroid.

The grass competition proved overwhelming for California sagebrush, which

had only about one-ninth the volume in the grass plots compared to the cleared

plots, with or without N fertilizer (fig. 6). Nitrogen did not shift the balance of

competition in favor or against California sagebrush, it only increased the growth

of California sagebrush with and without grass competition. Increased growth of

California sagebrush after fertilization should be a benefit, if this means that it can

survive competition from the grasses. But in fact most of the California sagebrush

seedlings in grass plots died during the normal summer drought that followed

these measurements, while many survived in the cleared plots. With or without N

the seedlings that experienced grass competition were still tiny, and apparently

not large enough, or with roots too shallow, to survive the summer drought. On

the basis of this experiment, we cannot expect that areas with N deposition will

have greater shrub seedling survival. In fact, we noted mortality of mature shrubs

in areas of high N deposition, and proceeded to examine potential negative effects

of N on shrub growth.

Figure 6 — Volume measurements of California sagebrush

(Artemisia californica) in weeded

(monoculture) and grassy

(mixture) plots in the field after

3 months of growth, fertilized

with 60 kg/ha N as NH4

NO3 or

unfertilized.

––

Monoculture

Mixture

Fertilized Unfertilized

Artemisia volume (cm

3) 500

400

300

200

100

0

–

–

–

–

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 137

Negative Plant Response to Nitrogen Hypothesis

Because the plasticity and competition hypotheses did not explain why N might

cause CSS decline, we continued to test alternative hypotheses. Shrubs in pollutionimpacted urban areas have been dying, as can be seen by a walk through Box

Springs Mountain County Park and other local reserves. Some 10 percent of the

shrubs in the polluted areas that we marked for experimental purposes died during

two growing seasons, but we did not experience shrub loss at the Lake Skinner

site. We do not yet have an estimate of the rate of mortality on a larger scale, but

Minnich and Dezzani [In press] have shown up to a 90 percent loss in shrub cover

since the 1930’s in urban areas of the Perris Plain (Riverside, California and

southward). Shrub loss is lower in the more rural southern Perris Plain. They did

their analyses from historic plots in CSS collected by the USDA Forest Service in

the 1930’s, which they resampled in 1993. Although they showed an increasing

pattern of shrub loss toward the urban areas, some factors confounded the results,

such as changes in soil type and CSS stand age since the most recent fire at the

time of sampling.

Wellburn (1990) explained that N deposition may more frequently harm

than fertilize plants, especially in the form of N oxides. We did a series of

experiments to determine if elevated N levels might have negative effects on

the shrubs. One was a greenhouse fertilization experiment with California

sagebrush, where the seedlings were fertilized with 50␣μg N/g as NH4

NO3.

Seeds were planted in a soil of low N, and after they were 3 to 4 months old

they were fertilized with the 50␣μg N/g soil. They were then fertilized every 2

months to maintain high soil N. During the first 6 months the seedlings grew

rapidly, as was noted above. Senescence of individual branches at 6 months,

and then complete mortality of all seven replicate plants between 6 and 9

months occurred. At 9 months, and even at 1 year, all of the plants that received

low N fertilizer levels were still healthy. These negative effects are difficult to

explain, as they appear to be a toxic effect. The soil N we maintained in the

greenhouse was not as high as the highest levels we measured in the field, up

to 86␣μg/g in soils of polluted sites during the dry season. The greenhouse

experiment was designed to maintain a concentration of 50␣μg/g in the soil

during the experiment, with two to three fertilizer additions during the 6 to 9

months. In the field, such a high level would build during the dry season and

be maintained only until the beginning of the next rainy season. When

vegetation begins to grow again in response to rain, the plants take up the

available soil N, and our measurements showed that extractable soil N dropped

to 10 ␣μg/g even in the most polluted sites.6 Leaching and denitrification might

also remove some of the deposited N from the soil, although both of these are

probably minor components of the N cycle in this semiarid shrubland. Such

high levels of soil N were not maintained throughout the year, so high soil N

would not cause as high a rate of shrub mortality in the field as in the

greenhouse. But these results do suggest that, over time, high soil N may be a

cause of shrub mortality.

The mechanisms by which shrub mortality occurred under high soil N is

not known. Horticulturists have long known that native species from soils of

low fertility will have short lifespans in a fertile garden, and advise that low

fertilizer levels be used for native California shrubs (Keator 1994). We can

discuss a number of alternative explanations for the adverse response of native

shrubs to high N. It is likely that California sagebrush was taking up large

quantities of N, because it has evolved only to take up this limiting nutrient in

the soils, not to exclude it. The metabolic requirements for maintaining and

detoxifying N in the tissues (NH4

+ is toxic in high concentrations) may be

expensive, and may lead to a shortened lifespan. Assimilation of NH4

+ requires

that carbon be shunted away from sugar synthesis into amino acid synthesis.

This would leave a plant deficient in carbohydrates and consequently energy

6 Unpublished data on file, Department

of Botany and Plant Sciences, University of California, Riverside.

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
138 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

to carry out other biochemical functions. We plan to pursue research on this

question, and currently are testing other species of CSS shrubs to determine if

this direct effect is generalizable.

Atmospheric N not only causes increased soil N, it results in exposure of

vegetation to elevated levels of gaseous and particulate forms of N that may

be deposited on leaf surfaces and interact directly with them (Krywult and

others 1996). Fumigation experiments have not yet been carried out on CSS

species, but they have been done on ponderosa pine (Pinus ponderosa Dougl. ex

Laws.) and black oak (Quercus kelloggii Newb.). Fumigation with ambient peak

summer levels of nitric acid (50 μg/m3) caused cuticular lesions and stomatal

collapse. These adverse responses by leaf surfaces could result in reduced

stomatal control and increased exposure of the leaf to other stresses. In addition,

nitric acid fumigation caused induction of nitrate reductase and increases in

amino acid levels in leaf tissue (Padgett and others 1995). These negative effects

can only be postulated for CSS species until similar research is done. Because

up to 90 percent of the leaves of CSS shrubs senesce in the summer, there are

fewer opportunities for foliar interaction with airborne N at peak summer

pollution levels. The CSS plants would be subject to lower spring airborne nitric

acid levels.

Conclusions

The decline of CSS vegetation is caused by a combination of complex factors,

beginning with direct destruction of CSS for agriculture and urban construction,

possibly including past grazing, and also including increased fire frequency in

lands that are adjacent to urban areas. But even when these effects are held

constant by examining CSS natural reserves that have not been impacted, these

shrublands are still declining. The lack of response of the vegetation at Box

Springs Mountain to additional N fertilization suggests that it is already N

saturated, one response that is cited by Aber and others (1989) as being an

indicator of N saturation. However, two forms of air pollution, ozone and N

compounds, are likely both central players in the Los Angeles Air Basin and

the Perris Plain. We have not discussed ozone in this paper, although it may

also play a role in CSS decline. Ozone is known to increase the mortality of

conifers in the local mountain ranges (Bytnerowicz and Grulke 1992, Miller

and others 1963). Westman (1990) hypothesized that ozone may also be a cause

of CSS decline because it reduced the growth of well-watered seedlings in

greenhouse experiments at simulated ambient summer concentrations (Stolte

1982). However, during peak summer air pollution conditions stomates of CSS

shrubs are closed and many leaves have senesced. Spring ozone concentrations

are relatively low, so ozone effects in the field are likely smaller than greenhouse

experiments would suggest. The effects of ozone may not be as serious as

previously thought in CSS vegetation. However, until our work began, all of

the air pollution effects were attributed to ozone and none to N deposition on

this vegetation type.

Our studies showed that the annual grasses that replace CSS vegetation

are not more nitrophilous than the shrubs, as they have equal plasticity in

response to N. However, one species we have tested to date, California

sagebrush, suffers a 100 percent mortality rate in the greenhouse in fertilized

soils with available N at levels that are no higher than field soils in polluted

sites. We are currently conducting experiments with two additional shrub

species, brittle-bush and California buckwheat. We have not seen similar results

from studies on ozone that would show such a rapid mortality rate. Most studies

on ozone report only decreased growth rates at ambient concentrations, not

mortality. In addition, the study on toxic effects of nitric acid vapor on pine

and oak leaves suggests that more studies of deposited N on CSS shrubs are

Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 139

needed. Increased mortality of CSS shrubs in N-polluted areas may be a cause

of CSS decline in the Los Angeles Air Basin. CSS may be the first vegetation

type in the western United States that exhibits stage 3 symptoms of N saturation,

which is defined as toxic effects on the vegetation (Aber and others 1989).

Acknowledgments

We thank Shiela Kee, Lucia Vasquez, and Lidia Yoshida for field assistance. This

research was funded by National Science Foundation grant no. DEB-9408079 and

USDA-National Research Initiative grant no. 9503036. We also thank Laurie Dunn

for technical editing of this manuscript.

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nitrogen from fossil fuel combustion may stress the biosphere. BioScience 39: 378-386. nitrogen from fossil fuel combustion may stress the biosphere

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Bytnerowicz, A.; Miller, P. R.; Olszyk, D. M. 1987. Dry deposition of nitrate, ammonium and sulfate to a Dry deposition of nitrate, ammonium and sulfate to a Ceanothus crassifolius

canopy and surrogate surfaces. Atmospheric Environment 21: 1749-1757. canopy and surrogate surfaces

Chapin, F. S. III 1980. The mineral nutrition of wild plants. Annual Review of Ecology and Systematics 11: 233-260. The mineral nutrition of wild plants

Davis, C. M. 1994. Changes in succession after anthropogenic mechanical disturbance in coastal sage scrub. San Diego, Changes in succession after anthropogenic mechanical disturbance in coastal sage scrub

California. San Diego State University; 65 p. M.S. thesis.

DiSimone, S. 1995. California’s coastal sage scrub. Fremontia 23: 3-8. California’s coastal sage scrub

Fenn, M.E. 1991. Increased site fertility and litter decomposition rate in high- pollution sites in the San Bernardino

Mountains. Forest Science 37: 1163-1181. Mountains

Fenn, M. E.; Bytnerowicz, A. 1993. Dry deposition of nitrogen and sulfur to ponderosa and Jeffrey pine in the San Bernardino

National Forest in southern California. National Forest in southern California Environmental Pollution 81: 277-285. 

Fenn, M. E.; Dunn, P. H. 1989. Litter decomposition across an air-pollution gradient in the San Bernardino mountains. Soil Litter decomposition across an air-pollution gradient in the San Bernardino mountains

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southern California. Forest Ecology and Management. southern California

Freudenberger, D. O.; Fish, B. E.; Keeley J. E. 1987. Distribution and stability of grasslands in the Los Angeles basin. Bulletin of Distribution and stability of grasslands in the Los Angeles basin

Southern California Academic Sciences 86: 13-26.

Jennings, D. H.; Trewavas, A. J. 1986. Plasticity in plants. Cambridge: Symposia of the Society for Experimental Plasticity in plants

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Keator, G. 1994. Complete garden guide to the native shrubs of California. San Francisco, CA: Chronicle Books; 303 p. Complete garden guide to the native shrubs of California

Krywult, M.; Hom, J.; Bytnerowicz, A.; Percy, K. E. 1996. Deposition of gaseous nitric acid and its effects on foliage of ponde- Deposition of gaseous nitric acid and its effects on foliage of ponderosa pine (Pinus ponderosa Dougl ex Laws.). In: Proceedings of the 16th international meeting for specialists in air pollu- Dougl ex Laws.)

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Minnich, R. A.; Dezzani, R. J. [In press]. Historical decline of coastal sage scrub in the Riverside-Perris Plain, California. In: Historical decline of coastal sage scrub in the Riverside-Perris Plain, California

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Session I1 Nitrogen Deposition Effects on Coastal Sage Vegetation Allen, Padgett, Bytnerowicz, Minnich

SESSION III

Air Pollution/Climate Change Effects on Forests:

Results of Gradient and Case Studies

Reginald Noble and Alicja Breymeyer, Chairs

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 143

Alicja Breymeyer2

Abstract

The responses of pine forest to changing climate and environmental chemistry were studied along two transects following

the pollution and continentality gradients in Poland. One axis begins on the western border of Poland, crosses the

country along the 52nd parallel, and ends on the eastern border of Poland in the area of Bialowieza National Park,

Biosphere Reserve. The second axis begins in industrial Upper Silesia in southwestern Poland and ends at the same

point in the Bialowieza area on the eastern border of Poland. The west-east latitudinal transect follows the gradient of

cooling and continentality, whereas the Silesian transect follows the pollution gradient. The extension of the continental

transect to the west (Germany) and the east (Belarus) was possible after 2 years of the program. This “large transect”

covers 20° of latitude, with a difference in mean annual temperature of 3.5 °C. The general assumptions of this program,

e.g., philosophy of gradient/transect studies, selected global change scenarios, current climatic conditions, and air

pollution exposure in central Europe, are reviewed. The International Geosphere Biosphere Program on Global Change

predicted temperature increases of 1.5-2 °C in our climatic zone, which are close to the difference in thermoclimate

registered along the transects in this study. In the description of transects, Polish systems of forest monitoring and

procedures used for stand selection are introduced. The phytosociological characteristics of the studied stands and

their position in the classification of vegetation has been determined, as well as some of the present changes in pollutant

distribution.

Introduction

Because pollution in Silesia is clearly elevated but has steadily decreased in recent

years, it is important to understand changes in local sources of contamination and

their impact on Polish forests. Our transect studies are an attempt to relate the field

test of pine forest responses to global warming. Because central European pine

forests are seriously affected by air contamination, we try to include in the same

program the influence of pollutants on forest behavior. Sites distributed along twoaxis transects were studied (fig. 1). One axis begins on the western border of Poland,

crosses the country along the 52nd parallel, and ends on the eastern border of Poland

in the area of Bialowieza National Parks, Biosphere Reserve. The second axis begins

in industrial Upper Silesia in southwestern Poland and ends at the same point in

the Bialowieza area on the eastern border of Poland. The west-east latitudinal

transect follows the gradient of cooling and continentality, whereas the Silesian

transect follows the pollution gradient. The extension of the continental transect to

the west (Germany) and the east (Belarus) was possible after 2 years of the program.

This “large transect” covers 20° of latitude, with a difference in mean annual

temperature of 3.5 °C.

This paper studies the phytosociological characteristics of pine stands, their

position in the classification of vegetation, and some of the present changes in pollutant distribution in order to analyze the process of pollution transect selection

and assumptions about its use as a research tool.

General Assumptions

The change in global climate anticipated and most probably occurring now is

expected to produce changes in the functioning of terrestrial ecosystems. The very

rich ecological literature dealing with this subject has resulted in the development

of a variety of proposed methods to evaluate changes in ecosystems. These include

Transect Studies on Pine Forests Along Parallel

52° North, 12-32° East and Along a Pollution Gradient

in Poland: General Assumptions1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Ecologist, Institute of Geography and

Spatial Organization, Polish Academy

of Sciences, Twarda 51/55, 00-818

Warsaw, Poland.
144 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

laboratory treatment of individual organisms with differing doses of CO2 and

different thermic regimes; experiments more closely approximating natural

conditions (especially through the widespread “open chambers” in which various

simulated climates are applied to plants or habitat fragments enclosed in plastic

tubes open at both ends); studies of the reactions of whole ecosystems to artificially

manipulated changing climatic conditions (e.g., through heating of tundra soil); or

studies on natural gradients of rising mean monthly air temperatures. Studies on

comparative analysis of the same type of ecosystem functioning over suitably long

periods of time in different climates are probably the most accurate. Certainly

the use of such methods avoids the problems linked with extrapolating results

from laboratory or semi-laboratory conditions to the field, and reduces the risk

that the ecosystem response to sharp experimental intervention may be

unpredictable or chaotic.

In gradient-related ecosystem research, stability is assumed in all environmental factors except the one studied, which varies gradually but sufficiently to modify

the functioning of the ecosystem. The correlation between the changes in the factor

being studied and observed departures in the functioning of the ecosystem are

considered to express cause-effect relationships. Ideally, ecosystems that are selected carefully to be similar may also be treated like the same ecosystem subjected

to gradually-changing intensities of a given factor. Gradients in space are thus considered, within limits, as substitutes of gradients in time, which would affect the

ecosystem if it experienced the increasing action of the factor under consideration.

Published scenarios and considerations on global warming (table 1) have been

quite consistent in their prognoses concerning “the geography of climate change.”

The most clear impact of warming is expected in the Northern Hemisphere in the

zones of temperate and cool climate. Changes in atmospheric precipitation in these

zones are difficult to predict, but the zones in question should experience a clear

shift in the present ranges of the extensive deciduous and coniferous forest biomes.

Table 1 Table 1 — Predictions of temperature response to change in C0 Table 1 2

 concentration (Bolin and others 1986).

Model Globally averaged surface temperature

response to CO2 doubling

WPC (1981) 1.5 - 3.5 °C

CDAC (1983) 1.5 - 4.5 °

C

EPA (1983) 1.5 - 4.5 °

C

Clark and others (1982) 2 - 3 °

C

Julich (1983) 1 - 3 °

C

Present assessment 1.5 - 5.5 °

C

Figure 1 — (A) - “Large” climate

transect along the parallel 52°

 N.

Position of 16 stands measured

by Geographic Positioning

System (GPS) Geo Explorer,

Trimble Navigation, Model 17

319. (B) - Two axes of climatic

and pollution transects on the

schematic map of Poland. Site

names numbered according to

Institute of Forest Research

monitoring system; “K” =

climatic transect, “S” = Silesian

transect. Site “KS011” is

common to both transects; it is

the coolest site in the climatic

transect and the “cleanest” site

in the Silesian transect.

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer

K092

K098

K061

K023 K055 K117 KS011

K126

S023

S002

S060 S005

S126 S028

S011

Atlantic

Ocean

52 °N

AFRICA

ASIATICCONTINENT

EUROPE

POLAND

BELARUS

LITHUANIA

RUSSIA LATVIA

GERMANY

CZECH

REPUBLIK

UKRAINE

RUSSIA

52°39'14°32'

52°49'19°00'

52°44'15°42'

52°48'19°19'

52°49'20°19' 52°40'

21°42'

52°55'23°37'

52°56'28°53'

52°48'27°08'

52°52'28°25'

53°14'30°12' 53°25'

31°06'

N

E

53°06'12°25'

52°47'16°26'

53°20'32°35'

53°20'32°37'
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 145

Attempts to predict global changes over time have been made by Boryczka

and Stopa-Boryczka (1992), who expect annual air temperatures to decrease between 1990 and 2050, and then to rise between the years 2050 and 2100. The overall

rise in temperature between 1990 and 2100 is an anticipated 0.5 °C, a figure significantly lower than that suggested in the various computer simulations and experiments linked with the International Geosphere Biosphere Program and usually involving a doubling of carbon dioxide concentrations by 2030 and a consequent

warming of 2-3 °C (table 1).

Climatic conditions in central Europe are determined by Arctic influences along

the north-south axis (cooling in the direction of the Pole), as well as by oceanic

influences of the Atlantic along the west-east axis (greater extremes of temperature

and cooling towards the interior of Asia) (fig. 2). Smialkowski (1995) described the

climatic characteristics from meteorological stations near research sites along the

transects (fig. 3). The mean annual air temperature declined from 8.3 °C in the west

to 6.7 °C in the east, while the total annual precipitation amounted to just over 500

mm throughout. The Upper Silesian transect was characterized by a fall in temperature from 8.1 (Silesia) to 6.7 °C at the northeastern end. The mean annual amplitude of air temperature (based on monthly average temperatures), a measure of

the continentality of climate, increased from 18.8 °C in western Poland to 22.3 °C

near the border with Belarus.

Description of the Transects

Scots pine (Pinus sylvestris L.) dominates 70-80 percent of Poland’s forests (in area

covered or tree biomass) (Trampler and others 1987). The forests are very sensitive

to pollution, and as a zonal community, must react in a distinct way to warming of

the climate. It was accepted initially (Breymeyer 1994) that two complementary

transects should be designated within Poland: a climatic (west-east) transect along

parallel 52° N and a southwest-northeast transect running from polluted Upper

Silesia to Bialowieza (fig. 1). Information on the state of Poland’s forest across this

large area has been collected by the forestry services. Monitoring of the state of

Poland’s forests is performed by various institutions. We used the network directed

Figure 2 — (A)-Climatic

differentiation of Europe: height

of the timberline (in meters,

isolines) in European mountains

as an index of northward cooling;

eastwards continentality expressed as increasing difference

between temperatures of coldest

and warmest month (in °

C,

shadowed surfaces) (Mayer

1984). (B)-Duration of the

vegetation growing period (days)

in various forests. Numbers

placed on the lines show the

difference between the warmest

stand “Slubice” and individual

stands (Falinski 1986).

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer

A B

ROMINTY F.

AUGUSTOW F.

165

215

180

KNYSZYN F.

180

BIALOWIEZA F.

185

POLESIE

180

ROZTOCZE

195

SLUBICE
146 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

by the Forest Research Institute. Basic data are collected on permanent research plots

(SPOs) covering the entire country and established in 1989 in stands of Scots pine,

Norway spruce, fir, oak, beech and birch greater than 40 years old. The selection and

siting of SPOs reflects the cover, age and species structure, as well as the condition of

Poland’s forests. Over 1,000 SPOs were established, each with 20 marked trees of the

dominant species observed annually for the following characteristics:

● Morphological characteristics of the crown;

● Defoliation and the coloration/discoloration of foliage and assignment

of trees to the appropriate damage classes;

● A phytopathological evaluation and an assessment relating to insect pests;

● Tree diameter measurements every 5 years.

The results of all measurements are recorded in a uniform manner in a databank.

Selection and Characterization of Sites

This research was based on a two-axis transect reflecting to some extent the

influences of climate and pollution on the forests studied. The “Silesian” axis traced

the gradient of air pollution from Silesia to the Bialowieza Forest, while the climatic

axis traced the gradient of continentality along parallel 52° N from Poland’s western

to eastern borders, again terminating near Bialowieza on the same KS011 stand

(fig. 1). Research sites were selected from a total of approximately 1100 SPOs based

on the criteria of identical type of forest habitat, soil type, age of pine, indices of

forest cover and stand quality, and similar aspect, slope, and elevations. The last

stage of the selection process was implemented by site visits leading to the final

choice of 15 permanent research plots.

Forests are complicated ecosystems; their functioning is conditioned by a

long history (usually several generations of trees), by current and past management, and by a whole mosaic of habitat factors that vary considerably across

Poland. Thus, the choice of sites along the transect was difficult, and despite

considerable care and effort, we could not completely avoid the influence of

confounding factors.

Figure 3 — Climatic characteristics of transects: mean annual

air temperatures (lines), precipitation totals (bars) and annual

amplitudes of temper-atures.

Long-term data collected from

meteorological stations in the

vicinity of 15 studied sites

(Smialkowski 1994). –

–––––––

1000

800

600

400

200

Precipitation totals (mm)

0

8.5

8

7.5

7

6.5

6

At temperature (C)

K092 K098 K061 K023 K055 K117 K126 KS011

Site number

P T

––––––––

1000

800

600

400

200

Precipitation totals (mm)

0

8.5

8

7.5

7

6.5

6

At temperature (C)

S126 S005 S028 S060 S002 S011 S023 KS011

Site number

23

22

21

20

19

At temperature Amplitude (C)

18 K092 K098 K061 K023 K055 K117 K126 KS011 –

–––––––

Site number

23

22

21

20

19

At temperature Amplitude (C)

18 –

–––––––

Site number

S126 S005 S028 S060 S002 S011 S023 KS011

P T

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 147

Ecosystems were mainly treated as functional units. According to Breymeyer

(1981, 1984) and Breymeyer and Uba (1987), we assumed that the responses of ecosystems to environmental stresses can be measured via responses in the main processes occurring within them, including the cycling of matter (i.e., its production

and decomposition). Responses to change by these processes may be intensified or

reduced in strength under different conditions, and their proportions may change.

However, the processes must continue and maintain the cycling of matter in

the ecosystem. If these processes were to discontinue, the ecosystem would

similarly cease to exist. Because changes in the intensity of ecosystem processes

are measurable, the responses of ecosystems to stress are also measurable. Along

the transects studied, the functioning of ecosystems was evaluated by measuring litter fall and the rate of decomposition of organic matter in the litter layer

and in underlying soil layers.

The structural features of forest ecosystems are composed of soil, vegetation

and different groups of organisms. The composition and structure of vegetation

were used to group plant communities according to phytosociological classification. The forests of Poland and central Europe have been described and classified

by using the Braun-Blanquet system, and maps of the potential vegetation of Poland and Europe have been established. The forests studied in our program have

been described in accordance with this classification. All of the sites studied belong

the Dicrano-Pinion alliance and include two associations of pine forest: PeucedanoPinetum (sub-continental pine forest) and Leucobryo-Pinetum (sub-oceanic pine forest), as well as mixed forest of Querco-Pinetum type. Pine forests occupy the same

place along the scale of humidity, but differ in the degree of continentality

(Matuszkiewicz and Matuszkiewicz 1973) (fig. 4). The communities of the LeucobryoPinetum association cover a large area and are the most common types of forest in

Poland, showing little regional differentiation. Further east, the sub-oceanic type is

replaced by the sub-continental Peucedano-Pinetum, which occurs to the east of the

line formed by the Bug and Lower Vistula Rivers. Sub-continental forest is much

richer floristically and characterized by the presence of continental species, mainly

perennials of a somewhat xerothermic nature, as well as by limited regional variation.

Some of the sites had a deciduous component, so they were classified ultimately as

mixed pine forest of the Querco- Pinetum type (fig. 4).

In the series of stands described above, the changes in behavior of pine ecosystems were measured by groups of specialists. The following indexes of ecosystem

Figure 4 — Regional diversity

of pure coniferous (A) and mixed

(B) pine forests in Poland

(Matuszkiewicz 1987).

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer
148 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

function and structure were selected for study:

● Soil characteristics such as vertical structure and physio-chemical

properties;

● Cotton decay in various soil layers as an index of the stratification

of biological activity;

● Litter fall and decomposition as an index of cycling rate of organic matter

(OM budget);

● Tree rings increment as an index of stand history influencing its

present condition;

● Potential vegetation (sensu Tuxen 1956) of the stand as an index of stands

similarity and position in the vegetation classification system;

● Selected characteristics of present plant cover: diversity of species and

forms, biomass, and chemistry (nutrients and pollution) of dominant

species of herbs and shrubs.

All these indexes are related to simultaneously evaluated climatic factors and

inputs of pollutants.

The fundamental idea of the double transect, with gradients focused mainly

on two groups of factors (climate, pollution), has since been affected by certain

changes. At the time the research began it was obvious that the pollution transect

must run from the south-west (Upper Silesia region) to the north-east (Bialowieza

forests). In the literature from the previous 20 years, very high levels of pollution

were reported for the industrial Silesia region. Furthermore, mathematical modeling of the origin and distribution of pollution throughout the country demonstrated

that Silesia was the dominant pollution source region (Juda-Rezler and Abert 1994;

fig. 5). However, recent years show a clear improvement in air quality, with a progressive decline in air pollution in the province of Katowice, Silesia’s largest city

(table 2). Recent years have also seen a distinct decrease in air pollution at “background” monitoring sites in regions far from emission sources (Przbylska 1995;

fig. 6). Some time will doubtless pass before the first signs of improvement in the

condition of forests are noted, but such changes will certainly occur. Thus, the research reported here is expected to document changes in forest condition which

will certainly occur; and this research will coincide with a certain blurring in the

gradient of pollution gradient along the axis between Silesia and northeastern

Poland. In such a situation, local pollution is inevitably of greater significance

than had been anticipated.

We decided to run the climatic gradient latitudinally along the 52nd parallel

for several reasons. This avoided the inclusion of sites conditioned by higher altitude characteristics for the southern half of Poland. In addition, the selected transect

is the longest one possible for the country’s coniferous forests, and it allows for

extension to the east or the west without changing the above conditions.

This extension was of value in the second year of the research, when a first

expedition to Belarus led to the designation of six new sites between the PolandBelarus and Belarus-Russia borders (fig. 6). The third year of the study, two more

Figure 5 — Mean annual

concentrations of S and N

compounds over Poland in 1990

(Jura-Rezler and Abert 1994).

(A)-Concentration of NO,

isolines: 0.1-1.5 μg N per m3

.

(B)-Concentration of S02

 (μg S

per m3

), isolines: 2, 3, 4, 5, 10,

15, 20, 25, 30, 35.

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 149

sites were added in Germany (at the western end of the transect). The extended

transect currently runs across 20° of longitude (from 12°25' E to 32°60' E), has a

range of annual temperatures of 3.5 °C (from 5 to 8.5°), and has annual amplitudes

of temperature ranging from 18.2 (west) to 26 °C (east). However, this study has

focused on results from the original Polish portion of the transect.

Figure 6 — Concentration in

atmospheric precipitation (A)

wet depositions (B) of some

elements, hydrogen ions and

pH as measured in five

meteorological stations located

across Poland. The program of

observations is comparable to

the international systems WMO,

EMEP, BMP, as well as to the

Polish national monitoring

system (Przybylska 1995).

Table 2 Table 2 — Emission of air pollutants in the Katowice District (1985-1992) (Katowice Voivodship Statistical Table 2 

Manual 1993).

Emission of Pollutants

(thousand tons / year) (thousand tons / year)

 Year Dust Gases

1985 453.9 1540.9

1986 418.7 1509.9

1987 389.3 1505.2

1988 327.9 1306.3

1989 305.3 1309.2

1990 227.1 1003.3

1991 186.7 850.2

1992 127.1 738.1

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer mgSI-1 pH units mgS/l mgN/l mgH+/m2yr gS/m2yr gN/m2yr
150 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Acknowledgments

The study was done within the framework of a project entitled “U.S.-Poland

Cooperative Project on the Status and Long-Term Trends in Forest Ecosystems:

Climate, Pollution and Forest Health,” and a project “Effects of Atmospheric

Deposition and Climate Change on Forest Ecosystems in Poland and the United

States.” The projects for this study were financed by the U.S. Environmental

Protection Agency; the USDA Forest Service; the USDA Foreign Agriculture Service

International Cooperation and Development (ICD); the Polish Academy of Sciences;

the Polish Bureau of Global Environmental Facilities (GEF); the Polish Forest

Research Institute; Michigan Technological University, Houghton, Michigan;

Bowling Green State University, Bowling Green, Ohio; and the Institute of

Geography and Spatial Organization of the Polish Academy of Sciences, Warsaw.

References

Bolin, B.; Doos Bo, R.; Jager, J.; Warrick, R.A. 1986. The greenhouse effect, climate change and ecosystems. SCOPE 29. The greenhouse effect, climate change and ecosystems

Chichester: John Wiley and Sons; 541 p.

Boryczka, J.; Stopa-Boryczka, M. 1992. Atlas wspolzaleznosci parametrow meteorologicznych i geograficznych w Polsce. (At- Atlas wspolzaleznosci parametrow meteorologicznych i geograficznych w Polsce. (Atlas of interdependence of meteorological and geographical parameters in Poland). Vol. 7. Warsaw. las of interdependence of meteorological and geographical parameters in Poland)

Breymeyer, A. 1981. Monitoring of the functioning of ecosystems. Environmental Monitoring and Assessment 1: 175-183. Monitoring of the functioning of ecosystems

Breymeyer, A. 1984. Ecological monitoring as a method of land evaluation. Geographia Polonica: 371-384. Ecological monitoring as a method of land evaluation

Breymeyer, A. 1994. Functioning of forest ecosystems in gradients of climate and pollution: Project for comparative ecosys- Functioning of forest ecosystems in gradients of climate and pollution: Project for comparative ecosystem studies on transects. In: Solon, J.; Roo-Zielinska, E.; Bytnerowicz, A., eds. Climate and atmospheric deposition stud- tem studies on transects

ies in forests. Warsaw, Poland: Institute of Geography and Spatial Organization, Polish Academy of Sciences, Conference

Papers 19: 101-107.

Breymeyer, A.; Uba, L. 1984. Monitoring ekosystemow: program ekologicznej kontroli stanu ekosystemow ladowych na

obszarze Polski. (Ecosystems monitoring: the program of ecological control of terrestrial ecosystems condition in Po- obszarze Polski. (Ecosystems monitoring: the program of ecological control of terrestrial ecosystems condition in Poland). Czlowiek i Srodowisko (Man and Environment): 471-495. land)

Falinski, J.B. 1986. Vegetation dynamics in temperate lowland primeval forests. Geobotany 8: 1-537. Vegetation dynamics in temperate lowland primeval forests

Juda-Rezler, K.; Abert, K. 1994. Rozklad stezen i strumieni zanieczyszczen powietrza w Karkonoszach na podstawie modelowych

badan symulacyjnych. (The distribution of concentrations and streams of air pollution in the Karkonosze Mountains on

the basis of simulation research). In: Fischer, Z., ed. Ecological research in the Karkonosze. Dziekanow Lesny: Institute of the basis of simulation research)

Ecology PAS; 29-62.

Katowice Voivodeship Statistical Manual Katowice Voivodeship Statistical Manual. 1993. Katowice. Katowice Voivodeship Statistical Manual

Matuszkiewicz, J. 1987. Geobotaniczne zroznicowanie lasow iglastych Polski. (The geobotanical differentiation of Poland’s

coniferous forests). In: Breymeyer, A., ed. Coniferous forests in Poland. Institute of Geography and Spatial Organization, coniferous forests)

Polish Academy of Sciences; Dokumentacja Geograficzna 3: 46-72.

Matuszkiewicz, W.; Matuszkiewicz, J. 1973. Przeglad fitosocjologiczny zbiorowisk lesnych Polski. Cz.2. Bory sosnowe. (A

phytosociological review of Poland’s forest communities. Pine forests). phytosociological review of Poland’s forest communities. Pine forests). Phytocoenosis 2 (4): 273-356. phytosociological review of Poland’s forest communities. Pine forests).

Mayer, H. 1984. Walder Europeas. Stuttgart: Gustav Fisher. Walder Europeas

Przybylska, G. 1995. Wieloletnie zmiany zanieczyszczenia atmosfery w Polsce. (Long-term changes in atmospherical pollu- Wieloletnie zmiany zanieczyszczenia atmosfery w Polsce. (Long-term changes in atmospherical pollution in Poland). In: Ocena i stan srodowiska przyrodniczego Polski i innych krajow. Miedzynarodowe Sympzjum Naukowe tion in Poland)

(Evaluation of natural environment in Poland and other countries). International Symposium. 1995 October 10-13;

Jachranka; 112-114a.

Solon, J. 1995. Diversity of herb layer in pine forest community along climatic and pollution gradients. In: Forest ecosystems Diversity of herb layer in pine forest community along climatic and pollution gradients

in Central and Eastern Europe and the United States. Report 1994-1995. Warsaw: Institute of Geography and Spatial

Organization, Polish Academy of Sciences; 46-64.

Smialkowski, J. 1995. Differentiation of thermic and precipitation conditions as an effect of altitude and distance from the

Atlantic Ocean on the “climatic” and “Silesian” transects. In: Effects of atmospheric deposition and climate change in Atlantic Ocean on the “climatic” and “Silesian” transects

forest ecosystems in central and eastern Europe and the United States, 1994-1995 Report. Warsaw: Institute of Geography and Spatial Organization, Polish Academy of Sciences; 24-25.

Trampler, T.; Girzda, A.; Dmyterko, E. 1987. Zroznicowanie zasobow i produkcji biomasy drzewnej w lasach iglastych w

Polsce, (Differences in resources and the production of tree biomass in Poland’s coniferous forests). In: Breymeyer, A., Polsce, (Differences in resources and the production of tree biomass in Poland’s coniferous forests)

ed. Lasy iglaste na obszarze Polski (Pine forests in Poland). Institute of Geography and Spatial Organization, Polish Academy of Sciences, Dokumentacja Geograficzna 3: 46-72.

Tuxen, R. 1956. Die heutigepotentielle naturliche Vegetation als Gegenstad der Vegetations Kartierung. Angewandte Die heutigepotentielle naturliche Vegetation als Gegenstad der Vegetations Kartierung

Pflanzensoziologie 13: 5-42.

Session I1I Transect Studies on Pine Forests Along Parallel 52° North, 12-32° East Breymeyer
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 151

Ewa Roo-Zielinska, Jerzy Solon2

Abstract

The influences of a geographical location on floristic composition, horizontal structure, and biomass of the herb layer

in pine and mixed pine forest communities along climatic and pollution gradients in East Germany, Poland, and

Belarus were determined. Phytosociological records were collected in permanent plots in May 1995. Each record covered

an area of 400 m2

. The floristic composition of the study area provided the basis for the determination of the

phytosociological differentiation of the plots along the transect. Similarity index between plots was calculated by

using the Czekanowski index; a similarity dendrogram was constructed according to Ward’s method. Biomass was

determined in six 0.1 m2

 replications sampled in each permanent plot. All of the identified study areas were representative

of the Dicrano-Pinion alliance and included two communities of pine forest (Peucedano-Pinetum and LeucobryoPinetum) as well as the community of mixed pine forest (Querco roboris-Pinetum). On the basis of similarity indices,

two geographical groups of plots were distinguished: suboceanic and subcontinental. Geographical differentiation of

the stands is related to the presence of the groups of species of different geographical distribution type. For the western

part of the transect, characteristic species are Deschampsia flexuosa and Leucobryum glaucum. Central and eastern

parts are characterized by the presence of the following species: Melampyrum pratense, Luzula pilosa, Veronica

officinalis, Lycopodium clavatum, Peucedanum oreoselinum and Scorzonera humilis. On the stands located on

the easternmost part of the transect (Belarus) frequent species are: Pirola chlorantha, Chimaphila umbellata,

Koeleria pyramidata and Hypochoeris maculata. The ground flora of the identified areas was not uniform; rather

it was characterized by a mosaic of sparse populations of plants. In general, three types of these systems can be

distinguished: one is dominated by a cover of mosses, the second is dominated by dwarfshrubs, and the third by

species of grass. The ground cover biomass on the west-east transect ranged from 49 to 544 g/sq. m, but for pine forests

the correlation between geographical longitude of the stand and biomass of mosses (corr. coeff. 0.701) and total biomass

of the ground cover (corr. coeff. 0.779) was statistically significant.

Introduction

This article presents a geobotanical analysis of the floristic composition of the herb

layer and its structure and biomass in East Germany, Poland, and Belarus. It

constitutes part of long-term, in-depth community research aimed at defining the

influence of geographical location on pine forest ecosystems, in particular pine

forests and mixed/pine forests (Breymeyer 1996, Degórski 1996, Smialkowski 1996).

The subjects of geobotanical analyses consisted of 17 permanent plots lying

along a west-east transect following the gradient of continentality. These are mature or almost mature pine stands, basically single-species and single-aged (60-95

years). All the analyzed forest communities represent (in accordance with phytosociological typology) the Dicrano-Pinion alliance, and within it two communities of

pine forest, Peucedano-Pinetum and Leucobryo-Pinetum, as well as the community of

mixed pine forest, Querco roboris-Pinetum. In accordance with the principles of forest typology these are different forms of pine forest occurring on sandy substrates.

Their economic use does not differ significantly.

The level of differentiation of the herb layer is one of the more important structural characteristics of forest phytocoenoses. On the one hand it influences the spatial differentiation for various processes like the production and decomposition of

organic matter, and on the other it may be an indicator of habitat

microdifferentiation (in particular topography, fertility, and humidity). A way of

describing the level of diversity of the herb layer is through the identification,

Geographical Differentiation of the Floristic Composition

and Structure of the Herb Layer of Forest Permanent Plots

in East Germany, Poland, and Belarus1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Botanists, Institute of Geography and

Spatial Organization, Polish Academy

of Sciences, 00-927 Warsaw,

Krakowskie Przedmiescie 30, Poland.
152 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

analysis, and typology of synusiae. Synusia is a grouping, within one layer of a

community, of species characterized by similar life forms and similar ecological

requirements. The synusia is sometimes used as a basic spatial unit, as well as a

functional-temporal unit within a phytocoenosis (Gillet 1986, Julve and Gillet 1994,

Mavriscev 1980).

Because the herb layer reacts most actively to all environmental variables, we

attempted to define the influence exerted by climatic differentiation and habitat

(also microhabitat) conditions on the species composition, structure, and biomass

of the herb layer. The floristic composition and structural characteristics of the studied communities depend on many factors, such as:

● Geographical location, which determines both the richness of the local

flora (influencing the occurrence of species of a defined type of range), and

is an indicator of macroclimatic differentiation. Climatic characteristics (in

particular mean annual temperature, the temperature of the warm and

cold periods of the year and total precipitation) have a considerable

influence on the phenology and competitive abilities of

different populations;

● The type and properties of the habitat, mainly its fertility and humidity,

which influence the presence or absence of the species building up a given

phytocoenosis;

● Anthropogenic influences linked with the pollution of the environment,

the history of its use, and the maturity of phytocoenoses.

This paper describes a study to define the mutual similarities and homogeneity of

the study sites on the basis of the floristic composition of the herb layer in East

Germany, Poland and Belarus; presents the synusial differentiation of the herb layer;

draws up a typology of the pre-analyzed synusias, and defines the mean biomass

within the different types of synusia and within the study areas.

Methods

Floristic Composition

The basis for the description of the 17 study sites used in this study was a floristic

characterization of the patch of vegetation representing a given type of community

and described in the form of a phytosociological record. This contained the following

basic information on a given patch of vegetation: the species composition; cover

(i.e., the proportion of the area occupied by each species); and layer structure (vertical

structure of the community), identified most often as A–trees, B–shrubs, C–herbs,

D–ground cover of mosses and lichens. The majority of phytosociological records

were taken in May 1995. Each record covered about 400 m2. The quantitative

representation (cover) of species was estimated by using a modified 12-point scale

on which r, +, and 1, refer to 0.1 percent, 0.5 percent, and as high as 10 percent,

respectively, while the remaining values from 2-10 relate to the successive 10 percent

divisions. Vascular plant species were defined by using the key of Szafer and others

(1969) and the nomenclature of Rothmaler (1976), while nomenclature of mosses

followed Szafran (1957, 1961), liverworts followed Rejment-Grochowska (1950) and

lichens followed Faltynowicz (1993).

The phytosociological records obtained were presented in an unordered table,

which was interpreted and described in accordance with the system of classification and list of diagnostic species of Matuszkiewicz (1981).

By using Czekanowski’s measure of distance, we calculated the mutual similarity of the identified forest communities on the basis of the floristic composition

of the herb layer. This was presented with the aid of a dendrogram produced by

using Ward’s method (Solon 1994).

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 153

Synusial Structure

The basis for the description of the synusial structure of an area’s ground cover

was field mapping of the most typical 5 by 20 m rectangular areas in 9 out of 17

study sites (N1, K092, K098, K061, K023, K055, K117, K126 and K011) lying on the

west-east transect in Germany and Poland. In addition, floristic inventories were

carried out within the different synusiae. These were the bases for the identification

of types of synusia. Inclusions in the typological table were confined to those species

whose cover was greater than 10 percent. The mapping and inventory of plant

species was done in May 1995.

The similarity of the floristic compositions of different synusiae was calculated

by using Czekanowski’s measure of distance and presented with the aid of a dendrogram produced by using Ward’s method (Solon 1994).

Biomass

Biomass was determined once during September and October 1995 at all 17 sites.

Six circular biomass samples were taken from a 0.1 m2 area of each permanent plot.

The choice of the area for replicates was not random; rather, we attempted to take

replicates of the dominant types of synusiae at a given study site. The entire biomass was collected in the field and then sorted into living moss biomass and higher

plants. The dead parts of plants were removed. The samples were dried to constant

weight at a temperature of 90 °C and weighed to an accuracy of 0.01g. The results

obtained were averaged for the type of synusia and for the study area and recalculated per m2

.

Results

Floristic Composition

Areas K092 and K023 had Leucobryo-Pinetum pine forest in the typical form, while

area K061 had a more humid variant with Molinia caerulea. In contrast, the two

areas N1 and K098 were occupied by communities occurring in an acid beech forest

habitat (Luzulo-Fagion alliance) or by mixed-pine forest, although they are almost

identical to the communities representing Leucobryo-Pinetum. The Peucedano-Pinetum

community is represented by areas K011, B2, B3 and B5 (type or sub-boreal forms),

as well as by a more humid variant with Molinia caerulea on areas S023 and B7. The

remaining areas had mixed pine forest (Querco roboris-Pinetum), with the typical

form on areas K055, K126, B1, B4 and B6, and area K117 consisted of a degraded

habitat that was a poor form of oak-linden-hornbeam forest Tilio-Carpinetum.

Analysis of the similarity of the species composition of vascular plants between

the different sites clearly points to the occurrence of two large groups of communities: the subcontinental and the suboceanic. But the compositions of these groups

differ, depending on the manner in which similarity is defined, e.g., on the basis of

cover (quantitative representation) of species or on the basis of the presence or

absence of species. The core of the suboceanic group is comprised of areas N1,

K092, K098, K061 and S023. As cover (quantitative representation) of species, areas

K117 and K126 also belong to this group, but if defined as the similarity of the

species composition, these are part of the subcontinental group.

The division into groups of lower rank is also first and foremost geographical

in nature. Distinguished within the sub-Atlantic group is a western subgroup,

although on the basis of cover the habitat division is more important because it

clearly separates out the subgroup of sites of greater humidity (K061, S023)

characterized by a greater share of purple moor grass (Molinia caerulea). Two

geographical subgroups may also be clearly distinguished within the subcontinental

group. On the basis of the presence or absence of species, all the Belarus sites (B1-

B7) create one subgroup, while on the basis of quantitative representation sites B1

and B3 are more similar to the Polish sites.

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
154 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 1 — Presence of chosen

species along the the west-east

transect (Rothmaler 1976).

One of the factors in the spatial variation of the floristic composition of pine

forests dependent on geographical location is the differentiation in the ranges of

particular species of plant, which, at least theoretically, consist of vegetation

conditions that are appropriate for their existence. A clear suboceanic character is, for example, displayed by Deschampsia flexuos and Leucobryum

glaucum, while the areas of a subcontinental character are most strongly

associated with such species as Chimaphila umbellata, Hypochaeris maculata

and Koeleria pyramidata (fig. 1).

Synusial Structure

On the basis of dominant species, life forms, and the degree of cover of the area

with vegetation, it is possible to distinguish six main types of synusiae (table 1). The

first type includes synusiae with an absolute dominance of Vaccinium myrtillus, a

limited role of mosses, and the sporadic occurrence of various grassy species. The

second type is characterized by the dominance of Vaccinium vitis-idaea and compared

to the first type, it occurs in places that are drier locally. The third type includes

mossy synusiae with a predominance of Pleurozium schreberi. This is the type

occurring most frequently and at the same time most differentiated into subtypes

in relation to the presence and character of codominants and subdominants. In

particular, it is possible to distinguish pure mossy, mossy-cowberry, mossy-bilberry,

mossy-sedge, mossy-fescue, mossy-hairgrass, and mossy-hairgrass-fescue subtypes.

The fourth includes synusiae characterized by the codominance of three or four

species and the lack of a clear dominant. Within this type, the first subtype includes

a mossy-grassy synusia and the second a mossy-heather one. A fifth type includes

a synusia with sheep’s fescue (Festuca ovina) as dominant, while a sixth type is

characterized by a very thin vegetation cover not exceeding 20 percent.

Different study areas differ clearly on the basis of the set of synusial types as

well as in their spatial distribution (fig. 2). Mapped fragments of the areas have 2

to 4 types of synusia creating between 3 and 32 individual patches. However, mixed

pine forests have a much more mosaic-like herb layer because these sites are observed to have greater numbers of patches of synusiae of different types (fig. 2; for

example, compare sites K092 and K126).

LONGITUDE

12 15 19 22 27 33

Distribution Type

ozCIRCPOL

arct(subk)CIRCPOL

b.(oz)EUR-WAS

b.(oz)EUR-WSIB

b.(oz)EUR

temp.subozEUR

temp.(suboz)EUR

b.subkCIRCPOL

temp.subozEUR

b.(subk)EUR-WAS

SPECIES

Deschampsia flexuosa

Vaccinium vitis-idaea

Melampyrum pratense

Luzula pilosa

Veronica officinalis

Peucedanum oreoselinum

Scorzonera humilis

Chimaphila umbellata

Hypochoeris maculata

Koeleria pyramidata

Lecanora conizaeoides

Cladonia chlorophaea

Pseudevernia furfuracea

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 155

The geographical differentiation of synusiae is poorly marked and results from

differences in the ranges of the dominant species (Roo-Zielinska and Solon 1996).

Western areas are associated with mossy-hairgrass, mossy-hairgrass-fescue, and

mixed mossy-grassy synusiae. The majority of types of synusiae occur in both fresh

pine forests and mixed/pine forests. The only synusiae to be closely associated

with fresh pine forests are mixed mossy-grassy and mossy-heather, as well as mossybilberry and pure mossy synusiae. In contrast, mixed/pine forests more often have

mossy-fescue, mossy-sedge, or fescue synusiae.

Biomass Differentiation

Biomass in Synusiae

The different synusiae and their types are highly diversified in terms of biomass of

mosses and vascular plants. Degraded synusiae (type VI) have total biomass of

about 15 g/m2, while the mossy-cowberry subtype attains more than 600 g/m2.

The variability in the biomass of the different types of synusiae is also relatively

great, and the differences between the most luxuriant and the poorest may be as

large as 300 percent, as is the case for the mossy-bilberry subtype. This variability

may only be partly explained by reference to geographical variability (fig. 3). The

only statistically significant (p < 0.05) correlation (r=0.67) with longitude is noted

for the biomass of mosses within the mossy-bilberry subtype of synusia. In the case

of this subtype the correlations between longitude on the one hand, and the biomass

of vascular plants and total biomass on the other are relatively great (about 0.5) but

of very low significance. A similar situation is also noted in the case of pure mossy

synusiae. The low level of significance of the correlation is linked with the limited

abundance of the analyzed sample, and the results may attest to the limited

variability of biomass in the different types of synusiae in the west-east direction.

Several types of synusia have ratios of vascular plant to moss biomass that are

relatively constant and clearly different from those of other types. However, this is

not a consistent ratio. In the case of the bilberry and cowberry synusiae (I and II),

the ratio attains values between 4 and 5. For the mossy type (III), it is almost always

below 1 (only in a few cases is it greater than 1, reaching about 3). For the mixed

type (IV) it varies from about 1 to 3 and in the case of the fescue type (V) from about

3.5 to 5.5. The degraded type has values oscillating around 1, while the purple

moor grass type (VII) always has a value more than 2 and sometimes much more

than 10 (table 2).

Type

Description number

Vaccinium myrtillus

Vaccinium vistis-idaea

Entodon schreberi

Deschampsia flexuosa

Festuca ovina

Calluna vulgaris

Molinia caerulea

Carex ericetorum

Calamagrostis arundinacea

Lichenes

Bare ground

I.1b

055

b

2

1

4

I.1a

N1

a

5

2

1

1

1

II.1

098

c

3

1

1

3

061

a

4

3

 

1

III.2 

011

d

4

4

1

1

011

b

2

4

III.1d

023

b

3

5

061

d

2

5

 

1

055

c

5

 

1

023

a

1

4

 

1

III.1a 

126

a

1

1

5

1

011

c

1

1

5

1

1

III.1c

126

d

1

1

5

1

2

011

a

4

5

III.3 

055

a

3

4

1

III.4

055

d

3

5

4

126

b

5

3

III.5 

117

d

5

4

III.1e

092

b

5

1

2

III.6

092

a

5

3

1

III.7a

098

a

1

5

3

3

III.7.b

098

b

2

1

5

3

3

IV.1 

023

c

2

3

2

N1

b

1

3

3

3

V.1b

117

e

2

3

2

1

117

g

1

4

1

V.1a 

117

c

1

4

1

117

f

3

1

1

2

III.1b 

117

a

3

1

2

IV.2b

061

c

2

2

2

2

1

IV.2a

061

b

2

3

1

1

126

c

1

1

5

VI 

117

c

1

1

1

4

061

e

1

1

4

I.Vaccinium myrtillus type

I.1. -(a - typical; b - lose)

II.Vaccinium vitis - idaea type

II.1Vaccinium vitis - idaea

III.Pleurozium schreberi type

III.1.(a - typical; b - lose; c - with Calluna vulgaris; 

d- with Vaccinium vitis - idaea; 

e - with Festuca ovina)

III.2.Pleurozium schreberi - Vaccinium vitis - idaea subtype

III.3.Pleurozium schreberi - Vaccinium myrtillus subtype

III.4.Pleurozium schreberi - Calsmagrostis arundinacea subtype

III.5.Pleurozium schreberi - Festuca ovina subtype

III.6.Pleurozium schreberi - Deschampsia flexuosa subtype

III.7.Pleurozium schreberi - Festuca ovina - Deschampais flexuosa subtype

IV. mixed type

IV.1.mixed moss - grass subtype

IV.2.Pleurozium schreberi - Celluna vulgaris subtype

(a - typical; b - with Molinia caerulea

V.Festuca ovina type

VI.degraded

VI.1.strongly degraded 

Table 1 Table 1 — Floristic composition of synusiae on the west-east transect. Table 1 

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
156 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 2 — Synusial differentiation of study plots (see table 2

for legend).

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 157

Mean Biomass in the Community

The mean biomass of the herb layers and mossy layers at the study sites vary with

geographical location (fig. 3). The correlation coefficient for moss biomass against

longitude for all 17 sites on the west-east transect amounted to 0.701***, while the

values for fresh pine and mixed/pine forests were 0.701* and 0.712*, respectively

(in which ***, **, and * respectively indicate significance at the p < 0.001, 0.01, and

0.05 levels). Longitude is also correlated with the combined biomass of the herb

layer, mosses in pine forests (0.779**), and in all sites (0.779***). If the mean biomass

of the richest type of synusia is considered a site characteristic, then correlations

have similar values: the correlation coefficient for moss biomass and longitude for

all 17 sites on the west-east transect is 0.687***, while that for fresh pine forests is

also 0.687*. Moreover, the coefficient defining the relationship between longitude

and total biomass of mosses and vascular plants attains respective values of 0.847***

and 0.848***. Both mean biomass of the community and mean biomass of the richest

synusia showed no statistically significant correlation between the biomass of

vascular plants and longitude.

Discussion and Conclusions

Changes in floristic composition were detected along the west-east transect both

within pine forests and mixed-pine forests. Differences in floristic composition

correspond to differences in the habitat (connected with the division into pine forests

and mixed-pine forests, and the occurrence of typical forms and wetter forms with

Molinia caerulea) and also to the division into geographically varying communities

within pine forests (the division into Peucedano-Pinetum and Leucobryo-Pinetum).

The main spatial geographic factor in the variability of the floristic composition of

pine forests is differentiation in the ranges of understory plant species, which find

appropriate conditions for their existence in this type of vegetation. Sites in central

and southern Poland may be intermediates between the well-defined western and

eastern pine forests. The geographical and edaphic differentiation of the study areas

is reflected most strongly in the herb layer, although it is not very visible among

lichens. Differences in this group are influenced to a greater degree by local factors,

including the land use history of the area, than by microhabitat factors.

Pine forests and mixed pine forests did not have homogeneous herb layers;

they were differentiated into fragments creating a system of separate micro-areas

varying in the dominance of particular species and life forms. A detailed analysis

of the spatial variability of the biomass of the herb layer and of bryophytes was

Figure 3 — Changes in biomass

of study plots along the westeast transect: pine forests (A);

mixed forests (B). Variables:

makniem = mean biomass of

vascular plants in the richest

synusium of the plot; makm =

mean biomass of mosses in the

richest synusium of the plot;

makraz = mean biomass of

vascular plants and mosses in

the richest synusium of the plot;

srniem = mean biomass of

vascular plants on the plot; srm

= mean biomass of mosses on

the plot; srraz = mean biomass

of vascular plants and mosses

on the plot.

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon g/m2 g/m2
158 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Table 2 – Biomass differentiation (g/sq. m) of synusiae and permanent plots. Table 2 – Biomass differentiation (g/sq. m) of synusiae and permanent plots.

Herbs Mosses Herbs & Mosses

Site Date Type Number Mean Standard Mean Standard Mean Standard

1995 of deviation deviation deviation

replications

N1 14.09. IV.1 3 137.83 45.69 45.10 14.67 182.93 59.93

I.1 3 159.77 24.07 31.57 33.20 191.33 40.53

mean 6 148.80 38.13 38.33 26.54 187.13 51.33

K092 15.09. III.1 3 42.70 5.42 247.17 19.19 289.87 13.85

III.6 3 62.00 8.86 214.33 55.52 276.33 51.73

mean 6 52.35 12.13 230.75 44.66 283.10 38.47

K098 13.09. III.7 6 77.30 25.52 114.87 45.99 192.17 69.35

mean 6 77.30 25.52 114.87 45.99 192.17 69.35

K061 13.09. III.2 3 101.80 31.25 99.03 24.24 200.83 29.07

IV.2 3 126.73 52.14 102.53 62.24 229.27 88.69

mean 6 114.27 44.76 100.78 47.26 215.05 67.51

K023 12.09. III.1 6 72.00 31.22 112.08 66.32 184.08 59.78

mean 6 72.00 31.22 112.08 66.32 184.08 59.78

K055 12.09. III.3 3 107.87 33.10 64.10 17.56 171.97 39.39

I.1 3 79.97 19.38 16.33 11.80 96.30 26.10

mean 6 93.92 30.50 40.22 28.18 134.13 50.48

K117 12.09. III.5 3 34.93 18.66 46.63 3.23 81.57 18.51

VI.1 3 8.60 4.52 7.03 4.45 15.63 8.20

mean 6 21.77 18.91 26.83 20.18 48.60 35.94

K126 11.09. III.3 3 69.13 26.76 61.93 11.77 131.07 17.68

III.5 3 38.97 9.19 48.10 10.30 87.07 8.56

mean 6 54.05 25.05 55.02 13.04 109.07 26.02

K011 11.09. III.3 3 192.57 28.66 76.60 13.49 269.17 16.36

III.2 3 90.13 5.44 178.73 36.85 268.87 31.47

mean 6 141.35 55.22 127.67 58.12 269.02 25.08

S023 11.09. VII.1 3 138.90 7.07 78.77 52.91 217.67 49.15

III.3 3 114.77 34.71 127.20 38.66 241.97 71.85

mean 6 126.83 27.80 102.98 52.28 229.82 62.75

B1 24.09. III.3 3 278.83 39.85 344.33 86.88 623.17 49.99

III.2 3 68.00 10.37 279.00 68.86 347.60 68.15

mean 6 173.72 109.07 311.67 84.92 485.38 150.19

B2 23.09. III.3 3 175.90 38.77 215.00 11.22 390.90 45.67

IV.2 3 144.67 71.34 288.00 17.47 432.67 54.17

mean 6 160.28 59.50 251.50 39.34 411.78 54.28

B3 22.09. III.1 3 97.83 23.39 448.17 58.25 546.00 35.31

III.3 3 184.33 70.21 347.00 58.8 541.33 126.63

mean 6 146.08 71.18 397.58 77.38 543.67 92.99

B4 22.09. III.3 3 115.67 45.84 190.33 21.45 306.00 62.12

IV.1 3 139.83 27.48 163.67 50.33 303.50 76.46

mean 6 127.75 39.68 177.00 40.92 304.75 69.67

B5 21.09. III.3 3 113.00 12.68 270.10 18.61 283.10 28.39

IV.1 3 183.83 62.28 287.33 16.01 471.17 68.33

mean 6 148.42 57.22 278.72 19.38 427.13 68.38

B6 20.09. III.3 3 119.00 57.71 223.00 19.80 342.00 66.71

III.1 3 69.33 29.10 264.33 41.07 333.67 52.45

mean 6 94.17 52.01 243.67 38.30 337.83 60.15

B7 20.09. III.1 3 72.17 20.68 198.33 36.28 270.50 31.31

III.5 3 162.17 31.56 277.67 50.31 439.83 51.79

mean 6 117.17 52.32 238.00 59.14 355.17 94.87

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 159

done for various forest habitats in Puszcza Kampinoska, Poland (Kwiatkowska

and Dudziec 1974). This showed unambiguously that significant differences

existed both within and between phytocoenoses. This variability suggests that biomass should be measured in smaller, more uniform structural units of the herb

layer. However, the synusial method is suitable for the description and characterization of such spatial systems. Because of the high patchiness of the herb layer the

synusial approach to the typology of forest communities is inapplicable in the manner once used in Scandinavia (Du Rietz 1930, Lippmaa 1935) and also still used

occasionally in forest communities of other regions (Gillet 1986).

The majority of the types of synusiae described in this work occur in both pine

forests and mixed/pine forests. Among the 14 subtypes distinguished only 4 are

clearly associated with pine forests and three with mixed/pine forests.

Variability in biomass from east to west occurred in both the different types of

herb layer synusiae and in entire study areas. This variability is strongly expressed

in the case of the biomass of mosses, and much more weakly in the case of vascular

plants. Similarly, stronger and more significant correlations were found for pine

forests than mixed/pine forests.

The data presented here on differences in biomass within the synusiae and

different study areas along the west-east transect are a supplement to data from the

literature. The values obtained are lower than those given for biomass in areas

further to the east. Gabeev (1990) showed that pine forests in Western Siberia had

mossy-cowberry herb layers with biomass ranging from 110 to 200 g/m2

 (of this

the biomass of mosses was between 50 and 140 g/m2). The pine forests with bilberry had herb layer biomass in the range of 90 to 130 g/m2

, while the biomass of

mosses was lower – in the range of 15-20 g/m2. In contrast, pine forests had grassy

herb layers of highly variable biomass, with values as high as 304 g/m2 but with

minimal representation of mosses (about 1 g/m2). It should be emphasized that

mean biomass of the herb layer is influenced not only by geographical location, but

by the edaphic conditions of the particular sites. For example, site S023 is characterized by a relatively high mean biomass (230 g/m2

) and at the same time

by a higher (46.3 percent) proportion of species requiring moderately moist

soils. These sites are also characterized by a higher percentage (17.8 percent) of

nitrophilous species.

On the basis of vegetation features analyzed in these studies, we did not detect

any changes at any site that can be explained by air pollution. Future studies of

vegetation should concentrate on population level and such features of plant species as life form, phenology, and type of hemeroby.

Acknowledgments

This work was part of a project entitled “U.S.-Poland Cooperative Project on the

Status and Long-Term Trends in Forest Ecosystems: Climate, Pollution and Forest

Health.” This study was financed by the U.S. Environmental Protection Agency;

the USDA Forest Service, the USDA Foreign Agriculture Service (ICD); the Polish

Academy of Sciences; the Polish Bureau of GEF and the Forest Research Institute;

Michigan Technological University, Houghton, Michigan; Bowling Green State

University, Bowling Green, Ohio, and the Institute of Geography and Spatial

Organization of the Polish Academy of Sciences, Warsaw. We also thank Laurie

Dunn for technical editing of this manuscript.

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Matuszkiewicz, W. 1981. Przewodnik do oznaczania zbiorowisk roslinnych Polski (Guide to determine plant communities of Przewodnik do oznaczania zbiorowisk roslinnych Polski (Guide to determine plant communities of

Poland). PWN, Warszawa; 298 p. Poland)

Mavriscev, V.V. 1980. Sinuzialnaja struktura fitocenozov kislicnoj seri tipov lesa, avtoreferat (Synusial structure of Oxalis

series of forests). Instytut Eksperimentalnoj Botaniki AN BSSR, Minsk; 36 p. series of forests)

Rejment-Grochowska, J. 1950. Watrobowce (Liverworts). PZWS; 68 p. Watrobowce (Liverworts)

Roo-Zielinska, E. [in press]. Fitoindykacyjna ocena warunków siedliskowych stanowisk badawczych wzdluz gradientu

kontynentalizmu (12 - 32 

degrees dl. geogr. wsch.) i zanieczyszczenia powietrza, (Phytoindicational evaluation

of habitat conditions of permanent plots along continental gradient 12-32 degrees East and air pollutant gradi- of habitat conditions of permanent plots along continental gradient 12-32 degrees East and air pollutant gradient). D ent) okumentacja Geograficzna.

Roo-Zielinska, E.; Solon J. [in press] Charakterystyka geobotaniczna zbiorowisk lesnych na stanowiskach badawczych wzdluz

gradientu kontynentalizmu (12 - 32 degrees dl. geogr. wsch.) i zanieczyszczenia powietrza (Geobotanical characteriza- degrees dl. geogr. wsch.) i zanieczyszczenia powietrza (Geobotanical characterization of plant communities on permanent plots along continental gradient 12-32 degrees East and air pollutant gradient). tion of plant communities on permanent plots along continental gradient 12-32 degrees East and air pollutant gradient)

Dokumentacja Geograficzna.

Rothmaler, W. 1976. Exkursionsflora. Kritischer Band. Exkursionsflora. Kritischer Band. Verlag, Berlin: Volk und Wissen Volkseigener; 812 p. Exkursionsflora. Kritischer Band. 

Smialkowski, J. 1998. Transect studies along 52 parallel, 12-32 degrees E: PAR measurements in pure-pine and mixed-pine

forests in Poland. forests in Poland. In: Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan, technical coordinators. Proceedings of forests in Poland. 

the international symposium on air pollution and climate change effects on forest ecosystems; 1996 February 5-

9; Riverside, CA. Gen. Tech. Rep. PSW-GTR-166. Albany, CA: Pacific Southwest Research Station, USDA Forest

Service [this volume].

Solon, J. 1994. Krajobrazowe zroznicowanie roslinnosci rzeczywistej (Landscape differentiation of real vegetation). Krajobrazowe zroznicowanie roslinnosci rzeczywistej (Landscape differentiation of real vegetation). In: Krajobrazowe zroznicowanie roslinnosci rzeczywistej (Landscape differentiation of real vegetation).

Kostrowicki A.S., Solon J., eds. Studium geobotaniczno-krajobrazowe okolic Pinczowa. In: Geobotanical-landscape

studium of Pinczow surroundings). Dokumentacja Geograficzna 1-2: 83-94.

Szafer, W.; Kulczynski S.; Pawlowski B. 1969. Rosliny Polskie (Polish plant species). Rosliny Polskie (Polish plant species). PWN, Warszawa; 1020 p. Rosliny Polskie (Polish plant species).

Szafran, B. 1957. Flora Polska, Mchy (Polish Flora, mosses). Flora Polska, Mchy (Polish Flora, mosses). Part I., PWN Warszawa; 440 p. Flora Polska, Mchy (Polish Flora, mosses).

Szafran, B. 1961. Flora Polska, Mchy (Polish Flora, mosses). Part II., PWN Warszawa; 407 p. Flora Polska, Mchy (Polish Flora, mosses)

Session I1I Geographical Differentiation of the Floristic Composition and Structure of the Herb Layer Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 161

Ewa Roo-Zielinska and Jerzy Solon2

Abstract

The influence of climatic and pollution gradients was determined on species richness, species diversity, and values of

phytoindicators of the herb layer in pine and mixed pine forest communities in Poland. Geographical longitude was

used as a synthetic measure of geographical position. Ellenberg’s indices were used as synthetic measures of habitat

differentiation, calculated as weighed averages: L (light index), T (temperature index), K (continentality index),

F (moisture index), R (reaction index), N (nitrogen index). Richness and diversity variables were used: razem (total

number of species), gatB (number of shrub species), gatC (number of herb layer species), diver (Shannon’s diversity

index H = -∑plogp), simps (Simpson’s index S =∑p2

), beta (beta =[1- simps]/[1-1/gatC]). Additional dependent

variables included the proportion of number and cover of different Raunkiaer’s life forms as well as shares of number

and cover of species of different leaf anatomy and different syntaxonomic character. By using Ellenberg’s indices the

vegetation on consecutive sample plots was not strongly differentiated. All sites on the climatic transect occurred on

dry, strongly acidic or acidic soil, with low nitrogen content. Values for the index of continentality “K” reflected clearly

the climatic differentiation from west to east. The number of herb layer species depended on the geographical longitude

(correlation coefficient 0.851) for pine and mixed forests together, 0.840 for pine forests only, and 0.926 for mixed

forests only. All diversity indices were also strongly correlated with the longitude. Generally, the share of the number of

chamaephytes and therophytes was positively correlated with the longitude. In pine forests there was also a general

tendency for the number of scleromorphic species to decrease (and mesomorphic species to increase) in the west-east

direction. Meadow species also generally tended to increase in the west-east direction. For all the plots taken together

the cover of meadow species is correlated with the longitude (correlation coefficient 0.881), 0.905 for pine forests only,

and 0.910 for mixed forests.

Introduction

Species richness and species diversity are some of the most important structural

features of forest communities. They are influenced by a range of factors such as

large-scale geography and habitat (Cowling 1990, Day and others 1988, Grime 1979,

Jurko 1985, Prusinkiewicz 1970). Other important factors are the history of vegetation

development in the Holocene (Huntley 1993) and the history of use (Law and Morton

1993) to interpopulational and successional interactions (Dollar and others 1992,

Trevin and others 1993). The role of the different factors changes in relation to the

scale of the phenomenon considered.

In analyses on a scale larger than the region, overall geographical factors, including macroclimatic differentiation, are most important. Macroclimatic differentiation consists of many elements, including mean annual temperature, total precipitation, moisture deficit, the amplitude of annual temperatures, the degree of

continentality of the climate and the length of the growing season. These elements

are mutually interlinked and are dependent on geographical location. It is for this

reason also that many studies consider the link between structural features of the

flora and vegetation and geographical location, treating the latter as a synthetic

indicator of macroclimatic differentiation.

This paper defines the relationships between geographical location (which determines macroclimatic differentiation and reflects the history of development of

the vegetation), habitat conditions (Ellenberg 1974), and selected characteristics of

the species richness and diversity of the ground cover in two narrowly-defined

types of forest community, namely, within typical pine forests and within mixed

Herb Layer Diversity and Phytoindicative Evaluation of

Habitat Conditions of Forest Permanent Plots in Poland1

1 An abbreviated version of this paper was

presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Botanists, Institute of Geography and

Spatial Organization, Polish Academy

of Sciences, 00-927 Warsaw,

Krakowskie Przedmiescie 30, Poland.
162 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

forests. The subjects of analysis are 17 study areas lying along a west-east gradient

of continentality (Breymeyer, 1996).

Methods

The species diversity of the herb layer and the phytoindicators of habitat conditions

were obtained from phytosociological records of all species occurring on 400 m2 in

each of the 17plots. The majority of the records were taken in May 1995. A detailed

description of the methodology and of the characterization of the vegetation cover

is given in a separate study (Roo-Zielinska and Solon 1996).

Defined on the basis of information contained in each of the 17 phytosociological records were values for basic indices of diversity, namely: (a) the number of

species of vascular plants; (b) the Shannon Index (Shannon and Weaver 1949), H=

-∑plogp; (c) the Simpson Index (Simpson 1949), S=∑(p2

); (d) the beta index (Simpson

1949) beta=[1- simps]/[1-1/gatC]; and (e) the index of evenness HHmax in the

form HHmax=H/log(n) (where pi is the share of a species in the total cover of

vascular plants in the ground cover layer; n = total number of species in a record).

The number and proportion in cover of species representing different Raunkiaer

life forms, types of leaf permanence, and types of anatomical structure also were

calculated. Detailed characteristics of species were taken from Ellenberg (1974).

The phytosociological structure of the ground cover layer was also described by

using the list of characteristic species from Matuszkiewicz (1981).

The analytical method of indicative numbers from Ellenberg (1974) was used

to assess habitat-climatic conditions (insolation “L” and the degree of continentality

“K”), as well as habitat-edaphic conditions (the moisture “F” and acidity “R” of

soils, as well as their nitrogen content “N”). All the phytoindicative analyses were

done on the floristic composition of the ground layer, which reacts most vigorously

to changes in abiotic conditions, and in which anthropogenic influences become

visible most quickly. The index of insolation “L” indicates the relative strength of

insolation (from 1: species requiring full shade; to 10: plant species requiring full

light). The index of continentality “K” describes the resistance of a species to the

frequency and length of occurrence of dry periods in the growing season, as well as

to the length of the pre-frost period (from 1: the most marked features of an Atlantic climate; to 10: the most marked continental features). The index of humidity “F”

expresses the ecological reaction of species in relation to the humidity of the substrate in the growing season (from 1: those requiring extremely dry soils; to 12:

aquatic plants that are mostly completely submerged in water). The index of soil

reaction “R” expresses biologically the acidity of the substrate experienced by plants

(from 1: species requiring very highly acid soil [pH < 3.5]; to 10: those requiring

neutral or basic soils [pH > 6.5]). The nitrogen index “N” expresses the ecological

reaction of species to the content of nitrogen in the soil (from 1: species only occurring in soils poor in nitrogen; to 10: species only in soils very rich in nitrogen, such

as those intensively fertilized). The mean values of phytoindicators were calculated twice for each of the phytosociological records — the first value was obtained

only on the basis of the number of species of defined divisions of habitat requirements while the second (weighted mean) also included species cover. The method

of Ellenberg and its various applications have often been discussed in the literature. It is useful in both the presentation of the spatial differentiation of current

habitat conditions (Kostrowicki and Wojcik 1971, Roo-Zielinska 1982, 1994) and

very well-suited to the dynamic expression of phenomena (Roo-Zielinska 1993,

Roo-Zielinska and Solon 1990, 1994).

The relationships between geographical location and the values of

phytoindicators (independent variables) and indices of diversity, as well as the proportions of species representative of different ecological groups (dependent variables) were defined on the basis of correlation and regression analyses for three

groups of sites: for all 17 sites on the west-east transect; only for the 9 pine forest

sites on the west-east transect; and only for the 8 mixed forest sites on the west-east

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 163

transect. The Stepwise Variable Selection procedure was used in the choice of the

independent variables of strongest influence on changes in species diversity.

Results

The number of species and species diversity of the vascular plants in the ground

layer increased in the west to east direction within both pine forests and mixed

forests, with the value of the Shannon Index changing from 1.87 to 5.20, the value

of the Beta Index within the range 0.82 to 0.99 and the value of the Simpson Index

varying between 0.35 and 0.03. A similar range of variability was observed in the

index of uniformity (HHmax), which varied from 0.71 to 0.96. The relationships

between indices of richness and diversity and longitude were regularly statisticallysignificant. For all the indices analyzed the absolute correlation coefficients were

above 0.74 (table 1). The correlations between all the indices of richness and diversity

and the Ellenberg indices were considerably weaker when all areas were considered

together. All of them (except the number of lichens) were correlated with the

coefficient of continentality (table 2). In all three groups the relationships between

the value for the index of species richness of the ground layer and the independent

variables (longitude and phytoindicators) are best described by a straight line

relationship gatC=(a*dlugeo)+(b*fito), in which gatC is the number of species of

the ground layer; dlugeo the geographical longitude; fito the phytoindicative indices

denoting: the mean index for light requirements (L) for all species (in the case of all

17 areas considered together or for pine forest only) or the mean index for moisture

requirements (F) also calculated for all species only in the mixed forests (fig. 1).

Another situation arises in the case of the variability of values for the Shannon

Index. In all three groups of sites, the relationships between the value of the Index

and the independent variables is best described by a straight line with longitude as

the dependent variable. However, the modifying influence of local habitat factors

is minimal and not statistically significant (fig. 2).

Variability was low in the proportion of different life forms across the gradient.

Within mixed forests a strong positive correlation was found between the relative

cover of chamaephytes and geophytes in the overall cover and the longitude. A

similar, but weaker, relationship was also observed in the case of joint analysis of

all transect sites (table 1), while differences in habitat conditions exerted a weaker

influence on the number of different life forms (table 2).

Geographical variability in the proportion of species of different leaf longevity

was not apparent. Only in the case of the number and cover proportion of species

with green wintering leaves showed a clear correlation with longitude (table 1).

This relationship is best described by an exponential equation or regression the

form Y=exp(a+bX), albeit with the parameters of the line not differing significantly

between all the analyzed blocks (fig. 3).

Clear geographic variability was found in the proportion of species of different

anatomical types. Positive correlations were found between the quantitative shares

of meso- and hygromorphic species and longitude, while inverse correlations characterized the relationship between the quantitative shares of scleromorphic species

and longitude (table 1). This latter relationship was best expressed by an exponential regression (fig. 4).

Species characteristics of the class Vaccinio-Piceetea were the main components

of the ground cover layer of the areas studied. Their shares in the total number of

species varied from 20 to 50 percent, but they always constituted the dominant

group. The cover of this group varied from 21 to 74 percent of total cover.

The composition of the ground cover of the studied areas also included other

groups of species of which the most important were those characteristic

(Matuszkiewicz 1981) for the classes Molinio-Arrhenatheretea, Sedo-Scleranthetea,

Nardo-Callunetea, Querco-Fagetea, Trifolio-Geranietea, Festuco-Brometea and Epilobietea.

The geographical variation in the numbers of species of differing syntaxonomic

affiliation was different for different groups. Statistically significant correlation

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
164 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

relationships between longitude and the amount of a particular group were more

numerous and stronger in the case of areal representation than in the case of quantitative representation (table 1). The clearest relationship was observed in relation

to the cover proportion of species characteristic of the classes Molinio-Arrhenatheretea

and Vaccinio- Piceetea. This relationship was best expressed by the straight line relationship of the form Y=p1+p2X (fig. 5).

Table 1 Table 1 — Statistically significant correlations between geographical longitude (independent variable) and diversity Table 1 

indices and shares of different species groups.1

All plots Pine forests Mixed forest

on transect on transect on transect

 Number of plots 17 9 8

all vascular species 0.8422*** 0.8506*** 0.9246***

shrub species 0.7903*** 0.8528*** 0.9267***

herb layer

 vascular species 0.8509*** 0.8396*** 0.9260***

beta index 0.8978*** 0.9241*** 0.8823***

Simpson’s index -0.8812*** -0.9156*** -0.8977***

Shannon’s index 0.9058*** 0.9186*** 0.9502***

HHmax 0.8733*** 0.9004*** 0.8686***

by species by species by species by species by species by species

cover number cover number cover number

Herbaceous

 Chamaephytes 0.6047* 0.5013* 0.7919*

Geophytes 0.5209* 0.8212*

Terophytes 0.6911** 0.6953* 0.8363**

evergreen species -0.7409*** -0.9099*** -0.7489*

overwintering

 green species 0.7261*** 0.8363*** 0.7834* 0.8484** 0.8208* 0.8439**

summergreen

 species -0.6691** -0.7871*

helo- and

 higromorphic 0.7528*** 0.8571**

mesomorphic

 species 0.5902* 0.7621*

scleromorphic

 species -0.7856*** -0.8918**

EP 0.5165* 0.8619**

MA 0.8813*** -0.6079** 0.9046*** 0.9096**

NC 0.7567*** 0.7525* 0.8430**

QF 0.7297* 0.7328* -0.7466*

TG 0.6815** -0.7324* 0.8246*

VP -0.6542** -0.7900*

1* = p < 0.05

** = p < 0.01

*** = p < 0.001

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 165

Table 2 Table 2 — Statistically significant correlations between phytoindicative characteristics (independent variables) and Table 2 

species richness of C layer.1

All plots Pine forests Mixed forest

on transect on transect on transect

 Number of plots 17 9 8

by species by species by species by species by species by species

cover number cover number cover number

L -0.5277* -0.5277*

K 0.5702* 0.5563*

F 0.7770*

R 0.5682* 0.5682* 0.7073* 0.7073*

1* = p < 0.05

** = p < 0.01

*** = p < 0.001

Figure 1 — Regression lines

for the relationship between the

number of species of the herb

layer (y axis) and geographical

longitude and phytoindicative

indices (x axis). (A) All plots on

the west-east transect; (B) pine

forests on the transect; (C)

mixed forests on the transect.

Key: dluggeo = longitude; gat C

= number of species; Lsrl = mean

index for light requirements; and

Fsrl = mean index for moisture

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Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
166 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 2 — Regression lines

for Shannon index (y axis) on

geographical longitude (x axis)

according to the linear model

Y=p1+p2X. (A) All plots on the

west-east transect; (B) pine

forests on the transect; (C)

mixed forests on the transect.

Dluggeo = longitude; gat C =

number of species; Lsrl = mean

index for light requirements; and

Fsrl = mean index for moisture

requirement.

Figure 3 — Regression lines

for the share in cover of

overwintering green species (y

axis) on geographical longitude

(x axis) according to the model

Y=exp(p1+p2X). (A) All plots on

the west-east transect; (B) pine

forests on the transect; (C)

mixed forests on the transect.

Figure 4 — Regression lines

for the share of scleromorphic

species (y axis) on geographical

longitude (x axis) according to

the model Y=exp(p1+p2X). (A)

All plots on the west-east

transect; (B) pine forests

on the transect; (C) mixed

forests on the transect.

Figure 5 — Regression lines

for the proportion of cover of

characteristic species of the

Vaccinio-Piceetea class (y axis)

on geographical longitude (x axis)

according to the model

Y=p1+p2X. (A) All plots on the

west-east transect; (B) pine

forests on the transect; (C)

mixed forests on the transect.

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-

-

-

–

-

-

-

-

–

-

-

-

-

–

B –

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

corr = 0.9186

a = -0.3391

b = 0.1697

*

*

*

*

*

*

*

*

5.8

4.8

3.8

2.8

1.8

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

C –

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

corr = 0.9502

a = -0.5736

b = 0.1507

*

*

*

**

*

*

* *

**

*

*

*

*

**

pl

5.8

4.8

3.8

2.8

1.8

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

A –

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

*

Y = exp(a+bX)

corr. = - 0.7759

a = 0.167

b = - 0.072

*

*

*

*

*

*

*

*

*

5.8

4.8

3.8

2.8

1.8

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

B –

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

*

*

*

**

* *

5.8

4.8

3.8

2.8

1.8

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

C –

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

Y = exp(a+bX)

corr. = - 0.6625

a = 0.409

b = - 0.052

Y = exp(a+bX)

corr. = - 0.9218

a = 0.895

b = - 0.096

*

* *

*

*

*

*

*

*

*

* *

*

**

*

Isk

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

A

*

Y = exp(a+bX)

corr. = - 0.8120

a = -0.188

b = - 0.036

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

B Y = exp(a+bX)

corr. = - 0.9056

a = 0.112

b = - 0.048

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

C Y = exp(a+bX)

corr. = - 0.7173

a = -0.452

b = - 0.025

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

-

-

-

-

–

*

*

* *

* *

** *

*

*

*

*

*

*

*

*

*

*

*

*

**

*

*

* *

*

*

*

*

*

pVp

*

0.8

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

A

*

a = 0.816

b = - 0.015

corr. = - 0.6542

*

*

*

*

*

*

*

*

*

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

Isk

0.8

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

B a = 0.840

b = - 0.016

corr. = - 0.7900

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

Isk

0.8

0.7

0.6

0.5

0.4

0.3

0.2

12 16 20 24 28 32 36

longitude

–---–---–---–---–---–---–

–

---

–

---

–

---

–

---

–

---

–

---

–

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

C a = 0.804

b = - 0.015

corr. = - 0.5762

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–-

-

-

-

–

*

*

*

*

*

*

* *

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 167

Results and Conclusions

The transect on the 52° north latitude between 12 and 33° longitude east in Poland

had varying ground cover in sites with pine forest and mixed forest. From west to

east we observed an increase in the total number of vascular plant species per unit

area as well as in the species richness of vascular plants in both the shrub and the

herb layer. However, the number of species of lichens is not correlated with longitude

but is entirely determined by local habitat conditions and the degree of maturity of

the phytocoenosis. An increase was also calculated in values of the Shannon diversity

index and the beta index; and a simultaneous fall was found in the value of the

Simpson Index. In addition, increases were determined for the value of the index

of evenness HHmax, which attests generally to changes in the dominance structure

from species poor to species rich; the number cover of species with green wintering

leaves (with a simultaneous fall in the cover of evergreen species); the number of

mesomorphic species (and a fall in the number of scleromorphic species); and the

proportion of meadow species in the overall cover.

The results clearly confirm the assumption that within a single narrowly-defined

type of forest habitat, longitude is the main factor influencing the species richness and

diversity of the herb layer, as well as its ecological and phytosociological structure. In

spite of this finding, the relationships obtained should be treated as first approximations

of a general model linking the structural characteristics of the community with

environmental conditions. According to Baker (1990), differences in species richness

may only be explained on the basis of a multi-factor model; thus, with any hypothesis

linking species diversity with only one factor, this has limited significance.

In the case of forest communities, the number of species in the ground layer is

shaped not only under the influence of edaphic and geographic conditions, but is

also dependent on the age, character, and structure of the tree stand. In particular,

the character of the herb layer is often determined by the amount of solar energy

reaching the forest floor (Specht and others 1990, Specht and others 1991, Specht

and Specht 1993). Specht and Specht (1989b) showed that the influence of the light

infiltration through the crown is different for different ecological groups of the

herb layer. In particular, increased shading is associated with a linear decrease in

the number of sclerophilous species. Also, the amount, time, and spatial variability of the litter fall may affect the heterogeneity and richness of the herb layer

(Smith 1987). Other factors having a great influence on the richness and ecological

character of the ground cover layer is the dynamic stage and level of anthropogenic impact on the forest community. It has been shown many times that mature

forest communities and/or those little modified from the natural state are characterized by considerably lower proportions of annual and biennial plants when

compared with early developmental stages or phytocoenoses subject to strong

anthropogenic impacts (Dodge and others 1983, Dollar and others 1992, Kirby

1988, 1990, Trabaud and Lepart 1980, Trevin and others 1993, Vankat and Carson

1991). On the other hand, under moderate anthropogenic pressure overall species

richness may increase (Grime 1979, Reader and others 1991).

Acknowledgments

The work was done within the framework of a project entitled “U.S.-Poland

Cooperative Project on the Status and Long-Term Trends in Forest Ecosystems:

Climate, Pollution and Forest Health” financed by the U.S. Environmental

Protection Agency, USDA Forest Service, USDA Foreign Agriculture Service (ICD),

Polish Academy of Sciences, Polish Bureau of GEF, and Forest Research Institute.

Also participating in the financing of the work were: Michigan Technological

University, Houghton, Michigan; Bowling Green State University, Bowling Green,

Ohio, as well as the Institute of Geography and Spatial Organization of the Polish

Academy of Sciences, Warsaw. The present work is not reviewed by the financing

organizations and may not represent their views.

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
168 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

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Prusinkiewicz, Z. 1970. Die Zahl der auf dem Minimi-Areal Vorkommenden Gefasspflanzenarten als Mass fur die Fruchtbarkeit

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Roo-Zielinska, E.; Solon, J. 1996. Charakterystyka geobotaniczna zbiorowisk lesnych na stanowiskach Charakterystyka geobotaniczna zbiorowisk lesnych na stanowiskach badawczych wzdluz

gradientu kontynentalizmu (12 - 32 stopnie dl. geogr. wsch.) i zanieczyszczenia powietrza (Geobotanical characteriza- gradientu kontynentalizmu (12 - 32 stopnie dl. geogr. wsch.) i zanieczyszczenia powietrza (Geobotanical characterization of plant communities on permanent plots along continental gradient 12-32 degrees East and air pollutant gradient). tion of plant communities on permanent plots along continental gradient 12-32 degrees East and air pollutant gradient).

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Specht, R.L.; Clifford, H.T.; Arianoutsou, M.; Bird, L.H.; Bolton, M.P.; Forster, P.I.; Grundy, R.I.; Hegarty, E.E.; Specht, A.

1991. Structure, floristics and species richness of plant communities in southeast Queensland. Structure, floristics and species richness of plant communities in southeast Queensland. Proceedings of the Royal Structure, floristics and species richness of plant communities in southeast Queensland.

Society of Queensland 101: 27-78.

Specht, R.L.; Grundy, R.I.; Specht, A. 1990. Species richness of plant communities: relationship with community growth and

structure. structure. Israel Journal of Botany 39 (4-6): 465-480. structure.

Specht, R.L.; Specht, A. 1989a. Species richness of overstorey strata in Australian plant communities - the influence of overstorey

growth rates. Australian Journal of Botany 37(4): 321-336. growth rates

Specht, R.L.; Specht, A. 1989b. Species richness of sclerophyll (heathy) plant communities in Australia - the influence of Species richness of sclerophyll (heathy) plant communities in Australia - the influence of

overstorey cover. overstorey cover. Australian Journal of Botany 37(4): 337-350. overstorey cover.

Specht, A.; Specht, R.L. 1993. Species richness and canopy productivity of Australian plant communities. Biodiversity and Species richness and canopy productivity of Australian plant communities

Conservation 2(2): 152-167.

Specht, A.; Specht, R.L. 1994. Biodiversity of overstorey trees in relation to canopy productivity and stand density in the

climatic gradient from warm temperate to tropical Australia. climatic gradient from warm temperate to tropical Australia. Biodiversity Letters 2: 39-45. climatic gradient from warm temperate to tropical Australia.

Trabaud, L.; Lepart, J. 1980. Diversity and stability in garrigue ecosystems after fire. Vegetatio 43: 49-57. Diversity and stability in garrigue ecosystems after fire

Trevin, J.O.; Rodney, K.P.; Glasgow, A.; Weekes, N. 1993. Forest change in a subtropical moist forest of St. Vincent. Forest change in a subtropical moist forest of St. Vincent. West Forest change in a subtropical moist forest of St. Vincent. 

Indies: the King’s Hill Forest Reserve. 1945-1990. Commonwealth Forestry Review 72(3): 187-192.

Vankat, J.L.; Carson, W.P. 1991. Floristics of a chronosequence corresponding to old field-deciduous forest succession in

southwestern Ohio. III. Post-disturbance vegetation. southwestern Ohio. III. Post-disturbance vegetation. Bulletin of the Torrey Botanical Club 118(4): 385-391. southwestern Ohio. III. Post-disturbance vegetation.

Session I1I Herb Layer Diversity and Phytoindicative Evaluation of Habitat Conditions Roo-Zielinska, Solon
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 169

Marek Degorski2

Abstract

The lithological and petrographical characteristics of soil pedogenesis was determined, and the spatial and vertical

distribution of some soil physico-chemical properties (including heavy metal content) were studied along two transects

in Poland. The genetic horizon for 22 soil profiles were described for particle size and petrographic composition, quartz

grain abrasion, transformation of organic substance, carbon and nitrogen characteristics, base saturation,

exchangeable cations, and acidity. The results showed that the morphogenesis of parent rock in all studied profiles is

characterized by the glaciofluvial sedimentation of fine sands. All research plots were situated on valley terraces or

outwash planes connected with glaciofluvial accumulation of redeposited, polygenetic sand formation. Soils belonged

to three subtypes of Spodosols (podsolic class). All soil profiles were characterized by a very acid reaction (pH 3.0-4.5

in horizon A), and the degree of acidity corresponded to soil type. All other defined physico-chemical properties indicated

that soils had low nutrient status. Mineral-humus layers of most of the soils were characterized by a high carbon:nitrogen

ratio (from 10.6 to 53.7). This suggests limited biological activity of the soils and slow humification and mineralization

rates. Overlying humus has three distinct subtypes (mor, moder/mor, mor/moder). In all soils the ratio of humic acids

to fulvic acids was less than one, with the highest content of fulvic forms in humus compounds. Total exchangeable

cations averaged 2.5 meq per 100 g of soil from the mineral-humus horizon and 0.5 meq per 100 g of soil in the parent

rock. The degree that the sorption complex was saturated with base cations was also very low (less than 10 percent).

Heavy metal spatial distribution in studied soils was not distinguished by region, but by local pollution sources or by

natural concentration in the soil.

Introduction

Pine forest ecosystems with associated podzolic soils are very common in central

and eastern Europe. For example, in Poland the potential natural plant association

of coniferous communities (Vaccinio-Piceetea) occupies about 25.3 percent of the total

area of the country (Matuszkiewicz 1991). Similarly, the same area of Poland is

characterized by podzolic soils (Spodosols). The highest concentration of podzolic

soils (39.7 percent) is in the central Polish lowland (Prusinkiewicz and others 1980).

All the study plots of both the climatic and Silesian transects in Poland (Breymeyer,

this volume) are located on podzolic soils. Research was done in 22 soil profiles

(Breymeyer and others 1995, Breymeyer 1996).

This study determined the geographical differentiation of lithological and petrographical conditions of the pedogenesis of soils and analyzed the spatial and

vertical distribution of some soil physico-chemical properties, including the content of heavy metals.

Methods

Base soil profiles and diagnostic excavations were done at study areas selected in

accordance with the concept of the research. The basis for pedological research was

the diagnosis of the different genetic horizons of 22 soil profiles on two research

transects. For each profile, soil material was collected for analysis from different

genetic horizons. A series of determinations were carried out concerning the

morpholithogenic properties of the soil substratum (Degorski 1994a, b; Degorski

[In press a]), the overlying humus and the organic matter in the mineral-humus

layers (Degorski and Jefriemow [In press]), the physico-chemical properties of soils

Spatial and Vertical Distribution of Soil Physico-Chemical

Properties and the Content of Heavy Metals in the

Pedosphere in Poland1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Geographer-Ecologist, Institute of

Geography and Spatial Organization,

Polish Academy of Sciences, 00-927

Warsaw, Krakowskie Przedmiescie 30,

Poland.
170 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

(Degorski 1995a, b; Degorski [In press b]), and the heavy metal content of the soils

(Gworek and Degorski [In press]) by using standard soil science methods.

Differences and similarities between the morpholithological features of the study

sites were analyzed, as well as the physico-chemical features causing the breakdown

of heavy metals and trace elements. Euclidean distance and Ward’s method (Batko

and Moraczewski, 1987) were used. Trace elements and heavy metals (Zn, Mn, Fe,

Pb, Ni, Cr, and Cu) were determined using extraction with 20 percent HCl, after

the combustion of organic matter at a temperature of 480 °C. Concentrations of

these elements were determined by atomic adsorption spectrophotometry (AAS).

An accumulation index was calculated from the data for all mineral-humus

(A), eluvial (E) and illuvial (I) horizons. The accumulation index was calculated

as the ratio of the content of a given element in the genetic horizon and in the

parent rock, giving an indication of exogenic origin in the surface layers of the

soil (Gworek 1985).

Results

Physico-Chemical Characteristics of the Soils

All of the soils studied were developed in redeposited, polygenic sandy

formations that accumulated in the Pleistocene and Holocene. These

differed only in the age of sedimentation, which is associated with the late

Pleistocene (fig. 1).

The studied habitats of pine forests and mixed pine forests developed on formations of river accumulation terraces or glaciofluvial forms (Degorski [In press

a]). All of the studied soil samples were loose and poor-clay sands of moderate or

varied grain size and with clear features of presorting, as well as limited contents

of skeletal fractions (over 1 mm). The surface layers of soils located on terrace areas

(e.g., K092, K098, KB001, and S126) were characterized by increased proportions in

their mechanical composition of silty and dusty fractions. This was caused by an

increase in deflational processes that took place in these areas in periglacial periods. The fluvial character of the sediments and their limited surface aeolization

was confirmed by granulometric indices calculated on the basis of their mechanical composition (Degorski [In press a]). Cumulative curves characterizing the

fluvial material and constructed on Phi probability plots (Program Analiza

Uziarnienia 2.0 - Mechanical Composition 2.0) depart clearly from the log-normal

distribution. This characterizes a leptokurtic breakdown (GSP 1.5-1.9) because curves

for sands with features scattering over a considerable interval resemble straight

lines and have a distribution close to the mesokurtic (GSP 1.3-1.4) (fig. 2). All of the

studied soils are Spodosols. Morphological features and chemical criteria allow for

their inclusion within three subtypes: podsols, rusty podsols, and podsolic rusty.

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski

Figure 1 — Age of last sedimentation for soil substratum.

Age of sedimentation

Germany West Central East Belarus

 Poland Poland Poland

600

500

400

300

200

100

0

thousand years

10.9 13 18 340 500 -

----
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 171

All of the profiles studied are characterized by very acid reactions. For the mineralhumus layers, pH values ranged from 3.0 to 4.5, with the degree of differentiation

corresponding to the type of soil (fig. 3). In addition to acidic reactions, other analyzed physico-chemical properties of the soils also indicate low nutrient status.

Total exchangeable cations (V) amounted on average to 0.5 meq per 100 g of soil in

the mineral-humus layer. The degree of saturation of sorption complexes with base

cations is very low in the mineral-humus horizons and around 20 percent in the

parent rock (fig. 4). All of the soils studied on both transects are also characterized

by very broad ratios of organic carbon (C) to total nitrogen (N). The C:N ratio in

mineral-humus horizons amounts to values of about 10.6 to 53.7, indicating very

limited biological activity of the soil and slow processes of humification and mineralization (fig. 5).

Figure 2 — Soil texture

characteristics defined for

research site KS011, horizon-B.

a - i = particle size fractions:

a = stones and gravel (over 1

mm);

b = coarse sand (0.5 - 1 mm);

c = medium sand (0.25 - 0.5

mm);

d = fine sand (0.1 - 0.25 mm);

e = very fine sand (0.05 - 0.1

mm);

f = coarser silt (0.02 - 0.05 mm);

g = silt (0.005 - 0.02 mm);

h = fine silt (0.002 - 0.005 mm);

i = colloidal clay (below 0.002);

perc. = percentile.

Pct. 1 = - 2.82

Pct. 5 = - 1.00

Pct. 16 = 0.26

Pct. 25 = 0.62

Pct. 50 = 1.31

Pct. 75 = 1.86

Pct. 84 = 2.41

Pct. 95 = 3.86

(c) P.P.

(PHI)

99 pct.

95

50

5

1

0.01

- 6 - 4 - 2 0 2 4 6 8 10

100 pct.

80

60

40

20

0

–

–

–

–

–

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski

a b cd e f g h i
172 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Analysis of organic matter also pointed to the poverty of the studied soils and

slow processes of humification. The ratio of humic to fulvic acids is less than 1 in all

the studied soils, indicating humus of the first type in the Kononowa classification

(1968), i.e., that with a low degree of condensation of aromatic ring compounds

(Degorski 1994a, b; Degorski 1995a, b). On the basis of statistical analysis of the

results for physico-chemical properties, properties of organic matter, humus type

and properties of the substrate, the studied soils may be divided into three subgroups differing from one another in a statistically significant way (figs. 6, 7). These

groups are linked with the genetic type of the soil (podsols, rusty podsols, or podsolic

rusty), with the type of the overlying humus (mor, moder/mor, mor/moder), and

with the mean characteristics of physico-chemical properties.

The first group includes podsol soils with mor-type humus and the lowest pH

values (3.0-3.5). These are characterized by the lowest rates of mineralization and

humification (the greatest thickness of overlying humus and the widest ratio for

humic to fulvic acids). The second group are rusty podsol soils, with moder/mor

type humus of lesser thickness and with better sorption properties. The third group

are rusty with poorly-marked processes of leaching (a lack of a clear eluvial horizon). The overlying humus is of the mor/moder or moder types and has the least

thickness and the highest rate of mineralization and humification of organic compounds (the narrowest C:N ratio). The low carbon content together with the high

Figure 3 — Relationship between pH and base saturation

(V) on two transects.

--------

25

20

15

10

5

0

Silesian Transect

Climatic Transect Poland

126 023 005 060 002 011S 011K 028

pH v

6 6.9 6.1

9.9

7.5

21.7

2.5 3 3.6 3.6 3.2 3.5 4.5 4.2

2.5 3.3

--------

16

14

12

10

8

6

4

2

0 K098 K061 K126 K023 K055 K092 K1117 KS011

pH v

3.2

6.3 7

8.5 8.8 8.6

13.2 13.2 13.9

3.3 3.7 3.8 3.8 3.9 4 4.2

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 173

degree of humification and the greater content of humates and humic acids (the

narrowest Ch:Cf ratio) all indicate soils with significantly more advanced evolution of organic matter. The sorption properties of this group of soils are also the

best among those studied. The degree of saturation of the sorption complex with

base cations increases from 10-20 percent in the mineral-humus layers to about

30 percent in the parent rock, while the reaction (pH) varies from about 4 to 4.5 in

the mineral-humus horizons (fig. 7).

Figure 4 — Percent base

saturation for whole soil

horizons (A, E, B, C) and

research plots.

Figure 5 — C:N ratio in soils

for each soil horizon.

50

40

30

20

10

0

pct

S060

S028

K092

S011

S005

S002

S126

A

B

E

Plots C

C:N

K098

K061

K023

K055

K117

K126

KS011

S023

V

30

25

20

15

10

5

0

pct

S060

S028

K092

K098

K061

K023

K055

K117

K126

KSO11

S023

S011

S002

S005

S125 A

B

E

C

Plots

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski
174 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The Breakdown of Heavy Metals and Trace Elements

There is no statistically-significant basis for claims regarding geographical-scale

variation in contents of heavy metals and trace elements along the two transects.

The only differences observed were in relation to regional distributions. The increase

in the contamination of a soil with some element is most often caused by some

local source of pollutant emissions. A good example of this situation is the spatial

distribution of the lead content in soils along the two transects (fig. 8). The highest

level of lead was recorded at point S028 at Klucze near Olkusz. This is traditionally

Figure 6 — Similarity of the

soil properties of two transects

(Euclidean distance/Ward’s

Method) (Batko and

Moraczewski 1990). a = climatic

transect, b = Silesian transect.

Figure 7 — Selected physical

and chemical properties of the

mineral-humic horizon defined

for selected soil groups, pH =

acidity determined in H2

O; V =

saturation degree of the sorption

complex with cations of basic

character in percent; Hum =

contents of humines; A =

thickness of the A horizon in

cm; O = thickness of the O

horizon in cm; SH = humification

degree in percent; Ch:Cf = the

ratio of humine to fulvic acids x

100 percent. A = podzolic soil,

B = rusty-podzolic soil, C = rusty

soil

A

B

1 -9 -8 -7 -6 -5 -4 -3 -2 -1 0

-----------

––––––––––

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . .

092

011

117

098

061

023

126

055

100

1 -9 -8 -7 -6 -5 -4 -3 -2 -1 0

-----------

–––––––––––

S126

S005

S023

S060

S002

S028

S011

K011

60

50

40

30

20

10

0

A B C

Ch:Cf

Degree of hum.

C:N

O (cm)

A (cm)

Hum.

V (pct)

pH

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 175

and remains a region in which zinc and lead ores are extracted and processed. The

contents of lead in the remaining points analyzed were linked to both the location

of industries and transportation routes.

The indices of accumulation defined for all of the studied soils confirmed ideas

concerning the spatial distribution of the contents of heavy metals and trace elements in the profiles. The greatest concentration occurs in the levels of overlying

humus (O) and of the accumulation-humus horizon (A). The highest values for the

index were attained for lead (to about 15 times the content of this element in the

parent rock) and manganese (to about 8 times the content) (fig. 9). The distribution

of concentrations of heavy metals in the upper layers of the soil differed from those

of the chemical background in the parent rock, indicating specific geographical

relationships. The content of zinc is clearly higher in Upper Silesia in connection

with the concentration of industry in this region. In contrast, the contents of nickel,

iron, and chromium were clearly greater in the mineral-humus horizons of western Poland. The higher content of these three elements in the soils of the western

part of the country may point to the transfer of pollution from western Europe,

which evidently has an impact on the heavy metal contamination of the soils in this

part of Poland. Such a finding is in accordance with the dominant wind direction

in Poland (westerly and north-westerly).

Conclusion

All of the research sites were linked morphogenetically with the accumulation of

redeposited, polygenetic sand formations that occurred in the Pleistocene. Soils on

both transects belong to three subtypes of Spodosols: podsolic, rusty podsolic, and

rusty. All of the studied profiles were characterized by high acidity (pH of 2.5 to

4.5) and a degree of variation in this feature that corresponds to soil type. All other

defined physico-chemical properties were also indicative of the fact that soils were

of low nutrient status. Many of the soils were characterized by mineral-humus

layers with a high C:N ratio (ranging from 10.6 to 53.7). This attests to the low

nutrient cycling rates of the soils and the slowness of the processes of humification

and mineralization. The lowest values of humic acids and humines were determined

in podsolic soil, suggesting that mineralization rates are lowest in these soils.

Figure 8 — Spatial variability

of lead distribution on two

research transects. Bars are

showing Pb mg/1000 g of soil.

70

60

50

40

30

20

10

0

S028 K098 K061 K023 S060 K100 S126 K011 S005 K126 K055 K117 S023 S011 K092 S002 -

---------------

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski
176 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 9 — Accumulation index of heavy metals and trace elements.

250

200

150

100

50

0 K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

-

-

-

-

-

- -

--------------

10

8

6

4

2

0 K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

-

-

-

-

-

- -

--------------

B E A O

plots

-

-

-

-

-

-

B E A

plots

K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126 -

--------------

2.5

2

1.5

1

0.5

0

2

1.5

1

0.5

0

-

-

-

-

-

100

80

60

40

20

0 K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

-

-

-

-

-

- -

--------------

B E A O

-

-

-

-

-

-

-

-

7

6

5

4

3

2

1

0 K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126 -

--------------

plots

10

8

6

4

2

0 K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

B E A O

plots

-

-

-

-

-

- -

--------------

Zn Ni

Mn Cr

B E A O

plots

B E A

plots

K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126 -

--------------

Fe Cu

B E A O

plots

Pb

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski

pct

pct

pct

pct

pct

pct

pct
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 177

Acknowledgments

This study was part of a project entitled “U.S.-Poland Cooperative Project on

the Status and Long-Term Trends in Forest Ecosystems: Climate, Pollution and

Forest Health.” This was financed by the U.S. Environmental Protection Agency,

USDA Forest Service, USDA Foreign Agriculture Service (ICD), Polish Academy

of Sciences, Polish Bureau of GEF, Forest Research Institute, Michigan

Technological University, Bowling Green State University, and the Institute of

Geography and Spatial Organization of the Polish Academy of Sciences,

Warsaw. I thank Laurie Dunn for technical editing of this manuscript.

References

Batko, A.; Moraczewski, I. 1990. Tytan 97, Pakiet programow realizujacy analize skupien. wersja 1.2, Warszawa; 48 p. (in Tytan 97, Pakiet programow realizujacy analize skupien

Polish).

Breymeyer, A. 1996. Badania borow na transekcie klimatycznym wzdluz rownoleznika 52° (12-32°) i na transekcie

zanieczyszczenia: zalozenia ogolne, uwarunkowania geograficzne i ekologiczne. zanieczyszczenia: zalozenia ogolne, uwarunkowania geograficzne i ekologiczne. Warszawa: Dokumentacja Geograficzna. zanieczyszczenia: zalozenia ogolne, uwarunkowania geograficzne i ekologiczne.

Breymeyer, A. 1998. Transect studies on pine forest along parallel 52°N, 12-32°E and along a pollution gradient: general E and along a pollution gradient: general

assumptions. In: Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan, technical coordinators. Proceedings of the assumptions

international symposium on air pollution and climate change effects of forest ecosystems; 1996 February 5-9; Riverside,

CA. Gen. Tech. Rep. PSW-GTR-166. Albany CA: Pacific Southwest Research Station, USDA Forest Service [this volume].

Breymeyer, A.; Degorski, M.; Roo-Zielinska, E.; Solon, J.; Smialkowski, J. 1995. Biodiversity and matter cycling in forest eco- Biodiversity and matter cycling in forest ecosystems in the gradient of air pollution in Poland; results of introductory research. systems in the gradient of air pollution in Poland; results of introductory research. In: Paschalis, P.; Rykowski, K.; systems in the gradient of air pollution in Poland; results of introductory research.

Zajaczkowski, S., eds. Protection of forest ecosystems biodiversity of Bialowieza primeval forest. Warsaw; 175 - 191.

Degorski, M. 1994a. Characterization of soils on research transect associated with Poland’s (West-East) climatic gradient. Characterization of soils on research transect associated with Poland’s (West-East) climatic gradient.

Report, Warszawa: IGiPZ PAN; 92 -142.

Degorski, M. 1994b. Charakterystyka gleb na transektach badawczych zwiazanych z gradientem zanieczyszczen w Polsce na

linii poludniowy-zachod, polnocny-wschod (Characteristic of soils situated on the transect connected with pollution dis- linii poludniowy-zachod, polnocny-wschod (Characteristic of soils situated on the transect connected with pollution distribution in Poland). In: Bioroznorodnosc i krazenie materii w ekosystemach lesnych ustawionych w gradiencie tribution in Poland)

zanieczyszczenia powietrza na obszarze Polski. Report, Warszawa: IGiPZ PAN; 150 - 169.

Degorski, M. 1995a. Some aspects of soil conditions on a research transect connected with distribution of pollution in Po- Some aspects of soil conditions on a research transect connected with distribution of pollution in Poland, In: Preliminary studies of polluted forest ecosystems in Poland (from Upper Silesia to Bialowieza). Report, Warszawa: land, In: Preliminary studies of polluted forest ecosystems in Poland (from Upper Silesia to Bialowieza)

IGiPZ PAN; 46 - 56.

Degorski, M. 1995b. Differentiation of the soil types and characteristic of their basic properties as an effect of climatic and

pollution gradient (“climatic” and “Silesian” transects) pollution gradient (“climatic” and “Silesian” transects) In: Effects of atmospheric deposition and climate change in forest pollution gradient (“climatic” and “Silesian” transects) 

ecosystems in Central and Eastern Europe and the United States, Report, Warszawa: IGiPZ PAN; 27 - 30.

Degorski, M. [In press a]. Charakterystyka morfo-litogeniczna i glebowa siedlisk borow i borow miesznych na transektach

badawczych. badawczych. badawczych. (Morpho-lithological characteristic and soil description of pine and mixed pine forest on two research

transect). transect). Warszawa: Prace Geogr. transect).

Degorski, M. [In press b]. Zroznicowanie fizyko-chemicznych wlasciwosci gleb siedlisk borow i borow mieszanych

na transektach badawczych. na transektach badawczych (Differentiation of physico-chemical soil properties of pine and mixed pine forest). (Differentiation of physico-chemical soil properties of pine and mixed pine forest).

Warszawa: Prace Geogr.

Degorski, M.; Jefremov, A. [In press]. Wlasciwosci prochnicy nadkladowej, biomasa mikroorganizmow i zawartosœ kwasow

nukleinowych w glebach siedlisk boru i boru mieszanego na transekcie zwiazanym z gradientem kontynentalizmu. nukleinowych w glebach siedlisk boru i boru mieszanego na transekcie zwiazanym z gradientem kontynentalizmu (Humus properties, biomass of microorganisms and the content of nucleic acids in soils of pine and mixed pine forest on the

transect along continentality gradient). transect along continentality gradient). Warszawa: Prace Geogr. transect along continentality gradient).

Gworek, B. 1985. Pierwiastki sladowe w glebach uprawnych wytworzonych z glin zwalowych i utworow pylowych polnocno- Pierwiastki sladowe w glebach uprawnych wytworzonych z glin zwalowych i utworow pylowych polnocnowschodniego regionu Polski. (Trace elements in cultivated soils developed from boulder loams and silty formations of the

northeastern region of Poland). northeastern region of Poland). Rocz. Gleb. 36(2): 43-59. northeastern region of Poland).

Gworek, B.; Degorski, M. [In press] Przestrzenny i profilowy rozklad pierwiastkow sladowych i zelaza w glebach zbiorowisk

borowych. (Spatial and vertical variability of trace elements and iron in soil of pine association). borowych. (Spatial and vertical variability of trace elements and iron in soil of pine association). Roczniki Gleboznawcze. borowych. (Spatial and vertical variability of trace elements and iron in soil of pine association).

Gworek, B. 1985. Pierwiastki sladowe w glebach uprawnych wytworzonych z glin zwalowych i utworow pylowych polnocno- Pierwiastki sladowe w glebach uprawnych wytworzonych z glin zwalowych i utworow pylowych polnocnowschodniego regionu Polski. (Trace elements in cultivated soils developed from boulder loams and silty formations of the

northeastern region of Poland). northeastern region of Poland). Rocz. Gleb. 36(2): 43-59. northeastern region of Poland).

Kononowa, N. 1968. Substancje organiczne gleby, ich budowa, wlasciwosci i metody badan (Soil organic matter, their struc- Substancje organiczne gleby, ich budowa, wlasciwosci i metody badan (Soil organic matter, their structure, properties and study methods). Warszawa: PWRiL. ture, properties and study methods)

Matuszkiewicz, W., 1991. Szata roslinna Szata roslinna (Vegetation). (Vegetation). In: Starkel, L., ed. Geografia Polski, srodowisko przyrodnicze, PWN, (Vegetation).

Warszawa: 445-494.

Prusinkiewicz, Z.; Bednarek, R.; Pokojska, U. 1980. Gleby bielicoziemne w Polsce. Gleby bielicoziemne w Polsce (Podzolic soils in Poland). (Podzolic soils in Poland). Roczn. Gleb. 32 (Podzolic soils in Poland).

(1): 103-113.

Session I1I Spatial and Vertical Distribution of Soil Physico-Chemical Properties and the Content of Heavy Metals Degorski

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 179

Alicja Breymeyer,2 Marek Degorski,2 and David Reed3

Abstract

The relationship between litter decomposition rate, some chemical properties of upper soil layers (iron, manganese,

zinc, copper, lead, mercury, nickel, chrome in humus-mineral horizon-A), and litter (the same eight elements in needle

litter fraction) in pine forests of Poland was studied. Heavy metal content in organic-mineral horizon of soils was

highly correlated with needle decomposition rate for copper (correlation coefficient 0.901), zinc (corr. coeff. 0.901),

nickel (corr. coeff. 0.834), and iron (corr. coeff. 0.850). Heavy metal content in needle litter was highly correlated with

needle decomposition rate (iron—corr. coeff. 0.83; zinc—0.80; lead—0.82; nickel—0.89), and with mixed litter

decomposition rate (iron—corr. coeff. 0.71; zinc—0.66; lead—0.68; nickel—0.72). Significant correlations were not

found between the rate of decomposition of wood or cones and their content of metals, except for chromium. When

comparing decomposition rates with degree of litter pollution, it was shown that in the case of needle decomposition

for six significant correlations, two were positive; in the case of mixed litter decomposition for five significant correlations,

two were positive. In both cases decomposition reacted positively to the presence of iron and lead. When comparing

decomposition rates with the degree of soil pollution, it was shown that in the case of needle decomposition, five

significant correlations were positive; in the case of mixed litter decomposition, only one significant correlation was

positive.

Introduction

Although the level of pollution in Poland has declined steadily in recent years

(Breymeyer 1996, Przybylska 1995, Statistical Yearbook of Katowice Voivodeship

1992), pine stands have continued to be damaged by it (Wawrzoniak and

Malachowska 1996, 1997). The saturation of litter layers and soils with heavy

metals is still at a level harmful to the proper functioning of the forest ecosystem

(Laskowski 1994).

Research on soils and the rate of decomposition of forest litter was carried out

at 15 sites with pine forest (Leucobryo-Pinetum and Peucedano-Pinetum) or mixed

pine forest (Quercoroboris-Pinetum) growing on the podsolic soils developed on the

glaciofluvial and fluvial deposits (Degórski 1996a). The sites were distributed along

two transects: one west-east across Poland at parallel 52° N (climatic or continental

transect), and the other from Upper Silesia to the Bialowieza area (Silesian transect)

(Breymeyer 1996).

The sites were slightly different in annual precipitation, while mean annual

temperature was distinctly lower to the east (Smialkowski 1996).

The chemical properties of overlying humus layers were studied, as well as the

mineral-organic horizons of soils, where organic matter accumulates and is mineralized and humificated. The sorption complexes are most vulnerable to the accumulation of heavy metals, and their buffering properties determine the geochemical balance of the whole ecosystem. Permanent circulation of soil organisms between upper soil horizons, humus, and litter creates one functional unit of layers.

This study analyzed the relationship between the rate of decomposition of forest litter and the chemical characteristics of the mineral-humus horizon-A of soils

by using deposition measurements of pollutants from local emissions in both Upper Silesia and the continental transect sites in Poland.

Transect Studies on Pine Litter Organic Matter:

Decomposition and Chemical Properties of Upper

Soil Layers in Polish Forests1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Ecology, and Soil Scientist, respectively, Institute of Geography and Spatial Organization, Polish

Academy of Science, Krakowskie

Przedmiescie 30, 00-927 Warsaw.

3 Professor of Forest Biometrics, Michigan Technological University,

Houghton, Michigan 49931.
180 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Methods

Rates of decomposition were measured using the common litter bags method (i.e.,

Pan-European Forest Ecosystem Research Program; Berg and Co 1993). Weighed

portions of litter dried to constant weight were enclosed in bags (10 by 10 cm)

made from 1 mm nylon netting. Bags were then located at the sites so that

decomposition could take place. The surfaces on which the bags were placed were

cleaned of moss and small shrubs to ensure that the exposed litter made contact

with the upper layer of the soil. Sixty litter bags were placed simultaneously at

each site: 20 with 1g of pine needles each, 20 with 1g of mixed litter, 10 with a single

pine cone, and 10 with 1g of small twigs. The exposure of litter bags always began

in autumn (October), with samples collected after two time periods: half after winter

(in March or April) and the remaining half during the following autumn (October),

when samples for the next year were placed on the soil. In every case only litter

that had fallen to the forest floor during the previous year was used (litter plots

were cleaned before the counted fall). Litter was collected from special demarcated

areas, ensuring that the litter collected was always “of the same age.” After field

incubation, litter bags were transferred to the laboratory and cleaned of soil and

plants overgrowing the netting. Litter was then dried to constant weight and

weighed. The mass loss during incubation provided a measure of decomposition

expressed in percentage terms or as the so-called “k” coefficient, which was

calculated by applying the formula:

ln (final mass / initial mass) = -k

Sinusoidal transformation of the results obtained for decomposition was also applied:

(sin-1 (x1/2).

The pollution of the organic-mineral horizon-A of the soil was estimated on

the basis of analyses of soil samples collected on a single occasion from 10 points

on each mineral-organic horizon of a given profile, with the mean value presented.

Contents of heavy metals, such as zinc (Zn), manganese ( Mn), iron (Fe), lead (Pb),

nickel (Ni), chromium (Cr), and copper (Cu), in soil organic matter were determined using an atomic adsorption spectrophotometer (AAS) after digestion of

samples in 20 percent HCl. An index of accumulation was calculated on the basis of

the results obtained for all mineral-humus horizon-A as a measure of their enrichment by the heavy metals. The index of accumulation was calculated as the ratio of

the content of a given element in the organic-mineral horizon-A and in the parent

rock, giving an indication of an exogenic origin n the surface layer of the soil

(Gworek 1985; Degorski, [In press]).

Litter (needles, twigs, and cones) collected annually from all sites was chemically analyzed for the concentration presence of eight metals as described above

for soil organic matter. Statistical calculations involved analysis of variance for the

10 elements and the 3 fractions.

Results

The Pollution of Litter

The concentrations of Pb and K were not statistically different among the litter

fraction (p > 0.05). Concentrations of the remaining elements (Fe, Mn, Zn, Cu, Cd,

Ca, Mg, and Na) did vary significantly from one litter fraction to another (p < 0.05).

Some trends in the composition of litter along both transects were visible. Two

clear groups of elements were present in pine needles (the dominant fraction in

litter): Zn, K, and Na were at significantly lower levels along the transect of

continentality; and Mg, Mn, Cu and Cd occurred at lower levels along the Silesian

transect. The remaining three elements (Pb, Fe, and Ca) did not show clear trends.

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 181

The three fractions of litter (needles, twigs, and cones) did not show significant

differences in the contents of the elements Pb and K (p > 0.05). Contents of all the

remaining elements were variable.

It is not possible to determine whether the different concentrations of metals in

the different fractions of litter resulted from the accumulation of elements within

tissues or on the surfaces of dead fragments of the plants forming it; litter was not

washed prior to analysis, so both possibilities remain. Therefore, we focused on only

the existence of different accumulations of metals by the three litter fractions.

The Pollution of Soils

Indices for the accumulation of eight metals in the organic-mineral horizon-A of

soils were determined from all 15 study sites (fig. 1). In western Poland and Silesia,

manganese, nickel, and copper occurred in concentrations exceeding those in the

parent rock (about six times), while Fe, Zn, and Cr did not. The lowest concentrations of the six metals were observed in eastern Poland. Lead accumulated more

intensively in soil than the other metals (fig.1). Lead concentrations may be 100 to

200 times higher in soil than in the substratum. The pollution of the environment

by lead is primarily caused by vehicles using leaded gasoline. However, none of

the sites in this study were in the vicinity of busy stretches of road, only within

forested areas. The source of the lead in these soils must be from another local

source.

The indices of accumulation defined for all of the studied soils confirmed the

spatial distribution of heavy metals and trace in organic-mineral horizon; the highest values for the index were attained for Pb, Mn, and Cu (fig. 1).

Figure 1 — Accumulation index for organic-mineral horizon-A of some chemical elements (A) with volume between 0-10 percent, and

(B) with value over 10 percent (Breymeyer 1996). K= continental transect S= Silesian transect.

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed

1

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

Silesian transect, 10.94-03.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

-0.8

Silesian transect, 10.94-09.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

Climatic transect, 10.94-03.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

-0.8

Climatic transect, 10.94-09.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

NUTRIENTS HEAVY METALS

needles wood cones mixed

CORRELATION INDEX
182 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The Decomposition of Litter

The annual rate of decomposition of litter amounts to just over 30 percent and is

higher in the first half of the exposition year (fig. 2). The mean annual rates of decomposition for the two transects were not statistically different. The course of these

means over time showed a 4 percent difference between the transects (with more

rapid decomposition on the climatic transect) after half a year. This difference had,

however, disappeared after 12 months. The sites along the Silesian transect showed

greater differences in the course of decomposition.

Analysis of the rates of decomposition of the different fractions of the litter

revealed that the rate of breakdown of the woody fractions (cones and twigs) was

clearly lower than the mixed litter, especially needles (fig. 2). After 12 months needle

litter mass decreased more than 30 percent, compared to about 20 percent of mixed

litter and 10 percent of the mass loss of cones and twigs.

Figure 2 — Decomposition

rates during the incubation year.

The incubation year always begins

in October when previous- year

litter-fall packed into nylon-net

bags is exposed in the stands; the

bags are collected after 6 months

and 12 months. (A) Average

differences among litter fractions

across all sites on continental

transect. Solid line represents the

mean values. (B) Differences

among sites on Silesian transect.

10

8

6

4

2

0

-

-

-

-

-

- -

--------------

Pct

K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

PLOT NUMBERS

Fe Mn Zn Cu Ni Cr

A

250

200

150

100

50

0

-

-

-

-

-

- -

--------------

K92 K98 K61 K23 K55 K117 K126 KS11 S23 S11 S02 S60 S05 S28 S126

Pct

B

PLOT NUMBERS

Pb

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 183

Concentration of Metals in the Mineral-Organic

Horizons of Soils and in Litter

Correlation index values between accumulated heavy metals in soils and the rates

of litter decomposition were calculated (figs. 3, 4). The first consistent and notable

result was the high prevalence of positive correlations between the breakdown of

litter and the concentration of a metal in the soil. Among the calculations for material from the two transects for decomposition in winter and the whole year, a negative correlation was only noted for lead (fig. 3). Among the calculations for each

transect separately, negative values for the coefficient were again noted for lead, as

well as for nickel and chromium (fig. 4). In consequence, the rate of litter decomposition and the accumulation of a metal in the upper layer of the soil are most often

correlated positively.

Correlation index values between rates of litter decomposition and concentrations of the 8 elements were determined (table 1). A prevalence of positive correlations was again noted, but greater negative values were found than for soil correlations. Thus, pollution of the litter may hold back the process of decomposition much

more often than the pollution of soils.

Figure 3 — Correlation index

between heavy metals in organicmineral soil horizon-A and litter

decomposition rate (litter bags).

Mean values for two transects.

1 A sin-1(X1/2) transformation was performed before analyses; x = only winter (1995) exposure period.

2 -p<0.001.

3 -p<0.05.

Table 1 Table 1 — Correlations between needle litter element concentrations and decomposition rate of different forms of organic Table 1 

deposition for the annual (1994-1995) exposure period on both the climatic and Silesian transects.1

Element Element Mixed litter Needles Cones Wood

 in litter

Fe 2 0.71 2 0.83 0.06 - 0.38

Mn 3- 0.42 3 0.44 0.01 0.13

Zn 2- 0.66 2- 0.80 -0.08 0.4

Cu - 0.25 - 0.34 -0.08 0.24

Pb 2 0.68 2 0.82 0.08 - 0.4

Cd 0.06 0.13 0.07 - 0.14

Ni 2

- 0.72 2- 0.89 -0.08 0.36

Cr 2- 0.51x 2- 0.42x 2- 0.56x 2- 0.63

Remaining Biomass (Pct)

100

90

80

70

60

50

–

-

–

-

–

-

–

-

–

-

–

0 6 12

–––

Months of Exposure

Needles Wood Cones Mixed

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed
184 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

These results indicate that a higher concentration of some of the heavy metals

in the studied organic and mineral-organic soil horizons of poor coniferous forest

habitats positively influence the transformation of organic matter.

Conclusions

The study found that decomposition of pine needles is similar along the climatic

and Silesian transects. The mean rate is higher along the climatic transect in the

first 6 months of the year, but mean rates for the transects do not differ significantly

after 12 months. The different fractions of litter decompose at different rates: the

slowest breakdown characterizes lignified parts (i.e., cones and twigs) resulting in

statistically significant differences (fig. 2).

The accumulation of heavy metals in the upper layer of the soil showed that at

the majority of sites, only lead occurs in amounts several tens or even several hundreds of times greater than in the parent rock (fig. 1). High concentrations of heavy

metals in litter are most frequent in Silesia, and in the western sites along the climatic transect ( i.e., in the western-border area of the country). The highest correlations between heavy metal contents in litter were found for needle litter decomposition and Fe (corr. coeff. 0.83), Zn (corr. coeff. 0.80), Pb (corr. coeff. 0.82), Ni (corr.

coeff. 0.89), as well as for mixed litter decomposition and Fe (corr. coeff. 0.71), Zn

(0.66), Pb (0.68), and Ni (corr. coeff. 0.72). There were not significant correlations

Figure 4 — Correlation index

between nutrients and heavy

metals in organic-mineral soil

horizon-A and litter decomposition rate (litter bags) in the

Silesian and climatic transect

for 1994-1995 study.

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

Two transect, 10.94-09.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

NUTRIENTS HEAVY METALS

needles wood cones mixed

CORRELATION INDEX

0.8

0.6

0.4

0.2

0

-0.2

-0.4

-0.6

Two transect, 10.94-03.95

Fe Mn Zn Cu Pb Nl Cr

–––––––

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 185

between the rate of decomposition of wood or cones and the content of metals in

litter, except for Cr (table 1). The highest correlation between heavy metals content

in organic-mineral horizon of soils and needle decomposition was determined for

Cu (corr. coeff. 0.901), Zn (corr. coeff. 0.901), Ni (corr. coeff. 0.834), Fe (corr. coeff.

0.850) (fig. 4).

In some cases the correlations between decomposition rate and metal contents

were positive. Comparing decomposition rates with the degree of litter pollution

showed that in the case of needle decomposition for six significant correlations,

two were positive; in the case of mixed litter decomposition for five significant

correlations, two were positive. In both cases decomposition reacted positively to

the presence of Fe and Pb. Decomposition rates compared with degree of soil pollution showed that in the case of needle decomposition for five significant correlations, all were positive; in the case of mixed litter decomposition, only one significant correlation was positive.

Thus, these results suggest that the pine forest ecosystems can react positively to the addition of pollution elements to the substratum and litter by

enhancing the cycling of matter in the ecosystem as measured by the rate of

litter decomposition.

References

Berg, B.; Berg, M.P.; Bottner, P.; Box, E.; Breymeyer, A.; Calvo de Anta, R.; Couteaux, M.; Escudero, A.; Gallardo, A.; Kratz, W.;

Madeira, M.; Malkonen, E.; McClaugherty, C.; Meentemeyer, V.; Munoz, F.; Piussi, P.; Remacle, J.; Virzo de Santo, A.

1993. Litter mass loss rates in pine forests of Europe and Eastern United States: some relationships with climate and litter

quality. quality. Biogeochemistry 20. Netherlands: Kluwer Academic Publishers; 127-159. quality.

Breymeyer, A. 1996. Badania borow na transekcie klimatycznym wzdluz rownoleznika 52o

N (12-32o E) i na transekcie

zanieczyszczenia; zalozenia ogolne, uwarunkowania geograficzne i ekologiczne. (Investigation of pine forest along cli- zanieczyszczenia; zalozenia ogolne, uwarunkowania geograficzne i ekologiczne. (Investigation of pine forest along climatic transect matic transect 52o

N (12-32oE) and pollution transect: general assumptions, geographical and ecological conditions). E) and pollution transect: general assumptions, geographical and ecological conditions).

Warszawa: Dokumentacja Geograficzna.

Degorski, M. 1996a. Charakterystyka morfolitologiczna siedlisk borow i borow mieszanych na tansektach badawczych. Charakterystyka morfolitologiczna siedlisk borow i borow mieszanych na tansektach badawczych

(Morpholithological characteristic of pine forest and mixed pine forest along studies transects). (Morpholithological characteristic of pine forest and mixed pine forest along studies transects). Warszawa: Dokumentacja (Morpholithological characteristic of pine forest and mixed pine forest along studies transects).

Geograficzna.

Degorski, M. 1996b. Zroznicowanie fizyko-chemicznych wlasciwosci gleb siedlisk borow i borow mieszanych na transektach

badawczych: klimatycznym (12-32oE) i zanieczyszczeniowym. E) i zanieczyszczeniowym. E) i zanieczyszczeniowym.(Differentiation of physico-chemical soil properties of pine

forest and mixed pine forest along studies transects). forest and mixed pine forest along studies transects). Warszawa: Dokumentacja Geograficzna. forest and mixed pine forest along studies transects).

Laskowski, R.; Niklinska, M.; Marynski, M. 1994. Heavy metals in forest litter: a chemical time-bomb. Heavy metals in forest litter: a chemical time-bomb. In: Solon J.; Roo- Heavy metals in forest litter: a chemical time-bomb. 

Zielinska E.; Bytnerowicz A., eds. Climate and atmospheric deposition studies in forest. Nieborow: International Conference, 1992. Conference Papers 19. Institute of Geography and Spatial Organization PAS.; 211-217.

Przybylska, B. 1995. Wieloletnie zmiany zanieczyszczenia atmosfery w Polsce (w:) Ocena i stan srodowiska przyrodniczego

Polski i innych krajow. Miedzynarodowe Sympozjum Naukowe. Polski i innych krajow. Miedzynarodowe Sympozjum Naukowe. (Long-term changes of air pollution in Poland). Jachranka Polski i innych krajow. Miedzynarodowe Sympozjum Naukowe.

10-13 pazdziernika; 112- 114.

Rocznik Statystyczny wojewodztwa katowickiego.1992. (Annual statistical yearbook of the Katowice Voivodeship). Katowice, Annual statistical yearbook of the Katowice Voivodeship)

Poland.

Smialkowski, J. 1996. Zroznicowanie warunkow klimatycznych na transekcie Christdorf- Chocimsk biegnacym pomiedzy 52

a 53∞ szerekosci geograficznej wschodniej) i zanieczyszczenia powietrza. szerekosci geograficznej wschodniej) i zanieczyszczenia powietrza (Differentiation of thermic and precipitationm

conditions as an effect of altitude and distance from the Atlantic Ocean on the climatic and Silesian transect). conditions as an effect of altitude and distance from the Atlantic Ocean on the climatic and Silesian transect). Dokumentacja conditions as an effect of altitude and distance from the Atlantic Ocean on the climatic and Silesian transect).

Geograficzna.

Wawrzoniak, J.; Malachowska, J. 1996. Spatial distribution of foliar chemistry of Scots pine (Pinus silvestris L.) related to air

pollution and forest health in Poland. pollution and forest health in Poland. Proceedings of XVI international meeting for specialists in air pollution effects on pollution and forest health in Poland.

forest ecosystems. 1994 Sept. 7-9; Fredericton, New Brunswick, Canada: Con. Forest Service, Atlantic Centre; 267-273.

Wawrzoniak, J.; Malachowska, J. 1997. Stan uszkodzenia lasow w Polsce w 1995 r na podstawie badan monitoringowych

(Degree of Polish forest damage evaluated on the basis of 1995 monitoring). Warszawa: Biblioteka Monitoringu (Degree of Polish forest damage evaluated on the basis of 1995 monitoring)

Srodowiska, PIOS.

Session I1I Transect Studies on Pine Litter Organic Matter: Decomposition and Chemical Properties Breymeyer, Degorski, Reed

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 187

200

180

160

140

120

100

80

60

40

20

0

-

-

-

-

-

-

-

-

-

-

- -

--------------------------

1970 1975 1980 1985 1990 1995

-

-

Years

Annual emission, ton x 1000

Abstract

The Niepolomice Forest is located near the city of Krakow in southern Poland. Since the erection of large iron works in

the 1950’s, the forest has suffered from heavy pollution with SO2

 and industrial dusts containing heavy metals. During

the past 10 years, the ecology of the Niepolomice Forest has been intensively studied and the impact of industrial

pollution has been significantly reduced. With the advent of modern computer intensive techniques, data gathered in

the past have been reanalyzed with respect to the spatial and temporal variations of the forest ecosystem response to

industrial pollution. In addition, the effects of natural conditions (soil, vegetation) and industrial pollution (heavy

metals, sulfur dioxide) upon the pine stands (tree volume increment, crown injuries) in the Niepolomice Forest were 

studied by using a geographic information system. Procedures of statistical analysis involving bootstrapping were 

developed. Results suggest the apparent fertilization of forest stands as a result of industrial pollution—an effect not

revealed in former studies.

Introduction

The Niepolomice Forest, an ancient forest complex situated close to the urban and

industrial area of Kraków in southern Poland, may be regarded as an object of a

large-scale ecological experiment. In the 1950’s, the forest was suddenly exposed to

enormous amounts of industrial pollution that increased steadily until the end of

the 1980’s and then decreased (fig. 1). This pattern was caused by the extensive

development of the metallurgic industry and subsequent production limitation

and technological improvements—a pattern clearly reflecting the political

changes in the country. These changes provide a unique opportunity to study

the response of forest ecosystems to such manipulations.

Geographic Information System (GIS) Analysis of Ecosystem

Response to Industrial Pollution in the Niepolomice Forest in

Southern Poland1

January Weiner,2 Stefan Fredro-Boniecki,2 David Reed,2 Ann Maclean,3 Marshall Strong,3

and Michael Hyslop3

1 An abbreviated version of this paper

was presented at the International Symposium on Air Pollution and Climate

Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Ecology and Graduate Student, respectively, Department of Ecosystem Studies, Institute of Environmental Biology, Jagiellonian University,

ul. Ingardena 6, 30-060 Kraków,Poland.

e-mail: weine@eko.eko.uj.edu.pl

3 Professor of Forest Biometrics, Associate Professor of Remote Sensing/

GIS, Graduate Researcher, and GIS

Technician, respectively, School of

Forestry and Wood Products, Michigan Technological University,

Houghton, MI., USA 49931.

Figure 1 — Changes in the total emissions of industrial dusts (filled bars) and sulfur dioxide (open

bars) in the Krakow agglomeration (Turzanski and Wetrz 1995). Note the distinct decrease of

pollution during the last 10 years and an increased proportion of dust and SO2

 emissions in the last

5 years which may lead to increased acidification.
188 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The Niepolomice Forest is a relatively large forest complex (about 110 km2)

which includes a variety of soils and vegetation types. For many years the forest

ecosystems of Niepolomice were the subjects of research for many scientific institutions in Kraków. This activity resulted in hundreds of publications (Banasik 1978),

including two synthesizing volumes (Grodzinski and others 1984, Kleczkowski

1981). However, the vast body of data could not be fully exploited. With the advent

of efficient computer-intensive methods, such as geographic information systems

(GIS), a more advanced analysis of the spatial and temporal variation can be undertaken and an explanation of the emerging patterns can be attempted. However,

the data have not been collected expressly for the purpose of such analysis and

therefore a number of methodological problems must be solved.

This paper reports a study that reanalyzes the available data on Niepolomice

Forest ecosystems by using GIS to reveal possible spatial correlations between environmental conditions, pollution levels, and indices of tree stand quality.

Methods and Materials

As a basis for the GIS database, published topographic and thematic maps of

Niepolomice Forest area were used: 1:25,000 topographic map; 1:20,000 forestry

compartment map; 1:37,500 potential vegetation map (Gruszczyk 1981a); 1:37,500

present vegetation community (Cwikowa and Lesinski 1981); 1:37,500 forest soils

map (Gruszczyk 1981b); 1:37,500 site type map (Gruszczyk 1981a); and 1:37,500

overstory stand type map (Maczynski 1981). The GIS (ARC Info) methods were

developed at Michigan Technological University. After scanning and editing, the

maps were registered to Universal Transverse Mercator coordinates using numerous

global positioning system (GPS) readings made in the field (Weiner and others

1996). The database was converted to IDRISI 4.0 format (Eastman 1992)4

 and

subsequent data input and analysis was performed at the Jagiellonian University

in Krakow.

The basic information on the Niepolomice Forest is contained in the GIS layers

such as soils (fig. 2) or forest stands (fig. 3), which are the electronic representations

of previously published maps. New maps were produced based on published quantitative data. The sources available usually contain precise numerical information

about the variable of interest, but the spatial coordinates for each sampling location are quite general. In most cases only the number of a forest compartment or

subcompartment is provided and no better accuracy can be derived after the passage of 20 or more years since the field research was done. The forest

subcompartments in Niepolomice can be of various sizes and shapes, sometimes

not larger than 100 m2

, but a standard forest compartment is 400 by 800 m in size.

Moreover, sampling for different variables was performed in different locations,

although the sampling intensities were quite dense. Thus, a direct use of the data

for a multivariate analysis is impossible because one cannot order the field data in

sets of exactly corresponding spatial variates. An approximate analysis can be made

by comparison of maps produced by the spatial interpolation of point data. This

can be done only by presuming that the variables studied are continuous (forming

a spatial gradient), at least locally. It was assumed that this was the condition in the

case of the variables used.

Most of the data came from the larger, southern part of the Niepolomice Forest

(mostly pine stands; fig. 3). Data for industrial pollution and ecosystem response

were obtained from the index of tree crown injuries (Grabowski 1981). This index

(DAM) combines various estimates of damage to pine needles and malformations of

twigs and branches, most probably caused by SO2

 pollution (Grabowski 1981). Spatial

interpolation of 86 data points resulted in a map (fig. 4).

Data were also obtained from the index of heavy metal accumulation. The content of six heavy metals (Cd, Pb, Zn, Cu, Fe, Mn) was measured in mosses at 15

locations in 1975 and in 1992 (Godzik and Szarek 1993). From these point data

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop

4 Unpublished data on file at Department

of Forest Mensuration, Agricultural

Academy, Krakow, Poland.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 189

interpolated maps have been produced. The contents of all metals are strongly

intercorrelated; the first principal component (variable PC1) may constitute an index of contamination with industrial dusts (fig. 5).

Indices of tree volume increment were also used (Rieger and others 1987, 1989).5

A detailed dendrometric survey of pine stands was carried out at 114 locations of

the southern forest complex in 1981. The plots were located in younger (A: 20-45

years) and older (B: 60-130 years) stands. The data set, combined from published

and unpublished information, includes: tree age, height, diameter at breast height

(dbh.), and a numerical index of diameter increment dynamics during the last 5

years preceding the study, as related to the previous 5-year period (variables WZDA

and WZDB for younger and older age classes, respectively; fig. 6). But, the original

data on tree ring measurements were not accessible.

Another index was calculated from the data as a residual of the regression line

of tree volume on age (fig. 7). The tree volume was calculated from height and dbh.

according to allometric formulas and empirical parameters given for Scots pine

5 Unpublished data on file at Department

of Forest Mensuration, Agricultural

Academy, Krakow, Poland.

Figure 2 — The soils of the

Niepolomice Forest (an IDRISI

map): 1--proper brown (mesic)

soils (5.1 percent); 2--podsolic

and cryptopodzolic soils (13.0

percent); 3--brown podsolized

soils (8.1 percent); 4--moist soils,

black earths (23.7 percent); and

5--seasonal gleyish soils (50.1

percent) (Gruszczyk 1981).

Figure 3 — Forest stands

classified by dominant species

(IDRISI map; Maczynski 1981).

1

2

3

4

5

3744.1m

Pine

Oak

Black alder

Spruce

Birch

Hornbean

Ash

Poplar

3744.1m

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
190 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

(Sulinski 1993). The residuals make up an index of cumulative volume increment

during the whole life of a tree as related to the average tendency in the whole forest

(variables TVRA and TVRB).

The first index should represent the recent changes in tree condition (e.g., related to changing pollution), while the second one should depend more on the

natural stand quality. The map of both indices was obtained by a spatial interpolation of point data for younger (A) and older (B) age categories separately (figs. 8, 9).

Analysis and Discussion

Characteristic patterns of the spatial distribution of environmental and forest

response variables are apparent according to the GIS maps. The areas most exposed

Figure 4 — Index of tree crown

injuries in arbitrary units

(Grabowski 1981).

Figure 5 — First principal

component (PC1) of the

contents of six heavy metals in

mosses in 1975 (Godzik and

Szarek 1993).

0

4

9

13

17

22

26

30

35

39

43

48

52

56

61

65

3744.1m

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

3744.1m

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 191

to heavy metal pollution demonstrated an accelerated tree volume increment (figs.

5, 6). We did not find visible association between the index of heavy metal

contamination and the injuries of tree crowns (figs. 5, 6).

The variables represented on maps by values obtained from spatial interpolation cannot be directly subject to a statistical analysis because they have extremely

high autocorrelations (Moran’s I in the range of 0.95 to 0.99 for different variables

studied; Eastman 1992). The occurrence of autocorrelation is inherent in empirical

distributions of variables of this kind (Legendre 1993) and an additional

autocorrelation component is introduced by the procedure of spatial interpolation. This may cause an overestimation of significance of test statistics (Legendre

1993). To minimize this effect a randomization (“bootstrap”) procedure was used

(Noreen 1989). The variables of interest were sampled from a limited set of randomly selected point locations. Simulations revealed that with a decrease of the

number of sampling points (from the original total number of pixels of about

870,000), the coefficient of autocorrelation (Moran’s I ) steadily decreases to a value

Figure 6 — Index of tree ring

diameter dynamics, WZD

(Rieger and others 1987, 1989).

Figure 7 — Regression of pine

tree volume on age in the

southern part of the

Niepolomice Forest (double

logarithmic plot).6

 Residuals for

a given age represent a

cumulative tree volume

increment index (TVR).

6

 Rieger and others 1987, Sulinski 1993). 

1977 - 1981

1972 - 1976

WZD = 1977 - 1981

1972 - 1976

x 100

2.0

1.0

0.5

0.3

0.2

0.1

–

–

–

–

–

–

–

20 30 40 50 60 70 8 90 100 150

––––

–

–

–

––

–

Tree age (years)

Tree volume (m3)

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
192 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

of about 0.7 at 1,000 points and then its variation increases (fig. 10). Accordingly,

the sample size was set at about 1,500 points. The statistical analysis of association

between variables or analysis of variance by categories was performed on these

samples. The procedure was repeated (with different sets of random point locations) and average statistics were calculated.

The associations between single variables were examined by using Spearman

rank correlation coefficient rs. Correlation coefficients were averaged (± standard

deviation, range) for 12 randomly sampled sets of data (table 1). A statistically significant positive correlation (rs␣>␣ 0, p␣ <␣ 0.05) was r evealed between the tree growth

Figure 8 — Cumulative index

of tree volume increment (TVR)

in the southern part of the

Niepolomice Forest (arbitrary

units): (A) - pine stands 20-45

years old; and (B) - pine stands

60-130 years old.

1 TVRA = tree volume index for young stands (20-45 yrs.). TVRB = tree volume index for older stands (60-130 yrs.).

WZDA = diameter growth index for young stands (20-45 yrs.). WXDB = diameter growth index for older stands

(60-130 yrs.). PC1 = first principal component of the heavy metal concentrations. DAM = crown injury index.

2 Average correlations that are different from zero (p < 0.05).

Table 1 — Average ( Table 1 ± standard deviation, range) Spearman rank correlation coefficients for dendrometric variables

(WZDA, WZDB, TVRA, TVRB), dust pollution index (PC1), and pine crown injury index (DAM).1

Item TVRA TVRB WZDA WZDB DAM

PC1

Average -0.098 2 0.320 2 0.492 2 0.330 2 0.071

Standard deviation 0.013 0.020 0.034 0.034 0.021

Minimum 0.115 0.294 0.444 0.297 0.035

Maximum -0.073 0.348 0.544 0.427 0.113

DAM

Average 2

- 0.175 2 - 0.247 2 0.243 2 0.331

Standard deviation 0.257 0.023 0.069 0.067

Minimum -0.987 -0.278 0.088 0.165

Maximum -0.065 -0.208 0.311 0.401

0

10

20

30

39

49

59

69

79

89

99

109

118

128

138

148

3744.1m

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 193

volume increment indices (WZDA, WZDB) and the first principal component (PC1)

of the heavy metal concentrations (i.e., the tree volume increments) (independent

of age class) were increased in the most polluted areas. The total volume of older

trees (TVRB) was also positively correlated with dust fall (table 1).

The tree crown injuries (DAM) were positively correlated with the indices of

tree growth dynamics in both age classes (WZDA, WZDB), but negatively correlated with the total tree volume (TVRA, TVRB; table 1). Thus, a malformation of

tree crowns was associated with an increased tree volume increment in the years

immediately preceding the field studies, but the trees with most damaged crowns

were also the smallest in the sample.

Five general types of soils have been distinguished in the southern part of the

forest covering different proportions of the area (fig. 2). Soil type exerts a significant effect upon the characteristics of tree stands. The index of increase in both age

classes (WZDA, WZDB) and the index of tree volume in older class (TVRB) is significantly higher in brown (mesic) soils. The total volume (TVRA, TVRB) tends to

be the smallest on dry (podzolic) soils (p < 0.01; Kruskal-Wallis). Tree crown damage is highest on brown mesic and in dry podsolic soils.

In the southern part of the forest the brown (mesic) soils are concentrated at the

western edge in the most polluted area. To discriminate the effects of soil type and

dust pollution upon the dendrometric variables, the same statistical analysis was

performed within the soil classes. Two distinct patterns persist after the effect of

soil type is removed: the positive correlation of the index of dust pollution (PC1)

with the indices of growth acceleration (WZDA, WZDB) and a negative correlation

of tree crown injuries (DAM) with tree volume, particularly in older age classes

(TVRB). In other words, the smaller the tree volume, the greater the damage. This

last phenomenon may indicate a synergistic effect of poor habitat and gaseous pollution. In the older age class the trees were exposed to pollution during the recent

one-third to one-fifth of their lifetime only. Thus, the relatively small volume must

Figure 9 — Index of tree ring

diameter dynamics (WZD,

percent) in the southern part of

Niepolomice Forest: WZD =

100 percent means an equal rate

of tree ring diameter increment

during 5-year periods 1977-1981

and 1972-1976; WZD > 100

percent denote acceleration of

growth, and WZD < 100 percent

is growth retardation. (A) - pine

stands 20-45 years old; (B) - pine

stands 60-130 years old.

3744.1 m -463

-406

-394

-292

-234

-177

-120

-63

-6

51

108

165

223

280

337

394

3744.1m

A

B

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
194 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

have been the effect of habitat conditions while the relatively strong malformation

of crowns in the weakest individuals is a secondary phenomenon. No correlation

was found between the tree crown injuries (DAM) and heavy metal contamination

(PC1; table 1).

A simultaneous effect of two variables (PC1 and DAM) upon the dendrometric

characteristics of pine trees was analyzed by using coefficient of determination (R2)

for a multiple regression:

Y = a0 + a1 PC1 + a2 DAM

in which PC1 is the dust contamination index, DAM is the pine crown injury index,

and a0, a1, a2 are parameters. The R2 values were calculated and showed an

indication of a positive (+) or a negative (-) effect of the independent variables

(the numerical values of parameters are omitted) (table 2). The two independent

Figure 10 — Changes in the

coefficient of autocorrelation

(Moran’s I) with decreasing

number of points sampled from

spatially interpolated data sets.

Grey bar indicates the number

of point locations used in

analysis.

1 TVRA = tree volume index for young stands (20-45 yrs.). TVRB = tree volume index for older stands (60-130 yrs.). WZDA =

diameter growth index for young stands (20-45 yrs.). WZDB = diameter growth index for older stands (60-130 yrs.). PC1 =

first principal component of the heavy metal concentrations. DAM = tree crown damage index.

2 Average R2 values differing from (p < 0.05).

Table 2 — Average ( Table 2 ± standard deviation, range) coefficients of determination (R2

) for a multiple regression for

dendrometric variables (WZDA, WZDB, TVRA, TVRB), dust pollution index (PC1), and pine crown injury index

(DAM). 1

Item TVRA TVRB WZDA WZDB

R2

Average 0.018 2 0.176 2 0.290 2 0.208

Standard deviation 0.0065 0.0203 0.0241 0.0241

Minimum 0.009 0.137 0.257 0.178

Maximum 0.028 0.211 0.346 0.250

Sign of a1 (effect of PC1) - + + +

Sign of a2 (effect of DAM) - - + +

1.0

0.9

0.8

0.7

0.6

0.5

0.4

-

-

-

-

-

-

-

-

+

+

+

+ +

+

+

+

+

+ +

*

*

*

*

*

*

*

*

*

*

*

100 1,000 100,000 100,000 1,000,000

-

---

-

-

--

-

-

---

-

-

-

-

-

-

-

-

-

-

-

-

---

-

-

-

-

-

--

-

-

WZDA TVRA PC1 DAM Average + *

Number of sampled point locations

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 195

variables account for 18 to 29 percent of variation in dendrometric indices (except

for TVRA, (i.e., total volume of trees in younger stands), for which the effect was

not significant). These results agree with the Spearman rank correlation patterns

for single variables.

The most probable explanation of this apparent paradox of growth acceleration of trees exposed to highest pollution lays in the high nutritional value

of industrial dusts for plants. These pollutants contain a great load of Ca, Mg,

N, P, and other nutrients (Grodzinski and others 1984) as well as heavy metals.

Moreover, the dust is basic and therefore it can compensate for low pH caused

by SO2 pollution.

These patterns describe the situation in the Niepolomice Forest in the late 1970’s.

Since, then, the level of pollution has changed dramatically (fig. 1). A new field of

study, GIS, repeated 20 years later, could help to confirm or to reject the hypothesized explanation of these effects.

Acknowledgments

This work was performed in connection with the “U.S.-Poland Cooperative Project

on Status and Long Term Trends in Forest Ecosystems: Climate, Pollution, and Forest

Health” that was funded by the U.S. Department of State, U.S. Environmental

Protection Agency, USDA Forest Service, USDA Foreign Agricultural Service (ICD),

Polish Academy of Sciences, and the Polish Ministry of Environmental Protection.

Jagiellonian University, Kraków, Poland, Michigan Technological University,

Houghton, Michigan, USA, and Bowling Green State University, Bowling Green,

Ohio, USA, have also provided support. We thank Stanislaw Orzel and Jerzy Raimer

for providing unpublished data on tree growth.

References

Banasik, Julita. 1978. Przyrodnicza bibliografia Puszczy Niepolomickiej ze szczegolnym uwzglednieniem lat 1947-1977. (Sci- Przyrodnicza bibliografia Puszczy Niepolomickiej ze szczegolnym uwzglednieniem lat 1947-1977. (Scientific bibliography of Niepolomice Forest with special focus on the period 1947-1977. In Polish). Studia Naturae A, 14: entific bibliography of Niepolomice Forest with special focus on the period 1947-1977. In Polish

205-223.

Cwikowa, Alicja; Lesinski, Jerzy A. 1981. ne zroznicowFlorystyczanie zbiorowisk aktualnej roslinnosci ne zroznicowFlorystyczanie zbiorowisk aktualnej roslinnosci lesnej Puszczy

Niepolomickiej (Floristic differentiation of communities of present vegetation of Niepolomice Forest. In Polish, English

summary). summary). Studia Osrodka Dokumentacji Fizjograficznej, IX; 159-196. summary).

Eastman, J. Ronald. 1992. IDRISI, Version 4.0. User’s guide and technical reference. Worcester, MA. IDRISI, Version 4.0. User’s guide and technical reference

Godzik, Barbara; Szarek, Grazyna. 1993. Heavy metal in mosses from the Niepolomice Forest, southern Poland—changes

1975-1992. 1975-1992. Fragmenta Floristica et Geobotanica 38(1): 199-208. 1975-1992. 

Grabowski, Andrzej. 1981. Zmiany morfologiczne koron sosny w Puszczy Niepolomickiej. (Malformations of crowns of Scots

pine in Niepolomice Forest. In Polish, English summary). pine in Niepolomice Forest. In Polish, English summary). Studia Osrodka Dokumentacji Fizjograficznej, IX; 357-368. pine in Niepolomice Forest. In Polish, English summary).

Grodzinski, Wladyslaw; Weiner, January; Maycock, Paul F. 1984. Forest ecosystems in industrial regions. Forest ecosystems in industrial regions. Springer Vlg. Heidel- Forest ecosystems in industrial regions.

berg; 277 p.

Gruszczyk, Andrzej. 1981a. Siedliskowe typy lasow w Puszczy Niepolomickiej. (Types of forest site of Siedliskowe typy lasow w Puszczy Niepolomickiej. (Types of forest site of Niepolomice Forest. In

Polish, English summary). Studia Osrodka Dokumentacji Fizjograficznej, IX; 205-220. Polish, English summary)

Gruszczyk, Andrzej. 1981b. Gleby Puszczy Niepolomickiej. (Soils of Niepolomice Forest. In Polish with English summary). Gleby Puszczy Niepolomickiej. (Soils of Niepolomice Forest. In Polish with English summary)

Studia Osrodka Dokumentacji Fizjograficznej, IX; 71-88.

Kleczkowski, Antoni S., Ed. Wartosci srodowiska przyrodniczego Puszczy Niepolomickiej i zagadnienia jej ochrony. (Re- Wartosci srodowiska przyrodniczego Puszczy Niepolomickiej i zagadnienia jej ochrony. (Resources of the natural environment of the Niepolomice Forest and problems with its protection). sources of the natural environment of the Niepolomice Forest and problems with its protection). Studia Osrodka sources of the natural environment of the Niepolomice Forest and problems with its protection). 

Dokumentacji Fizjograficznej, IX; 409 p.

Legendre, Pierre. 1993. Spatial autocorrelation: trouble or new paradigm? Ecology 74: 1,659-1,673. Spatial autocorrelation: trouble or new paradigm?

Maczynski, Marian 1981. Drzewostany Puszczy Niepolomickiej. (Stands of Niepolomice Forest. In Polish, English summary). Drzewostany Puszczy Niepolomickiej. (Stands of Niepolomice Forest. In Polish, English summary).

Studia Osrodka Dokumentacji Fizjograficznej, IX; 197-204.

Noreen, Eric W. 1989. Computer intensive methods for testing hypotheses. Computer intensive methods for testing hypotheses. New York: John Wiley and Sons; 229 p. Computer intensive methods for testing hypotheses.

Raimer, Jerzy; Grabczynski, Stanislaw; Orzel, Stanislaw; Rieger, Roman. 1987. Evaluation of the regularity in growth of trees

in selected scots pine (Pinus silvestris L.) stands in the L.) stands in theNiepolomice forest on the basis of selected growth functions. Niepolomice forest on the basis of selected growth functions. Acta Niepolomice forest on the basis of selected growth functions.

Agraria et Silvestria 26: 129-146.

Rieger, Roman. 1987. The loss of increment in dbh. as a basis for the estimation of a degree of degradation of scots pine (Pinus

silvestris L.) stands in the Niepolomice Forest. L.) stands in the Niepolomice Forest. Acta Agraria et Silvestria 26: 103-112. L.) stands in the Niepolomice Forest. 

Rieger, Roman; Grabczynski, Stanislaw; Orzel, Stanislaw; Raimer, Jerzy. 1987. The dynamics of diameter increment of scots

pine (Pinus silvestris L.) in stands of the Niepolomice Forest in the light of dendroecological research. Acta agraria et L.) in stands of the Niepolomice Forest in the light of dendroecological research

Silvestria 26: 114-127.

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Rieger, Roman; Orzel, Stanislaw; Grabczynski, Stanislaw; Raimer, Jerzy. 1987. Characteristics of diameter increment struc- Characteristics of diameter increment structure of scots pine (Pinus silvestris L.) stands in the Niepolomice Forest. L.) stands in the Niepolomice Forest. Acta agraria et Silvestria 25: 87-102. L.) stands in the Niepolomice Forest. 

Sulinski, Jozef. 1993. Modelowanie bilansu wodnego w wymianie miedzy atmosfera, drzewostanem i gruntem przy uzyciu

kryteriow ekologicznych. (Modeling of water balance in an exchange between atmosphere, stand and ground using eco- kryteriow ekologicznych. (Modeling of water balance in an exchange between atmosphere, stand and ground using ecological criteria. In Polish, English summary). logical criteria. In Polish, English summary). Zesz. Nauk. AR Krakow; 1-133. logical criteria. In Polish, English summary).

Turzanski, Konrad P.; Wertz, Jerzy, eds. 1995. Raport o stanie srodowiska w Wojewodztwie Raport o stanie srodowiska w Wojewodztwie Krakowskim w 1994 r. Raport o stanie srodowiska w Wojewodztwie (Environ- (Environmental protectioon in Cracov Region, report for 1994. In Polish). mental protectioon in Cracov Region, report for 1994. In Polish). Krakow: Biblioteka Monitoringu Srodowiska; 198 p. mental protectioon in Cracov Region, report for 1994. In Polish).

Weiner, January; Maclean, Ann; Reed, David; Strong, Marshall. 1995. The use of a geographic information system to investi- The use of a geographic information system to investigate atmospheric influences on forest growth in the Niepolomice Forest near Krakow, Poland. gate atmospheric influences on forest growth in the Niepolomice Forest near Krakow, Poland. In: Cox, R.; Percy, K.; gate atmospheric influences on forest growth in the Niepolomice Forest near Krakow, Poland.

Jensen, K.; Simpson, C., eds. Air pollution and multiple stresses: the 16th international meeting for specialists in air

pollution effects on forest ecosystems. Fredericton, New Brunswick, Canada: Natural Resources Canada, Canadian Forest

Service - Atlantic Centre; 275-282.

Session I1I Geographic Information System Weiner, Fredro-Boniecki, Reed, Maclean, Strong, Hyslop
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 197

Wojciech Dmuchowski,2 Ewa U. Kurczynska,2 Wieslaw Wloch3

Abstract

The impact of environmental pollution is defined for the chemical composition of Scots pine (Pinus sylvestris L.)

needles and cambial activity in the tree stems in Polish forests. The research investigated 20-year-old trees growing in

two areas in significantly different levels of pollution. The highly polluted area was located near the Warsaw steelworks,

while the area with relatively low level of pollution (control area) was in the Kampinos National Park, 10.5 km away

from the steelworks. The research revealed significant differences in chemical composition of needles between the two

areas. The increased content of almost all of the studied elements was observed in all of the typical pollutants such as

sulfur, chlorine, and heavy metals; and in the nutrients such as nitrogen, potassium, magnesium, calcium, and sodium.

Differences were also observed in relations between particular elements and accumulation of nutrient elements (ANE).

The changes in chemical composition of needles affected by environmental pollution were accompanied by a reduction

of cambial activity. This reduction was not expressed as a decrease in the number of cells in cambial zone, but as a

decreased number of cells on the phloem side, both at the tree-top level and at breast height. These differences were

particularly visible in the second half of the vegetative season.

Introduction

Air pollution is considered one of the main causes of forest decline in central Europe

(ECE 1984, Molski and Dmuchowski 1986, Nihlgard 1985, Schutt and Cowling 1985).

According to the results of national forest health monitoring, more than 50 percent

of trees in Poland represent the highest defoliation classes (Malachowska and

Wawrzoniak 1995). Among the European countries, only the forest condition in the

Czech Republic is worse than in Poland (United Nations Economic Commission

for Europe 1997).

The negative influence of air pollutants on tree growth has been well established (Smith 1990). Air pollution affects various physiological and biochemical

processes (Biggs and Davis 1981, Burton and Morgan 1983, Bytnerowicz and Grulke

1993), as well as morphology and anatomy of shoot and root system of trees (Heale

and Ormrod 1982, Kozlowski and others 1991, Kurczynska and others 1996, Smith

and Davies 1978). Air pollutants reduce shoot growth of seedlings, and height

growth of older trees may be reduced as well (Kozlowski and others 1991). The leaf

area of trees exposed to air pollutants may be reduced because of inhibition of leaf

formation, arrested leaf expansion, and accelerated leaf abscission (Kozlowski

and others 1991). Air pollutants also reduce diameter growth of seedlings and

mature trees (Thompson 1981). However, influence of pollution on cambial

activity and differentiation of its derivatives, processes which regulate tree

growth and development (Zajaczkowski and Wodzicki 1978), have not yet been

adequately investigated.

Recently published results showed that in the polluted environment physiological polarity of pine stems was affected. This resulted in changed polar transport of auxin at the cellular level of the cambial zone (Wodzicki and others 1990a).

Polar transport of auxin, in turn, is responsible for regulation of cambial activity

and differentiation of its derivatives (Larson 1994).

This study investigated the effects of industrial/urban air pollution on chemical composition of needles as well as cambial activity and differentiation of its derivatives, especially on the phloem side, of Scots pine (Pinus sylvestris L.) trees in

the polluted and control forest sites near Warsaw, Poland.

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Plant Chemist, Plant Physiologist, respectively, Botanical Garden of Polish

Academy of Sciences, ul. Prawdziwka

2, 02-973 Warsaw, Poland.

3 Anatomist, Department of Biophysics and Cell Biology, Silesian University, ul. Jagiellonska 28, 40-032

Katowice, Poland.

Chemical Composition of Needles and Cambial Activity

of Stems of Scots Pine Trees Affected by Air Pollutants in

Polish Forests1
198 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Materials and Methods

Investigations were carried out at two sampling areas along an air pollution gradient

(Molski and Dmuchowski 1990) during one growing season in 1992. These sampling

areas were the Kampinos National Park (a control area located upwind, 10 km

away from the Warsaw Steelworks) and a site in the vicinity of the Warsaw

Steelworks (a polluted area located upwind, 0.2 km away from the Warsaw

Steelworks). Sampling areas were established in relatively uniform and

representative portions of the stands.

Three trees were selected from each sampling area for anatomical investigation,

and six trees were selected for chemical analysis of needles (18-20-year-old; 8-9 m

height) of Scots pine (Pinus sylvestris L.) with approximately the same diameter at

breast height (30-32).

Foliar mineral composition of Scots pine (N, P, K, Mg, Ca, Mn, S, Cl, and heavy

metals) was determined as an indicator of health condition and contamination of

the experimental trees. The samples of needles were taken from the same six trees

and at the same time as in the anatomical study. Needles from the previous years’

growth from the second and third whorls were collected (Dmuchowski and

Bytnerowicz 1995).

Concentration of chemical elements were determined by using the following

analytical procedures: metals and phosphorus were assessed by spectrophotometry atomic absorption method (Perkin-Elmer 1990, Morton and Roberts 1991) on a

Perkin-Elmer 1100 fitted with a Flows Analysis System (FIAS-2000) and Graphite

Furnace (HGA-700).4

 Sulfur (total) was assessed by infrared absorption method by

using LECO SC-132 Sulfur System (LECO Corporation 1987) and chloride by potentiometric titration using selective chloride ion electrode (LaCroix and others 1970).

Content of elements in needles was presented as the sum of the accumulation

of nutrient elements (ANE). To calculate ANE, all parameters of the contents of

elements in needles were converted into gram-equivalent values. The basis of the

conversion was the atomic mass of elements and the valency of their ions (Ostrowska

and others 1988).

For anatomical analysis, samples included current-year wood, cambium, and

current-year phloem (once in the winter season of dormancy, and April 30, May 8,

June 5, July 3 and 31, August 28, and September 25) from tree-top level and from

breast height of each investigated tree. Samples were fixed, dehydrated, and embedded in Epon (Kurczynska and others 1996). Samples were cut into a series of

cross sections (3 m thick) with an ultramicrotome. The sections were attached to

microscope slides and stained with leucofuchsin (PAS-reaction) and toluidine blue,

mounted in Euparal, and investigated under light microscope. To determine the

number of wood, cambium, and phloem cells, 20 radial rows were counted on the

cross section from each tree. Data were analyzed by Student’s t-test (p<0.05) and

compared between sampling areas.

The soil at the two sites is of the same origin and represents the same type

podsolic (Dmuchowski and others [In press]). The differences between soil conditions are caused by over 30 years of pollutant emission from the Warsaw area. On

both sites the percentage of exchangeable cations in the sorption complex in the

organic level of soils was the same but differences were identified in the mineral

levels of soils. In the soil from the Kampinos area hydrogen was the dominant

cation, which is typical for coniferous forests, while in the soil from the Warsaw

area, calcium was dominant. In the organic layer of the soil from the Warsaw area,

2 times more iron was measured, 30 times more zinc, 6 times more copper, 20 times

more cadmium, and 3 times more chromium than in the Kampinos area.

4 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 199

Results and Discussion

For both areas the chemical analysis of pine needles did not reveal a deficiency of

any of 15 analyzed elements (table 1). Certain macroelements like magnesium, potassium, and phosphorus showed deficiencies in damaged trees (Huttl 1985,

Rehfuess 1983, Tomlison 1987, Zotll and Huttl 1989).

The chemical composition of pine needles at the two locations was much different, particularly in regards to elements recognized as typical pollutants (fig. 1).

For instance, 2-year-old needles from the Warsaw area contained eight times more

lead than the needles from the Kampinos National Park. The levels of lead, zinc,

and chromium defined as toxic for the Scots pine were exceeded. The sulfur content in plants in Central Europe reflects the impact of air pollution with sulfur dioxide. Both investigated areas differed considerably in this point. The high sulfur

levels in the needles from the steelworks are evidence of a detrimental impact of air

pollution with sulfur dioxide on the Scots pine (fig. 2). In the Kampinos National

Park the impact of pollution was not so strong. At present there are no areas in

Poland excluded from the impact of air pollution by sulfur dioxide.

Differences were also observed also in the contents of elements performing

basic physiological functions (fig. 1). The needles from the Warsaw area contained

more magnesium, calcium, potassium, and sodium, but less nitrogen and manganese than the needles from the Kampinos National Park. The level of phosphorus

was the same.

Figure 1 — Chemical composition of needles in the

polluted area shown as percent

of the concentrations in the

control areas.

Table 1 Table 1 — Concentration of macronutrients in Scots pine needles and normal and deficiency levels of these Table 1 

elements in plants.

Element Element Kampinos Warsaw Level Reference

 normal deficiency

N 1.32-1.66 1.83-1.44 1.3-1.6 <1.2-1.3 Wehrmann 1963; Gussone

(percent) 1964, Fiedler and others

1969, cit. Smidt 1988.

P 0.11-0.18 0.12-0.18 0.11-0.17 <0.11 Gussone 1964, Fiedler and

(percent) others 1969, cit. Smidt 1988.

K 0.31-0.50 0.44-0.66 0.35-0.66 <0.30 Zottl 1964; Heinzdorf 1967,

(percent) cit. Smidt 1988.

Ca 0.15-0.33 0.32-0.58 0.23-0.50 <0.05-0.10 Gussone 1964, cit. Smidt

(percent) 1988: Kraus 1965.

Mg 753-910 1207-1518 500-1300 <500 Wehrmann 1963; Fiedler and

(mg/kg) others 1969, cit. Smidt 1988.

Mn 538-916 76-139 175-810 Wehrmann 1963; Fiedler and

(mg/kg) 1 (40-2000) 1 (20) others 1969, Materna 1962;

Bergman1982; Knabe 1985;

Isermann 1986.

1 Picea abies

700

600

500

400

300

200

100

0

elements

S CI N P K Mg Ca Mn Cd Pb Cr As Zn Cu Fe

––––––––––––––

pct

Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
200 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

In order to identify the presence of possible ionic equilibrium disturbances in

the trees affected by environmental pollution, the ionic equilibrium indicators were

calculated: ratio of calcium to magnesium (fig. 3) and ANE total of all sample elements (fig. 4). The calculation revealed certain differences, but these were not statistically significant. It seems that anatomical changes in stems of Scots pine in the

Warsaw area are not caused by mineral deficiencies, but perhaps by toxic effects of environmental pollutants. Concentration of sulfur in needles was at

toxic levels. Concentrations of heavy metals were always higher than normal

levels in plants, but below toxic levels. Elevated concentrations of sulfur and

heavy metals can cause negative effects on trees, especially under synergistic

influences of other pollutants (table 2).

Figure 2 — Total sulfur content

in previous-year needles.

Figure 3 — Ratio Ca/Mg in

previous-year needles.

Figure 4 — Accumulation

nutrients elements (ANE) in

previous-year needles.

0.25

0.20

0.15

0.10

0.05

0.00

S (pct)

6.03

30.0415.0528.0512.0626.0610.0724.07

7.08

28.0811.0925.09

sampling date

Kampinos Warsaw

––––––––––––

–

–

–

–

–

–

2.50

2.00

1.50

1.00

0.50

Ca/Mg

6.03

30.0415.0528.0512.0626.0610.0724.07

7.08

28.0811.0925.09

––––––––––––

–

–

–

–

–

Kampinos Warsaw

(day, month)

2000

1800

1600

1400

1200

1000

ANE

(me/100g)

6.03

30.0415.0528.0512.0626.0610.0724.07

7.08

28.0811.0925.09

Kampinos Warsaw

––––––––––––

–

–

–

–

–

–

(day, month)

Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 201

During the growing season, cambial activity and the number of cells on the

xylem and phloem side were higher at the tree-top level than at breast height, independent on the sampling area. We found no statistically significant differences in

the number of cells in the cambial zone between trees grown in control and polluted areas (fig. 5), and in the number of xylem cells between trees grown in these

two areas (fig. 6). The number of phloem cells was significantly different between

trees grown in control and polluted area (fig. 7). The decrease in the number of

phloem cells was especially visible in the second part of the growing season. It is

possible that changes in the phloem are very important cause of tree decline in the

polluted environment. For instance, in normally growing trees the maximal mass

of functioning phloem appears in the second part of the growing season, when the

most intensive transport of sugars to the stem and roots occurs. Decreasing the

number of cells participating in that transport could lead to the disturbance in translocation of assimilates to different parts of trees and cause limited growth.

Table 2 Table 2 — Concentration of macronutrients in Scots pine needles and normal and toxic levels of these elements in plants. Table 2 

Element Element Scots pine Level Species Reference

Kampinos Warsaw normal toxic

S 0.10-0.14 0.15-0.20 0.06-0.09 >.12 Pinus sylvestris Huttunen and others 1985;

(percent) Dmuchowski and Bytnerowicz 1995.

Cd 0.18-0.36 0.39-0.60 0.01-0.3 plants Allen and others 1974;

(mg/kg) 0.05-0.2 >5 plants Kabata-Pendias and Pendias 1993;

4.8 Picea sp. Burton and Morgan 1983.

Pb 1.9-2.4 11.9-17.6 <10 >30 plants Kabata-Pendias and Pendias 1993;

(mg/kg) 19 Picea sitch. Burton and Morgan 1983.

As 0.05-0.12 0.29-0.54 >1 plants Macnicol and Becket 1985;

(mg/kg) >5 plants Kabata-Pendias and Pendias 1993.

Cu 3.3-5.1 5.8-8.9 2-20 >20 plants Kabata-Pendias and Pendias 1993;

(mg/kg) 2.5-25 plants Hewitt and Smith 1974;

25 leaved trees Linzon and others 1979.

Zn 31-42 68-104 10-100 >100 plants Kabata-Pendias and Pendias 1993;

(mg/kg) Karweta 1976;

226 Picea sitch. Burton and Morgan 1983;

250 leaved trees Linzon and others 1979.

Cr 0.30-0.98 2.47-4.72 0.1-1.0 >1 plants Kabata-Pendias and Pendias 1993

(percent)

Cl 0.02-0.04 0.04-0.08 0.02-0.10 Picea abies Keller 1972

Figure 5 — Number of cells in

the cambial zone during growing

season. Each bar represents the

average value (+ SD) from three

trees.

15

13

11

9

7

5

3

1

number of cells

––––––––

–

–

–

–

–

–

–

–

(day, month)

30.04 8.05 5.06 3.07 31.07 28.08 25.09

Kampinos - breast height Warsaw - breast height

Kampinos - tree top level Warsaw - tree top level

Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
202 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Conclusions

The Scots pine trees growing in the more polluted Warsaw Steelworks environment

differed considerably from the trees growing in the relatively unpolluted Kampinos

National Park environment. In the polluted environment, the needles contained

much higher concentrations of elements recognized as typical pollutants: sulfur

and heavy metals. And the levels of sulfur, lead, zinc, and chromium in the needles

of trees from the Warsaw area can be defined as toxic for the Scots pine.

The differences in the content of elements performing basic physiological functions were of minor importance. No deficiency of any element was identified. The

differences in chemical composition of needles were accompanied by differences

in ionic equilibrium. However, these differences were not statistically significant.

Tree stem tissues such as cambium, phloem, and wood at the tree-top level

were more sensitive to pollutants than tissues in older parts of trees (breast height).

The limited production of phloem (which was statistically significant) and xylem

cells of the trees growing in the polluted environment may have an essential impact on the disturbances in transport of water to tree-crowns and transport of assimilates from crowns to roots. This fact may cause the limitation of growth and

development of the trees growing in the polluted environment and may lead to

their mortality

The growth limitation of the trees from the Warsaw area seems to be caused to

a larger extent by environmental pollution from sulfur, heavy metals, and other

toxic agents (which were not examined, such as nitric oxide) rather than by the

disturbances in ionic equilibrium in trees.

Figure 7 — Number of phloem

cells during growing season. Each

bar represents the average value

(+ SD) from three trees. Bars

marked by * are significantly

different in area category.

Figure 6 — Number of wood

cells during growing season. Each

bar represents the average value

(+ SD) from three trees.

26

21

16

11

6

1

number of cells

––––––––

–

–

–

–

–

–

(day, month)

30.04 8.05 5.06 3.07 31.07 28.08 25.09

Kampinos - breast height Warsaw - breast height

Kampinos - tree top level Warsaw - tree top level

*

*

*

*

180

160

140

120

80

60

40

20

0

number of cells

––––––––

–

–

–

–

–

–

–

–

–

(day, month)

30.04 8.05 5.06 3.07 31.07 28.08 25.09

Kampinos - breast height Warsaw - breast height

Kampinos - tree top level Warsaw - tree top level

Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 203

Acknowledgments

This work was supported by the program “Effects of Atmospheric Deposition and

Climate Change on Forest Ecosystems in Central and Eastern Europe and the United

States,” sponsored by the USDA International Cooperation and Development (ICD).

We thank Laurie Dunn for technical editing of this manuscript.

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Session I1I Chemical Composition of Needles and Cambial Activity of Stems of Scots Pine Trees Dmuchowski, Kurczynska, Wloch
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 205

Barbara Kieliszewska-Rokicka, Maria Rudawska, Tomasz Leski2

Abstract

Ectomycorrhizae of Scots pine (Pinus sylvestris L.) trees grown in forests influenced by different levels of air pollutants

were investigated. Total numbers of mycorrhizal root tips in the soil horizons and the frequency of mycorrhizal

morphotypes were compared as indicators of ectomycorrhizal status. The studies were conducted in two comparable

young pine plantations in western Poland and in two mature forest ecosystems in southern Poland differing in pollution

level. Soil at the polluted young pine plantation (Lubon) had lower pH and increased availability of aluminum ions

than the control site (Kornik). During 3 years of observations a lower number (50 percent) of mycorrhizal root tips per

soil volume were found at the polluted site compared to the control site. The percentage of particular mycorrhizal

morphotypes in the two young pine plantations was similar. The mature pine stands represented heavily-polluted

(Niepolomice Forest) and moderately-polluted (Ratanica forest catchment) forest ecosystems. The total number of

mycorrhizae in the Niepolomice Forest was lower in sites located closer to the urban-industrial area than at more

distant plots. In the Ratanica Forest catchment the total number of mycorrhizal root tips varied between the sampling

sites; however, on an average it was considerably higher than in the Niepolomice Forest. The mycorrhizal diversity at

the heavily-polluted stand was reduced ( four mycorrhizal morphotypes) as compared to the moderately-polluted site

(eight mycorrhizal morphotypes). The results indicated that ectomycorrhizae of mature Scots pine trees were more

affected by environmental pollutants than mycorrhizae at young pine plantations.

Introduction

Acidification of forest soils caused by acid deposition of sulfur and nitrogen

compounds is considered the main factor of forest decline in Northern Europe

(Ulrich 1983). At low pH (< 4.5) the rates of dissolution of minerals and solid

complexes increase and the chemistry of soil changes (Ritchie 1995). Increased

amounts of soluble aluminum, manganese, and other metals can reach

concentrations that may be toxic to plants. At the same time, base cations (Ca2+,

Mg2+, Na+, K+) are also solubilized and easily leached from the upper rooting horizon

of the soil. It is believed that aluminum toxicity is related to increased Ca/Al ratios

in the forest soil rather than pure Al3+ concentration (Ulrich 1983).

Ectomycorrhizae are an integral part of the root system of Scots pine (Pinus

sylvestris L.), the dominant forest species in Poland. Mycorrhizae improve growth

of plants by enhancing uptake of water and nutrients (particularly phosphorus)

and are totally dependent on plant derived carbohydrates transported from shoot

to root. Mycorrhizal Scots pine seedlings have been shown to be more tolerant to

aluminum ions than non-mycorrhizal plants (Goransson and Eldhuset 1991). It is

considered that ectomycorrhizae may protect tree roots from the effects of environmental pollution, acting as the physical and/or chemical barrier. On the other hand,

ectomycorrhizae and ectomycorrhizal fungi may be negatively influenced by indirect factors, such as low photosynthetic rates after yellowing of needles or needle

loss, or direct factors, such as low pH and high soluble aluminum concentrations,

low availability of nutrients, elevated soil nitrogen, or drought. Soil texture, aeration, and humus compounds are also important factors affecting mycorrhizae

(Kottke and others 1993).

Changes in the chemical composition of soil affect the production of fine roots,

which potentially could develop mycorrhizae. A significant decrease of fine root

biomass in soils degraded by environmental pollutants was documented for Scots

pine (Jozefaciukowa and others 1995, Munzenberger and others 1995). Some authors indicated that environmental pollutants reduce the number of mycorrhizal

root tips and the diversity of mycorrhizal morphotypes (Dighton and Skeffington

1987, Kowalski 1987, Kowalski and others 1989).

Ectomycorrhizae of Young and Mature Scots Pine Trees

in Industrial Regions in Poland1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Plant Physiologist, Polish Academy of

Sciences, Institute of Dendrology,

Parkowa 5, 62-035 Kornik, Poland.
206 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Forest ecosystems in Poland have been under pressure of heavy industrial pollution, mainly SO2

, NOx, and heavy metals, for about half a century (Godzik 1990).

Although in recent years the levels of toxic pollutants emitted to the atmosphere

have decreased, the southwestern part of Poland remains one of the most polluted

areas in Europe (Tickle and others 1995). Toxic pollutants, such as heavy metals, accumulated in the contaminated soils and have affected the terrestrial

and aquatic ecosystems.

This paper describes a study that compares the mycorrhizal status of Scots

pine trees in forest ecosystems influenced by varying intensities of environmental

pollution, using total number of ectomycorrhizal root tips in a soil volume and the

diversity of mycorrhizal morphotypes as indicators.

Materials and Methods

Study Areas

The investigations were conducted at four Scots pine stands in Poland (fig. 1) influenced by different deposition rates of air pollution. At Lubon and Kornik young

Scots pine trees planted in 1984 as 2-year-old seedlings were studied. The plot at

Lubon is located about 2 km from a phosphate fertilizer plant established in 1917.

The plant has emitted continual increasing levels of toxic air pollutants (Hernik

1987). In 1980 the emission of toxic gases were estimated: sulfur dioxide at 2,806 t

yr -1, fluorides at 162 t yr -1, and nitric oxides at 850 t yr -1. In recent years (since 1987)

the amount of toxic pollutants emitted to the atmosphere has decreased considerably compared to previous years. In 1994 mean soil pH in the soil horizon 0-5 cm

was: pHwater = 4.2, pHsalt = 3.7, and the Ca/Al ratio was 0.14 (Rudawska and others

1995). The Kornik stand is situated 12 km to the southeast of Lubon in the experimental forest of the Institute of Dendrology. This area, free of direct industrial pollution, was a control site. Soil at the control site was more than 10-fold less acidic

than at the polluted site (mean pHwater = 5.8, pHsalt = 4.9). The Ca/Al ratio in the soil

horizon 0-5 cm was 17 (Rudawska and others 1995). At the two young Scots

pine plantations mycorrhizas were investigated, from autumn 1992 to autumn

1995, in the top 5 cm of the soil horizon. In 1995 additional samples from deeper

soil layers were taken.

At the Niepolomice Forest and at Ratanica forest mature Scots pine trees (50-80

years old) were investigated. The Niepolomice Forest ecosystem has been affected

for more than 40 years by gaseous and dust pollutants emitted by the adjacent

urban-industrial area of Krakow (15-30 km west of the margin of the Forest) and by

Figure 1 — Experimental Scots

pine plots in Poland.

Kornik

Lubon

Niepolomice Forest

Ratanica

'

'

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 207

the distant industrial area (Upper Silesian Industrial District — one of the most

polluted areas in Central Europe, 70 km west of the Forest). The forest complex has

been influenced by SO2, NOx, CO, CO2, and heavy metals, including Fe, Zn, Pb, Cu,

Cd, Ni (Manecki 1984). The average pH of soil in the upper 5 cm in 1994 was:

pHwater = 4.3, pHsalt = 3.3. Scots pine is the major tree species of the forest stand,

comprising 71 percent of the total area (Grodzinska 1984). Four sites (94, 124, 161,

and 232) located on a west-east transect from Krakow were chosen as the experimental plots. Mycorrhizae were assayed in 1994, three times per year, at 0-5 cm and

5-10 cm soil depth.

The Ratanica experimental plot is located 40 km south of the Krakow urban

area. It is a small forest catchment (88 ha) in the Carpatian foothills in the region of

a drinking water reservoir, surrounded by intensively managed agricultural land.

The catchment is influenced by gaseous and dust pollutants (containing SO2 , NO2,

Cl, Mn, Zn, Cu, Pb, Cd) transported there by winds from several distant industrial areas and by local agricultural emissions from surrounding villages

(Niklinska and others 1995, Szarek 1995). Average pH in the organic horizon

was reported as pHwater = 3.9 and in the mineral layer as pHwater = 4.3 (Grodzinska

and Szarek 1995). The level of air pollution in this area was qualified as moderate but

chronic (Grodzinska and Szarek 1995). The catchment is covered by a mixed forest

with a predominance of Scots pine (Pinus sylvestris L.) and beech (Fagus sylvatica L.).

Mycorrhizal Frequency

Fine-root sampling was carried out with a 16 cm3 soil corer from three places around

a pine tree, at about 60-80 cm distance from the trunk. Samples were taken three

times per year (May, August, October). At Lubon and Kornik 24 young trees at

each plot were tested. At Niepolomice Forest six mature trees in each of the forest

sites were investigated. At the Ratanica forest catchment three to seven mature

Scots pine trees were studied in each of five areas of the entire catchment. Samples

were taken to the laboratory and stored at 3 °C until analyzed. Roots were washed

over a mesh, sorted, and analyzed under a stereomicroscope. The number of mycorrhizal root tips was counted in the root samples.

Characterization of Ectomycorrhizal Morphotypes

Each root tip was characterized according to morphological traits (shape, color,

surface texture). The ectomycorrhizal morphotypes were distinguished and named

according to the main types of ectomycorrhizal systems presented by Agerer (1987-

1995) (fig. 2).

● Type A: Simple (unramified), elongate my corrhizal root

tips, with a thin fungal mantle, their diameters were similar to

non-mycorrhizal roots, yellow-brown in color, the surface was

more or less smooth.

● Type A:1 Simple, elongate, thick mycorrhizae, their diameters were mostly three times the diameter of the supporting

roots, light yellow in color, abundant fungal mantle with

extramatrical hyphae on the surface.

● Type B: Dichotomous, elongate, thin mycorrhizae, orange-brown in color. The fungal mantle thin with a very smooth

surface, no evidence of extramatrical hyphae.

● Type C: Shortened, abundantly dichotomizing mycorrhizae, light yellow, with extramatrical mycelium.

● Type D: Stumpy coralloid mycorrhizae with thickened

yellow-brown branches. The surface was more or less smooth,

without extramatrical hyphae. Older forms were darker in color

and had a wrinkled appearance.

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
208 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

● Type E: Cenococcum sp. characteristically jet-black mycorrhizae with abundant emanating black hyphae stiffly projecting around the root tip. Infection by the fungus was also overgrowing other types of mycorrhizae, as A and B. Type

● Type F: Tuberculate mycorrhizae with a thick mantle and

a smooth surface.

● Type G: Monopodial-pinnate mycorrhizae with thick fungal mantle, yellow or yellow- brown, with a smooth surface.

● Type H: Monopodial-pyramidal mycorrhizae with thick

mantle, white-yellow or yellow- brown, smooth or grainy surface, old forms brown, with wrinkled surface.

Statistics

Data were processed using Duncan’s multiple range test (ANOVA) or Scheffe test

(Statistix 4.1). Statistical significance was accepted at P < 0.01.

Results

Young Plantations of Scots Pine (Lubon and Kornik)

At the polluted site (Lubon) the total number of mycorrhizal root tips in the upper

soil horizon (0-5 cm), expressed per volume of soil sample, was significantly lower

than at the control site in Kornik (fig. 3), although at both sites non-mycorrhizal

roots were hardly found. The dry mass of fine roots (< 2 mm diameter) was slightly

lower in soil samples from the polluted than from the control site; however, the

differences were not significant (data not shown). In the 15-30 cm soil depth the

average number of mycorrhizal root tips per soil volume was a little higher (not

significantly) at the polluted than at the control plantation (fig. 4).

Mycorrhizal morphotypes distinguished on pine roots at the experimental plots

(fig. 2), corresponded to the main types of ectomycorrhizal systems described by

Agerer (1987-1995). At the control plantation at Kornik and the polluted site at

Lubon, a maximum of seven morphotypes (A, A1

, B, C, D, E, F) was found (table 1).

At both sites, the most frequent were the simple (unramified) mycorrhizae A and

A1. The frequency of the thin A type, considered as the less efficient, was similar at

both plots. The dichotomous mycorrhizae with abundant fungal mantle (type C)

and coralloid mycorrhizas (type D) were present at both polluted and unpolluted

plantations at low frequencies.

Figure 2 — Ectomycorrhizal

morphotypes distinguished on

Scots pine roots at the

experimental plots:

A, A1

 - simple mycorrhizae;

B, C - dichotomous mycorrhizae;

D - coralloid mycorrhizae;

E - mycorrhizae with Cenococcum

 sp.;

F - tuberculate;

G - monopodial-pinnate;

H - monopodial- pyramidal.

A A1 B C D

E F G H

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 209

Mature Pine Trees (The Niepolomice Forest and

Ratanica Forest Catchment)

In the Niepolomice Forest the total number of ectomycorrhizae was lower at the

sites located closer to the urban-industrial centers than at the more distant sites,

however, in August and October, in the soil depth 0-5 cm, the differences were not

significant statistically (fig. 5). Four mycorrhizal morphotypes (A, A1, B, D) were

found on pine roots in the Niepolomice Forest (table 2). Generally, Scots pine roots

developed at these forest sites mostly simple, thin mycorrhizae A (up to 84 percent). The dichotomous mycorrhizae (B) and coralloid morphotypes (D) were typically a low percentage of the mycorrhizae in the plots, with a few exceptions.

Considerable variations in the total number of mycorrhizae per soil volume

(from 360 to 2,030) were found among 22 pine trees tested in the Ratanica forest

catchment, however, the mean values count for various sites of the forest were

similar or differed slightly (fig. 6). The average number of mycorrhizal root tips in

the Ratanica forest catchment, in soil depth 0-5 cm, was more than 100 percent

higher in relation to the Niepolomice Forest.

Figure 3 — Total number of

ectomycorrhizal root tips in soil

samples (100 cm3

) taken from a

0-5 cm depth at the control

(Kornik) and the polluted

(Lubon) Scots pine plantations,

from 1992 to 1995, three times

per year. Data are means of 24

replicates. The differences

between the sites are statistically

significant (P < 0.01, Scheffe test).

Figure 4 — Total number of

mycorrhizal root tips in soil

samples (100 cm3

) taken from a

15-30 cm depth at the control

(Kornik) and polluted (Lubon)

Scots pine plantations in 1995.

Data are means of 24 replicates.

The differences between the

sites are not significant

statistically, according to the

Scheffe test.

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
210 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

1 Data are means + SD (n = 24).

Table 1 — Frequency of mycorrhizal morphotypes of Scots pine at two soil depths: 0-5 cm and 15-30 cm, at control (Kornik) and p Table 1 olluted (Lubon) plantations. Root

samples were observed in May, August, and October 1995.1

Time Site Percentage of mycorrhizal morphotypes

A A1 B C D E F

Soil depth 0-5 cm

05/95 Kornik 61 + 16 21 +11 7 + 3 1 + 1 5 + 5 0 5 + 5

Lubon 53 + 17 20 +14 13 + 8 1 + 1 10 + 3 0 3 + 3

08/95 Kornik 74 + 7 5 + 3 14 + 6 2 + 2 3 + 2 2 + 2 0

Lubon 60 + 13 8 + 7 23 + 10 1 + 1 7 + 7 1 + 1 0

10/95 Kornik 73 + 7 10 + 5 6 + 3 3 + 2 5 + 5 3 + 3 0

Lubon 59 + 15 14 + 7 9 + 3 7 + 7 8 + 3 3 + 3 0

Soil depth 15-30 cm

05/95 Kornik 67 + 22 10 +10 9 + 7 1 + 1 9 + 9 0 3 + 3

Lubon 60 + 24 11 + 11 12 + 9 0 17 + 12 1 + 1 0

08/95 Kornik 79 + 9 1 + 1 13 + 7 0 6 + 4 1 + 1 0

Lubon 60 + 13 1 + 1 17 + 4 0 20 + 10 0 2 + 2

10/95 Kornik 71 + 17 14 + 14 9 + 9 1 + 1 4 + 4 1 + 1 0

Lubon 47 + 27 12 +10 10 + 6 4 + 4 15 + 12 0 12 + 12

Figure 5 — Total number of

mycorrhizal root tips in soil

samples (100 cm3

) taken from

0-5 cm and 5-10 cm depths at

four sites (94, 124, 161, 232) in

the Niepolomice Forest, located

along a transect extending from

the Krakow area, the source of

air pollutants. Data are means of

six replicates. Means followed by

the same letter do not differ

significantly (P < 0.01, ANOVA,

Duncan’s test).

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 211

Eight morphotypes of pine ectomycorrhizae were distinguished in the Ratanica

forest catchment (table 3). The average frequency of the simple, thin mycorrhizas

(type A), considered as the less beneficial for a tree, was lower than 50 percent.

Occurrence of new morphotypes, compared to the Niepolomice Forest was observed: abundantly dichotomizing (C), monopodial-pinnate (G), monopodial-pyramidal (H), and Cenococcum sp. mycorrhizae (E).

Discussion

Mycorrhizal status of forest trees provides accurate information about changes in

the forest soil affected by environmental pollution. In this study reduced numbers

of Scots pine mycorrhizae were found at the severely polluted forest stands in Poland

(Lubon, the Niepolomice Forest) compared to the slightly or moderately polluted

sites (Kornik, Ratanica forest catchment) (figs. 3, 5, 6). Significant reduction in

mycorrhizae was found at 11-18 year-old Scots pine plantations (Kowalski and others

1989) and in mature Scots pine stands (Ohtonen and others 1990) influenced by

industrial pollutants. A decrease in numbers of mycorrhizae was also caused by

low pH and available aluminum in perlite/peat cultures (Metzler and Oberwinkler

1987). The influence of other important factors on mycorrhizae, like soil type and

soil moisture must also be considered (Kottke and others 1993).

Frequencies of ectomycorrhizal morphotypes are considered accurate indicators of the mycorrhizal status of forest trees. Dighton and Skeffington (1987) observed that artificial acidic precipitation and an increase of Al3+ concentration in

Time Site Percentage of mycorrhizal morphotypes

A A1 B D

I II I II I II I II

05/94 94 82 + 11 75 +19 10 + 7 7 + 7 4 + 3 16 +14 3 + 3 1 + 1

124 74 + 24 59 +25 17 + 23 8 + 7 5 + 3 6 + 5 4 + 4 27 +25

161 50 + 6 35 + 25 45 + 19 64 +32 3 + 3 1 + 1 2 + 2 0

232 65 + 16 68 +17 26 + 13 24 + 12 2 + 2 4 + 4 6 + 5 4 + 4

08/94 94 84 +13 77 +17 6 + 6 14 + 14 6 + 6 5 + 5 4 + 2 4 + 4

124 58 + 13 76 + 7 37 + 17 18 + 9 2 + 2 2 + 2 3 + 2 4 + 2

161 70 + 18 70 + 22 23 + 16 25 + 24 2 + 2 2 + 2 6 + 5 3 + 3

232 59 +30 80 + 21 32 + 12 17 + 17 4 + 3 1 + 1 5 + 4 2 + 2

10/94 94 60 + 10 73 + 9 25 +12 19 + 13 6 + 3 3 + 3 8 + 4 4 + 4

124 80 + 19 77 +127 7 +6 8 +8 5 + 5 8 + 7 9 + 8 6 + 5

161 70 +19 56 +27 28 +17 44 + 31 2 + 2 0 1 + 1 0

232 62 +19 81+13 25 +16 15 +10 8 + 5 1 + 1 5 + 4 2 + 2

1

 Data are means ± SD (n = 6)

Table 2 Table 2 — Frequency of mycorrhizal morphotypes of Scots pine at soil depths: 0-5 cm (I) and 5-10 cm (II) at four forest sites Table 2 in order of decreasing pollution exposure (94,

124, 161, 232) in the Niepolomice Forest. 1

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
212 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

soil solution significantly reduced the branching of fine roots of Scots pine and

decreased the occurrence of two coralloid mycorrhizal morphotypes. Mycorrhizal

roots of Scots pine at highly contaminated areas in Poland occur mainly as simple

mycorrhizae with a scant fungal mantle, whereas in forests with low-level industrial pollution, mycorrhizae are more commonly of the dichotomous morphotypes

(Kowalski 1987, Kowalski and others 1989).

In the present studies, no evident differences in the proportion of

ectomycorrhizal morphotypes were found between the polluted and the control

young Scots pine plantations (table 1); however, considerable differences in the

occurrence of mycorrhizal morphotypes were observed between the severe-polluted and moderately-polluted mature pine trees (tables 2, 3). Simultaneously,

at the polluted young pine plantation we recorded a more abundant occurrence of species and carpophores of ectomycorrhizal fungi (Rudawska and others [In press]). Termorshuizen and Schaffers (1987) observed that mycorrhizal

fruitbodies were not depressed by air pollution in young stands of Scots pine (5-13

years), but they were reduced in mature stands (50 years). This phenomenon could

be explained by relatively low susceptibility of mycorrhizal fungi species living in

symbiosis with young Scots pine trees (Shaw and others 1992). Ectomycorrhizal

fungi show differential responses to Al in pure cultures (Kieliszewska-Rokicka and

others [In press], Thompson and Medve 1984). Since Scots pine tolerates a wide

range of ecological conditions and it is also classified as relatively tolerant to soluble

Al, (Schaedle and others 1989) the mycorrhizae formation could be limited by the

sensitivity of fungi rather than of the plant (Metzler and Oberwinkler 1987).

Reduced mycorrhizal diversity in the Niepolomice Forest compared to the

Ratanica forest catchment suggests a lower biodiversity of mycorrhizal species at

the severely-polluted mature pine stand than in the moderately-polluted forest.

Kowalski and others (1989) reported that the reduction in diversity of mycorrhizal

morphotypes at a contaminated Scots pine plot was accompanied by significant

changes in the composition of mycorrhizal fungi species and a decrease in the production of fruitbodies. Termorshuizen and Schaffers (1991) have found a negative

correlation between SO2

 concentration in the air and the number of mycorrhizal

species and carpophores in mature Scots pine forests in The Netherlands. Carpophores of mycorrhizal fungi and fungal species have declined during this

century in Europe (Arnolds 1991). The disappearance of various species of

mycorrhizal fungi has been observed also in many areas in Poland (Wojewoda

and Lawrynowicz 1986).

Environmental pollutants undoubtedly affect mycorrhizae directly, inhibiting

the uptake of nutrients and causing nutrient imbalance in root and shoot (Janhunen

and others 1994) or limiting growth of the mycorrhizal root tips (Metzler and

Figure 6 — Total number of

mycorrhizal root tips in soil

samples (100 cm3

) taken from a

0-5 cm depth at five sites in the

Ratanica forest catchment. Data

are means of three to seven

replicates. Means followed by the

same letter do not differ

statistically (P < 0.01, ANOVA,

Duncan’s test).

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 213

Oberwinkler 1987). At young Scots pine plantation contaminated by soluble Al, an

increased Al/Ca ratio in needles was correlated with a decrease in the net

photosynthesis, biomass production, and allocation to roots (Reich and others 1994).

Simultaneously, a high, negative correlation between aluminum content in pine

needles and the number of mycorrhizal root tips in the upper 5 cm of soil has been

recorded for the young pine trees (Rudawska and others 1995). Other external

factors, like drought, frost, and pathogens could intensify the effects of

environmental pollutants.

Conclusions

The decrease in the number and diversity of mycorrhizae at the heavily polluted

sites is presumably related to a direct negative effect of aluminum and other metals

on fungal hyphae as well as the influence of pollutants on photosynthesis and

transport of carbohydrates to roots. Mycorrhizae of mature Scots pine trees seem to

be more sensitive to the stress factors than mycorrhizae of young trees. As Scots

pine changes ectomycorrhizal partners during its ontogeny, a modification of

mycorrhizal status with the age of the plantation could be possible.

Acknowledgments

This study was in part supported by the KBN (Poland) grant no. 569/PO4/95/08.

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of threatened macrofungi in Poland. Warsaw: PWN; 45-82.

Session I1I Ectomycorrhizae of Young and Mature Scots Pine Trees Kieliszewska-Rokicka, Rudawska, Leski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 215

Abstract

In the last 10 years, 3.5 percent of the tree population died annually in Poland’s largest and most polluted cities, which

is a problem of economic importance. Dieback of streetside trees in Warsaw is a long term process. It is an effect of

biological reactions of trees to unfavorable conditions in the urban environment, particularly air and soil pollution

and water deficiency. The process is being intensified by microclimatic changes leading to xerism of the environment

(lowering of air humidity and rising of air temperature). To study this problem Crimean linden (Tilia euchlora K.

Koch) trees were examined. Crimean linden, constituting over 40 percent of Tilia in urban areas, and used in the

reconstruction of urban plantings after World War II at the beginning of 1950. Observation of phenologic development

of trees indicated a shortening of their vegetation period (up to 30 days) depending on the proximity to the city center.

The dendrometric examinations revealed a reduction of trunk diameter of the affected trees. These changes were

accompanied by morphological dieback of leaves observed in the beginning of July. Significant correlations were found

between the leaf damage and chemical concentration of some nutrients and pollutant elements.

Introduction

During the last 15 years, urban forestry research in Warsaw, Poland, has focused on

applied problems. The results may be of interest for specialists evaluating specific

ecological conditions of urban areas and health of forest trees in central and eastern

Europe.

Change of climatic conditions in urban areas is a part of the recently observed

climatic changes on a global scale. Unfortunately, climatic changes caused by urbanization, have led to extensive die-back of urban trees. In the beginning of the

1980’s, in western Europe, the estimated number of trees dying annually was over

830,000 (Fluckinger and Braun 1981). Poland also faces the same problem: in the

last 10 years up to 3.5 percent of the tree population has died annually in the

country’s biggest and the most polluted cities (Suplat 1989).

During periods of rapid urbanization, local scale monitoring of changes in environmental pollution and their impact on trees can be achieved on a shorter time scale

than at global scales. The rate of changes in soil and air pollution is faster in urban

ecosystems than in forest ecosystems. Our research suggests that climatic and environmental changes adversely impact urban trees at a time scale of 1 to 2 decades.

This paper discusses our long term research to define the biological reaction

of trees to specific environmental pollutants and climatic conditions in urban

areas in Poland.

Objectives and Methods

The first phase of research was to determine biological responses to urban temperature anomalies. We applied a 15-year phenological monitoring program from

1973 to 1988 in the Warsaw area. This program was also part of a study to determine the chronic effects of pollution on urban trees.

More than 100 Crimean linden (Tilia euchlora K.Koch) trees, 30-40 years of age

and growing on 12 examined areas, were monitored from 1973-1988. The areas

represented different habitat conditions and included parks, city squares, and streets

(fig. 1). Observations were made of phenological development, chemical composition of leaves, and trunk growth by diameter at breast height (DBH)

(Chmielewski 1990).

Waldemar Chmielewski, Wojciech Dmuchowski, Stanislaw Suplat2

Impact of Urban Environmental Pollution on Growth,

Leaf Damage, and Chemical Constituents of Warsaw

Urban Trees1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Ecologist, Plant Chemist, Plant Physiologist, respectively, Botanical Garden, Polish Academy of Sciences, ul.

Prawdziwka 2, 02-973 Warszawa,

Poland.
216 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

In the second research phase, 143 Crimean linden trees were analyzed in 1995

for health (e.g., leaf damage) and their nutritional status. The trees were located in

an ecologically homogenous habitat and were exposed to extensive traffic and soil

pollution (such as de-icing salt).

Measurements included evaluation of tree condition by determining the index

of leaf damage in relation to tree-crown condition (United Nations Economic Commission for Europe 1994): the seasonal dynamics of the foliar nutrients, status of

trees showing various damage levels, and dendrometric measurements (such as

DBH). Every 2 weeks during the vegetative season (May-September), leaf damage,

their chemical constituents, and tree-stem diameter were measured.

Results and Discussion

The vegetation period is defined as the beginning of leaf-buds bursting to the date

of leaves falling. Results suggest that over the period 1973-1988, environmental

pollution affected the length of in-leaf season for urban trees (fig. 2). Twelve study

areas were arranged along an urban-suburban gradient. The in-leaf season for trees

located in the city center was shorter than for trees in suburban areas. The average

difference between habitats located in the city center (Konstytucji Square) and at

the outskirts (Ursynów) was 25 days. Similar results for other tree species were

reported by Feige and others (1980), Supuka (1988), and Zacharias (1972), who also

attribute the shortening of the vegetation season to urbanization and degradation.

Session I1I Impact of Urban Environmental Pollution on Growth, Leaf Damage Chmielewski, Dmuchowski, Suplat

Figure 1 — Distribution of

observation plots (habitats) in

Warsaw along the north-south

gradient. Outskirts and large city

parks: 1—Bielanski Forest;

2—Towarowa street;

8—Solders Cemetery;

9—Zwirki, Wigury street;

10—Ujazdowskie street;

11—Lazienki Park;

12—Ursynow.

Squares, streets, and parks in

the city center:

3—Saski Garden;

4—Krolewska street;

5—Marszalkowska street;

6—Konstytucji Sq.;

7—Unii Lubelskiej. Sq.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 217

For trees in streets and city squares, there was also a gradual shortening of the

vegetative period throughout 1973-1982 period. Average reduction in the length of

the vegetative period for 12 habitats was about 30 days, with considerable fluctuations between the individual years.

In the population of examined trees, the rate of leaf damage was increasing

throughout the vegetative season. By the end of July, only 30 of 143 examined trees

showed leaf damage while the index of damage ranged from 6 to 75 percent. Analyses of the chemical composition of tree leaves revealed differences between the

trees of various leaf damage indices.

A seasonal variation was found in the chlorine accumulation in the Crimean

linden (Tilia euchlora K.Koch) leaves (fig. 3). The trees with strongly damaged leaves

showed high chlorine concentrations immediately after the leaves emerged. Leaf

chlorine concentrations also increased during the vegetative season. In the group

of trees not showing leaf damage, leaf chlorine concentrations at the beginning of

the vegetative season were relatively low, but increased with the time. Leaf chlorine concentrations in healthy trees located at the city outskirts (e.g., Bielanski Forest) were more than three times lower than in unhealthy urban tree leaves that

were damaged (Zwirki, Wigury Street). At the suburban site, the rise in chlorine

concentrations during the vegetative season was slight. According to Pauleit (1988),

healthy Crimean linden leaves contain 0.27-0.30 percent chlorine. For damaged

leaves the level is 1.0 to 6.0 percent (Ruge and Stach 1968).

The results of the chemical analysis of leaves did not indicate any deficiency of

the nutrient elements (Chmielewski and others 1994). However, higher than normal physiological levels of chlorine, sodium, sulfur, lead, and copper concentrations were measured, suggesting that the ionic equilibrium of unhealthy trees was

disturbed. The measured chlorine concentrations could be characterized as toxic.

However, nutrient ratios are also significant indicators of crown condition

(Brogowski and others 1988, Packham and others 1992).

To characterize the ionic equilibrium of tree tissues, the gram-equivalent values were used. In the interpretation of research results of this study, the gramequivalent values we used included relation of bivalent cations to monovalent catFigure 2 — Two tendencies

(1973-1982) in the vegetation

period of Crimean linden trees

(Tilia euchlora K. Koch) in the 12

examined habitats in Warsaw.

DAYS

220

200

180

160

140

1973 1975

1977 1979

YEARS 1981

1

2

3

4

5

6

7

8

9

10

11

12

suburb-downtown-suburb

Session I1I Impact of Urban Environmental Pollution on Growth, Leaf Damage Chmielewski, Dmuchowski, Suplat
218 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

ions, total cations, relation of calcium to magnesium, and ANE (accumulated nutrient elements—total of all examined elements) (Ostrowska and others 1988).

Measurements were made of tree stem diameter at the breast height (DBH)

and the index of leaf damage from July and September. The DBH ranged from 71 to

172 centimeters and was positively correlated with the leaf damage index for September. The index of leaf damage was higher in the trees containing more leaf chlorine and calcium. The leaf damage index was inversely proportional to magnesium

content (table 1).

The September leaf damage index, the DBH, and the coefficient of ionic equilibrium (independent variables) were correlated (table 2). There was no correlation

between DBH and ionic equilibrium in trees. The leaf damage index was positively

correlated with the ANE indicator and with the high calcium to magnesium ratio

given in gram-equivalent values.

These are preliminary results. Detailed statistical analysis of current data,

including anatomic studies of cambium, phloem, and xylem should be done in

future studies to further determine the effects of urban environments on trees

in Poland.

Figure 3 — Seasonal changes

of chlorine accumulation in the

Crimean linden tree leaves in

three classes of leaf damage.

Session I1I Impact of Urban Environmental Pollution on Growth, Leaf Damage Chmielewski, Dmuchowski, Suplat
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 219

References

Brogowski, Z.; Kocon, J.; Kowalkowski, A. 1988. Preliminary studies on supply of mountain spruce-trees with mineral compo- Preliminary studies on supply of mountain spruce-trees with mineral components and their health condition. nents and their health condition. Polish Journal of Soil Science XXI/I: 51-60. nents and their health condition.

Chmielewski, Waldemar. 1990. Wykorzystanie wieloletnich obserwacji fenologicznych Tilia euchlora K.Koch. oraz analizy

skladu chemicznego li_ci do bioindykacyjnej oceny stopnia degradacji siedlisk miejskich w Warszawie. skladu chemicznego li_ci do bioindykacyjnej oceny stopnia degradacji siedlisk miejskich w Warszawie. Warszawa: Agri- skladu chemicznego li_ci do bioindykacyjnej oceny stopnia degradacji siedlisk miejskich w Warszawie.

culture University; 120 p. Ph. D. dissertation.

Chmielewski, Waldemar; Dmuchowski, W.; Suplat, S. 1994. Stan i rozwoj drzew w warunkach miejskich Warszawy. Stan i rozwoj drzew w warunkach miejskich Warszawy. Annual Stan i rozwoj drzew w warunkach miejskich Warszawy.

research report. Warsaw: Botanical Garden of the Polish Academy of Sciences; 85 p.

Feige, G.; Geyer, M.; Kremer, B.P. 1980. Die Linden der Stad Koln - Bioindikatoren des Stadklimas und der anthropogenen

Umweltvernderungen. Umweltvernderungen. Decheniana 133:197-198. Umweltvernderungen. 

Fluckinger, W.; Braun, S. 1981. Perspectives of reducing the deleterious effect of de-icing salt upon

vegetation. vegetation. Plant and Soil 63: 527-529. vegetation. 

 Packham, J.R.; Harding, D.J.L.; Holton G.M.; Stuttard. 1992. Functional ecology of woodlands and forests. Functional ecology of woodlands and forests. London: Chapman Functional ecology of woodlands and forests. 

and Hall; 353-354.

Ostrowska, A.; Szczubialka, Z.; Gawlinski, S.; Paul, M. 1988. Application of the ANE value and the share of Application of the ANE value and the share of particular ele- particular elements in this value for appraising the state of supplying pine trees (Pinus silvestris L.) with the nutrient elements. Annales

of Warsaw Agriculture University. of Warsaw Agriculture University. Forest and Wood Technology 37: 31-40. of Warsaw Agriculture University. 

Pauleit, S. 1988. Vitalitatskartirerung von Stadtbaume in Munchen. Vitalitatskartirerung von Stadtbaume in Munchen. Garten and Landschaft 7: 38-40. Vitalitatskartirerung von Stadtbaume in Munchen.

Ruge, U.; Stach, W. 1968. Injury to roadside trees caused by thawing salt. Injury to roadside trees caused by thawing salt. Angew. Botany 42: 69-77. Injury to roadside trees caused by thawing salt. 

Supuka, J. 1988. Comparative phenology of urban woody species—reflection of changed ecological conditions. Comparative phenology of urban woody species—reflection of changed ecological conditions. Folia Comparative phenology of urban woody species—reflection of changed ecological conditions.

Dendrologica 15: 267-285.

United Nations Economic Commission for Europe. 1994. Manual on methods and criteria for harmonized sampling, assess- Manual on methods and criteria for harmonized sampling, assessment, monitoring and analysis of the effects of air pollution on forests. ment, monitoring and analysis of the effects of air pollution on forests. Brussels, Geneva, Vienna. ment, monitoring and analysis of the effects of air pollution on forests.

Zacharias, F. 1972. Bluthphaseneintritt an Strassenbaumen besondere (Tilia euchlora K. Koch.) und Temperatueverteilung

in Westberlin. in Westberlin. Berlin: Der Freien Universitat Berlin. Fachbereich Biologie. 240 p. Ph. D. dissertation. in Westberlin. 

Table 1 Table 1 — Correlations between the index of damaged Crimean linden tree ( Table 1 Tilia euchlora K.Koch) leaves, the

tree-stem DBH (dependent variables), and the concentration of selected elements in leaves.1

 Dependent Chlorine Sodium Magnesium Calcium

 Variables

Index of leaf damage + 0 - +

Decrease in

tree-stem DBH + - + -

1

 0 = nonsignificant correlation

 + = positive significant correlation

 - = negative significant correlation

Table 2 Table 2 — Correlation between the index of damaged Crimean linden tree ( Table 2 Tilia euchlora K.Koch) leaves, the tree-stem

DBH (dependent variables), and the coefficient of ionic equilibrium (independent variables) in gram-equivalent values.

Dependent A1 B C D E

 Variables

Index of

leaf damage 200 + + +

Decrease in

tree-stem DBH 0000

1

A = ratio of bivalent cations to monovalent cations (Ca+2 + Mg+2/K+ + Na+)

B = sum of cations (Ca+2 + Mg+2 + K+ + Na+)

C = ratio of calcium to magnesium (Ca+2 / Mg+2)

D = ANE accumulation nutrient elements

E = tree-stem DBH.

20 = nonsignificant correlation

+ = positive significant correlation (p < 0.05)

- = negative significantcorrelation (p < 0.05)

Session I1I Impact of Urban Environmental Pollution on Growth, Leaf Damage Chmielewski, Dmuchowski, Suplat

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 221

Patrick J. Temple and Paul R. Miller2

Abstract

Ambient ozone was monitored from 1992 to 1994 near a forested site dominated by mature Jeffrey and ponderosa

pines (Pinus jeffrey Grev. & Balf. and Pinus ponderosa Dougl. ex Laws.) at 2,000 m in the San Bernardino Mountains

of southern California. Ozone injury symptoms, including percent chlorotic mottle and foliage retention, were evaluated

on 130 mature pines in September of 1991 to 1994 at a site 2 km from the ozone monitor. Symptom measurements were

combined in an ozone injury index (OII) that represented the average crown condition of each plot. Leaf stomatal

conductance and available soil water were measured biweekly from March to November at one of the plots from 1992

to 1994. Mean annual ozone concentrations were relatively constant for the 3 years. However, the distribution of

individual trees in the 0 to 100 range of the OII showed a shift from 1991 to 1994 to higher frequencies of trees in

classes representing greater amounts of injury. Compared with 1991, in 1993 and 1994 more trees remained in the 

higher injury classes. Maximum rates of stomatal conductance and maximum ozone uptake occurred in June of each

year, but the total annual ozone flux varied as the amount of available soil water varied with seasonal precipitation. 

The period of April to June or May to July, depending upon year, accounted for 70 to 75 percent of annual ozone flux,

confirming the high correlation observed between ambient ozone concentrations from April to June and development

of chlorotic mottle on the needles. Foliar injury to current-year needles, which served as a marker of ozone response for

a single exposure season, was greatest in 1992, coincident with higher flux of ozone to foliage in 1992, compared with

1993 and 1994. These data confirm that the period of maximum ozone impact on these pines is early in the growing

season, and that ambient ozone after July had less effect on the development of foliar ozone injury on older pine needles.

Year-to-year variation in ozone injury to pines was influenced both by the status of soil moisture availability in summer

at this location and the internal foliar dynamics of the trees as they recovered from drought. 

Introduction

Annual precipitation throughout the ranges of ponderosa and Jeffrey pines (Pinus

ponderosa Dougl. ex Laws. and Pinus jeffreyi Grev. & Balf.) in the mountains of

California is subjected to strong seasonal influences, with more than 90 percent of

total precipitation falling in the months of October to April. Annual precipitation

totals are also highly variable, and the long-term annual mean is composed of many

years with low total precipitation and a few years of much greater than average

precipitation (NOAA 1995). Summer drought is the norm, interrupted by occasional

brief, local thunderstorms. The growth cycles of the trees have become adapted to

these highly variable conditions, and gas exchange in the pines closely followed

patterns of soil water availability and vapor pressure deficit (Helms 1970, 1971).

Because the uptake of ozone by plants is controlled by stomatal conductance

(Runeckles 1992), annual and seasonal patterns of ozone uptake in ponderosa and

Jeffrey pines may also be mediated by environmental factors, such as soil water

availability and vapor pressure deficit, that control gas exchange (Helms 1971).

Intrinsic factors, such as leaf age (Helms 1970) and amount of foliar ozone injury

(Coyne and Bingham 1982), also influence the rate of gas exchange in ponderosa

pines. Those factors that influence stomatal conductance could also affect the response of the tree to ozone and could be used to model ozone uptake and foliar

injury responses of trees in the field. Thus, measurements of stomatal conductance

and calculations of ozone flux to leaves may be used to bridge the gap between

measurements of ambient ozone concentrations in the forest and measurements of

foliar injury symptoms on pines.

The objectives of this study were to measure seasonal ozone flux to the foliage

of mature ponderosa and Jeffrey pines as influenced by environmental and

physiological factors, and to correlate ozone flux with observed changes in ozone

injury measured on mature pines at the same location in the San Bernardino Mountains of southern California.

Seasonal Influences on Ozone Uptake and Foliar Injury to

Ponderosa and Jeffrey Pines at a Southern California Site1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside, California.

2 Project Consultant and Plant Pathologist, respectively, Pacific Southwest

Research Station, USDA Forest Service, 4955 Canyon Crest Drive, Riverside, CA 92507, U.S.A.
222 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Materials and Methods

Ambient Ozone Concentrations

Continuous measurements of ozone concentrations from January 1992 to December

1994 were obtained from a meteorological station located at Barton Flats at 2,000 m

in the mixed conifer forest zone of the San Bernardino Mountains in southern

California. Ozone air quality was measured with a certified Environmental

Protection Agency (EPA)-equivalent ozone analyzer, calibrated and maintained

according to standard air quality protocols (Miller and others 1996a). Precipitation

data were obtained from a meteorological station located at Big Bear Lake, 10 km

north of the site.

Pine Plot Evaluations

Ozone injury symptoms were evaluated each September from 1991 to 1994 on 5

branches of 130 mature ponderosa or Jeffrey pines located at an elevation of 2,000

to 2,200 m, 2 km from the meteorological station. The number of annual needle

whorls was counted and each whorl was evaluated for percent of needle area with

chlorotic mottle, percent of needles remaining in each whorl, and average needle

length of each whorl. The percent live crown was determined as the proportion of

the total crown with any live branches relative to the total tree height. These

parameters were combined into an ozone injury index (OII) by weighting chlorotic

mottle by 0.40, abscission by 0.40, needle length by 0.10, and percent live crown by

0.10 (Miller and others 1996b). Injury scores were averaged to obtain the average

crown condition for each plot for each year. The distribution of trees in 10 unit

increments of the 0 to 100 of the OII were also determined annually. Higher

numerical values of the OII mean greater chronic injury.

Measurements of Stomatal Conductance

Leaf stomatal conductance was measured twice monthly from March to November

and monthly from December to February of 1992 to 1994 at 25 m height in the

canopy of a stand of mature ponderosa and Jeffrey pines on one of the pine

evaluation plots. Access to the top of the trees was provided by a tower that

penetrated through the canopy to a height of 30 m. Branches from three trees were

accessible from the tower, and stomatal conductance measurements were taken on

all available age classes of needles on three branches from each of three mature

pines. Stomatal conductance was measured with a steady-state porometer (Model

LI-1600, Licor Inc., Lincoln, NE)3 equipped with a cuvette specifically designed for

conifer foliage. Needle surface area was calculated by using the diameter length of

the cuvette opening (5 cm) multiplied by the diameter of the needle fascicle and the

number of needles in the fascicle (usually three). Fascicle diameter was measured

with an electronic digital caliper (Mitutoyo Corp., Japan) to the nearest 0.01 mm.

Measurements were taken usually between 0900 and 1300, when rates of stomatal

conductance were at maximum. However, stomatal conductance measured hourly

throughout three complete diurnal cycles showed that conductance rates in mature

pines varied little during most of the daylight hours.4 Photosynthetically active

radiation, relative humidity, leaf and ambient temperature were measured

simultaneously with conductance measurements by sensors on the porometer

cuvette. Volumetric soil water was measured bi-weekly under the tree canopy, except

during months with snow cover, using a time domain reflectometry instrument

(TRASE Model 1010X, SoilMoisture Corp., Santa Barbara, CA) equipped with 15

cm soil probes.

Ozone Flux

Flux of ozone to pine foliage was calculated as:

Flux (O3)(μmol m-2 d-1) = [O3]a (μmol mol-1) x gs(O3) (mol m-2 d-1)

in which [O3

]a = ambient ozone concentration per day, averaged over the number

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

4 Unpublished data on file at the Pacific

Southwest Research Station, Riverside, Calif.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 223

of daylight hours from 30 min after sunrise to 30 min before sunset. Stomatal conductance to ozone (gs

(O3)) was calculated from gs(H2O) by multiplying by the ratio

of the molecular weights of O3 to H2O. Boundary layer resistance in the well-mixed

conifer cuvette was factory-set in the instrument at 0.07 s m-1. The actual boundary

layer resistance of the measured pine needles would depend upon wind speed, but

generally at the upper layers of the canopy wind speed > 5 m s-1. Other aspects of

ozone transport from the atmosphere to internal leaf surfaces (Runeckles 1992) were

ignored for the purposes of these calculations. Mean daily ozone flux (μg m-2 d-1),

averaged over each month, was calculated from mean monthly daylight ozone concentrations and mean daily rates of stomatal conductance for each month of the 3-

year study.

Results and Discussion

Average ozone concentrations were relatively uniform at Barton Flats during the

3 years of the study. Mean monthly daylight ozone concentrations ranged from 38

parts per billion (ppb) in January to 83 ppb in June (fig. 1). Highest mean ozone

concentrations occurred in June of 1992 and 1994, but ozone levels were lower in

July and August of 1992 because of unusually cloudy and rainy conditions during

the late summer of 1992. Ambient ozone averaged 18 percent lower in July 1992

relative to July 1994; but for the 5-month period from May to September, ozone

concentrations averaged 73 ppb, 74 ppb, and 74 ppb for 1992, 1993, and 1994,

respectively.

Annual precipitation totals were highly variable at the Barton Flats site (fig. 2).

For the 4-year period preceding the study, precipitation totals averaged 50 percent

below the long-term mean of 83 cm. This prolonged period of drought was reflected in the foliage of ponderosa and Jeffrey pines at the site, which were significantly shorter and lighter in weight than needles produced after the drought had

been relieved in 1991 (Miller and others 1996a). More than 90 percent of annual

total precipitation fell during December to March of each of the three years of the

study (Miller and others 1996a).

This pattern of several years of drought before the study, followed by rapid

recovery from drought stress beginning in 1991 was reflected in the amount of

Figure 1 — Monthly 12-hour

(0600-1800) ozone averages for

Barton Flats, San Bernardino

Mountains, California, 1992 to

1994.

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller
224 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

ozone injury on the ponderosa and Jeffrey pines at Barton Flats (table 1). In 1991,

the mature pines averaged more than 4 years of needle whorl retention and the

ozone injury index (OII) averaged 38.3. By 1992, the number of age classes of needles

had decreased to an average of 3.6 and the OII had increased significantly

(p < 0.001) to 47.5, an increase of 24 percent. The OII stabilized over the next

2 years, but the number of needle whorls retained continued to decline (table

1), as the fascicles retained during the drought were replaced by larger, more

physiologically-active needles.

Distribution curves of the frequency of trees in 10-unit classes of the OII (fig. 3)

showed a shift from a relatively flat distribution in 1991 to clustering in the higher

injury classes beginning in 1992 and continuing in 1993 and 1994. Injury on current-year needles in 1992 was 10 times greater than in 1991 and three times greater

than in 1993. Injury on current-year needles was very low in the drought year of

1994 (table 1). The effect of drought in reducing the impact of ozone on these mature pines in the San Bernardino Mountains was comparable to the responses of

drought-stressed ponderosa pine seedlings exposed to ozone under experimental

conditions (Temple and others 1992).

Figure 2 — Annual total

precipitation at Big Bear Lake,

San Bernardino Mountains, 10

km north of Barton Flats, 1985

to 1995.

1 Data are means of 5 branches from 130 trees in 3 plots at the site.

Table 1 — Ozone injury index (OII), percent chlorotic mottle (CM) on current-year needles, and number of age classes Table 1

of needles retained on mature ponderosa and Jeffrey pines at Barton Flats, San Bernardino Mountains, recorded in

September 1991 to1994.1

 Year OII CM (pct) CM (pct) Age classes retained

1991 38.3 0.076 4.2

1992 47.5 0.798 3.6

1993 47.5 0.258 3.4

1994 49.2 0.005 3.3

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 225

Seasonal trends in rates of stomatal conductance of pine needles generally followed the same patterns in 1992 to 1994, although variations in total precipitation

and summer rainfall patterns that influenced soil water retention altered conductance rates, particularly in mid- to late summer. In 1992, conductance in 1-year-old

needles peaked in June and gradually declined (fig. 4), although conductance increased again in late August in response to summer rains that had increased soil

water content 2 weeks earlier. Current needles showed measurable rates of gas

exchange in July, but conductance rates in these needles remained low, relative to

maximum rates of 1-year-old needles in June. In 1993, maximum conductance rates

of 1-year-old needles peaked in early June, then gradually declined, except for small

increases in July and September in response to increased soil water content 2 to 3

Figure 3 — Distribution of 130

ponderosa and Jeffrey pines in

10 unit classes of the ozone

injury index (OII) from 1991 to

1994 (higher OII scores mean

more injury).

Figure 4 — Seasonal trends in

maximum rates of stomatal

conductance in 1-year-old

needles of mature ponderosa

and Jeffrey pines (n=3), and

volumetric soil water content

under the canopy at Barton

Flats, San Bernardino Mountains,

1992.

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller CONDUCTANCE (mol m-2 s-1)
226 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 5 — Seasonal trends in

maximum rates of stomatal

conductance in 1-year-old

needles of mature ponderosa

and Jeffrey pines (n=3), and

volumetric soil water content

under the canopy at Barton Flats,

San Bernardino Mountains, 1993.

weeks earlier (fig. 5). Current-year needles developed measurable conductance in

mid-August, and conductance rates in current needles peaked in mid-September,

but maximum conductance rates in current needles were below peak conductance

rates in 1-year-old needles. In 1994, winter precipitation (1993-1994) was 30 percent

below average, and soil water content depleted rapidly during the summer. Rates

of stomatal conductance closely followed soil water content, with peak conductance in April and May, followed by rapid reductions in conductance as the soil

dried during the summer (fig. 6). Current needles became active in early August,

but conductance in current needles remained low for the rest of the year.

These results are similar to those reported for ponderosa pine in the Sierra

Nevada, where gas exchange in photosynthesis was influenced primarily by temperature and light intensity under favorable soil water conditions. In mid- to late

summer, under high temperature and low soil water, photosynthetic rates declined

in ponderosa pine in response to increased vapor pressure deficits (Helms 1970,

1971). However, reports of higher rates of gas exchange in Jeffrey pine during winter months compared with rates in September (Patterson and Rundel 1989) were

not confirmed by these results

Highest mean daily ozone flux to 1-year-old needles of mature pines at Barton

Flats for the study period was in 1992 (fig. 7), reflecting the high stomatal conductance rates measured in 1-year-old needles in 1992. The high rates of conductance

in these 1991 needles in 1992 may have been due to the high rate of physiological

activity in these needles, as previous years’ needles produced during the drought

years were significantly smaller than 1991 needles, and had already begun to senesce

by 1992 (Miller and others 1996a). Ozone flux peaked in June, and this month accounted for 28 percent of the total yearly ozone flux in 1992. In 1993, ozone flux

was more evenly distributed throughout the growing season, but total ozone flux

averaged 40 percent less than in 1992. Flux rates were higher in 1994, although

precipitation was lower than in 1993 (fig. 2). In response to rapid decreases in rates

of conductance (fig. 6), ozone flux was higher in April 1994 but lower in July, August, and September than in 1992 and 1993. For the 3 years of the study, the months

of April to July accounted for more than 60 percent of total annual ozone flux.

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller CONDUCTANCE (mol m-2 s-1)
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 227

These results are consistent with statistical analyses of ambient ozone concentrations and development of ozone injury symptoms on pines, in which highest correlations were found between ozone injury and ozone concentrations between April

15 and July 15 (Miller and others 1996a).

Figure 7 — Mean daily ozone

flux to 1-year-old needles of

mature ponderosa and Jeffrey

pines at Barton Flats, San

Bernardino Mountains, 1992 to

1994.

Figure 6 — Seasonal trends in

maximum rates of stomatal

conductance in 1-year-old

needles of mature ponderosa

and Jeffrey pines (n=3), and

volumetric soil water content

under the canopy at Barton Flats,

San Bernardino Mountains, 1994.

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller CONDUCTANCE (mol m-2 s-1) OZONE FLUX (umol m-2 d-1)
228 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Conclusions

Results of this study of ozone injury development relative to stomatal conductance

and ozone flux to foliage of mature ponderosa and Jeffrey pines in the San

Bernardino Mountains showed that ozone injury increased substantially in the year

following relief from a prolonged period of drought. The distribution of individual

trees in OII classes showed a shift from 1991 to 1992 to greater numbers of trees in

classes representing higher injury levels. Seasonal patterns of conductance followed

trends of diminishing soil water as the summer progressed, and highest rates of

ozone flux occured early in the growing season, with relatively little ozone flux

after July. However, the high level of variability in annual precipitation influenced

the size and persistence of annual needle whorls, particularly during and after

recovery from a prolonged drought. These internal foliar dynamics interacted with

external environmental parameters, such as winter precipitation and soil water

depletion, to influence rates of stomatal conductance and ozone flux to pine foliage,

which influenced development of ozone injury on these trees.

Acknowledgments

We thank Anthony Gomez and David Jones for technical support, and Susan

Schilling and Delparde Guthrey for assistance with data analyses. We thank Laurie

Dunn for technical editing of this manuscript. We also thank Brent Takemoto for

his support of this project. This research was sponsored by the California Air

Resources Board, Contract No. A032-180.

References

Coyne, Patrick I.; Bingham, Gail E. 1982. Variation in photosynthesis and stomatal conductance in an ozone-stressed ponde- Variation in photosynthesis and stomatal conductance in an ozone-stressed ponderosa pine stand: light response. rosa pine stand: light response. Forest Scienc rosa pine stand: light response. e 28: 257-273. e 28: 257-273.

Helms, John A. 1970. Summer net photosynthesis of ponderosa pine in its natural environment. . Summer net photosynthesis of ponderosa pine in its natural environment. Photosynthetica 4: 243-253. . Summer net photosynthesis of ponderosa pine in its natural environment. 

Helms, John A. 1971. Environmental control of net photosynthesis in naturally growing . Environmental control of net photosynthesis in naturally growing Pinus ponderosa Laws. Ecol- Laws. 

ogy 53: 92-101.

Miller, Paul R.; Chow, Judy; Watson, John; Bytnerowicz, Andrzej; Fenn, Mark; Poth, Mark; Taylor, George. 1996a Mark; . Assessment

of acidic deposition and ozone effects on conifer forests in the San Bernardino Mountains. Final Report, Contract No. of acidic deposition and ozone effects on conifer forests in the San Bernardino Mountains. Final Report, Contract No.

A032-180. A032-180. California Air A032-180. Resources Board, Sacramento, CA .

Miller, Paul R.; Stolte, Kenneth W.; Duriscoe, Daniel; Pronos, John, technical coordinators. 1996b. Evaluating ozone air pollu- . Evaluating ozone air pollution effects on pines in the western United States. tion effects on pines in the western United States. Gen. Tech. Rep. PSW-GTR-155. Albany, CA: USDA Forest Service; 79 p. tion effects on pines in the western United States. 

NOAA. 1995. Climatological Atlas for California. . Climatological Atlas for California. Washington, D.C.: National Oceanographic and Atmospheric Administration. . Climatological Atlas for California. 

Patterson, Mark T.; Rundel, Philip W. 1989. Seasonal physiological responses of ozone-stressed Jeffrey pine in Sequoia Na- . Seasonal physiological responses of ozone-stressed Jeffrey pine in Sequoia National Park, California. tional Park, California. In, Allen S., eds. Effects of air pollution on western forests. Pittsburgh, PA: Air and Waste Management Assoc.; 279-292.

Runeckles, Victor C. 1992. Uptake of ozone by vegetation. . Uptake of ozone by vegetation. In: Lefohn, Allen S., ed. Surface level ozone exposures and their . Uptake of ozone by vegetation. 

effects on vegetation. Chelsea, MI: Lewis Publ. Inc.; 157-188.

Temple, Patrick J.; Riechers, George H.; Miller, Paul R. 1992. Foliar injury responses of ponderosa pine seedlings to ozone, wet

and dry acidic deposition, and drought. and dry acidic deposition, and drought. Environmental and Experimental Botany 32: 101-113. and dry acidic deposition, and drought. 

Session I1I Seasonal Influences on Ozone Uptake and Foliar Injury to Ponderosa and Jeffrey Pines Temple, Miller
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 229

Nancy E. Grulke,2 Paul R. Miller,2 Theodor D. Leininger3

Abstract

Five species of western conifers (Pinus ponderosa, Abies concolor, Pseudotsuga menziesii, Abies lasiocarpa, and

Picea engelmannii) were exposed, in two standard open-top exposure chambers per treatment, to charcoal-filtered

air and a simulated diurnal ozone exposure profile (120 d sum of 136 ppm-h) to test their relative sensitivity. CO2

exchange rate (CER), stomatal conductance (gs), and total leaf nitrogen of current year foliage were measured at 1-

month intervals. P. ponderosa was the most adversely affected by ozone, based on reductions in mid- and late season

CER under saturating light (1.18 vs. 2.17 μmol CO2

 m-2 s-1), lower light compensation point (43 vs. 33 μmol quanta

m-2 s-1), and lower CER at a given gs value. P. menziesii showed reduced CER and higher gs by the end of the experimental

ozone exposure. A. concolor and P. engelmannii showed no reduction in CO2

 uptake and noclear effects of ozone

exposure on gs. Adverse ozone effects on biomass or growth in the year of exposure were seen for A. lasiocarpa and P.

ponderosa, whereas A. concolor, P. engelmanii and P. menziesii showed adverse effects only in the year after exposure.

Few consistent patterns in which plant component was affected were found, either in the year of exposure or the year

after exposure. An analysis of the number of chambers required or the difference between treatment means necessary

to obtain statistical significance with the population variance measure was an important result.

Introduction

Considerable variations in the response of trees to air pollutants may be caused by

differences in seed source, nutritional status of plants and other stressors, plant

age, and plant phenological stage (Bytnerowicz and Grulke 1992). Many

experimental exposure studies have analyzed the effects of pollutants on gas

exchange at one phenological period: near the end of the growing season when

cumulative experimental exposure to the pollutant is at a maximum. However,

because gas exchange rates may be at a minimum, treatment effects may be difficult

to distinguish statistically.

Plant response to ozone may not necessarily be cumulative, but relatively few

studies have studied this by examining seasonal responses during ozone exposures

(Byres and others 1992, Grulke and others 1989, Hanson and others 1994, Samuelson

and others 1996, Weber and others 1993). As long as concentrations are sufficiently

low, ozone may have little effect on gas exchange. However, high ozone uptake rates

have been found to reduce stomatal conductance (gs) (Weber and others 1993).

This study determined the relative susceptibility of five species of western

conifers — Abies concolor [Gord. & Glend.] Lindl., Abies lasiocarpa [Hook.] Nutt.,

Picea engelmannii Parry, Pinus ponderosa Dougl. ex. Laws, and Pseuotsuga

menziessii Franco — to ozone exposure by analyzing gas exchange characteristics (CO2

 exchange rate, stomatal conductance, and dark respiration), growth,

and biomass allocation.

Materials and Methods

Plant Culture

Two-year-old, bare-root A. concolor seedlings were obtained from the California

Department of Forestry nursery in Ben Lomond (seed zone 531.50). One-year-old,

container-grown P. ponderosa seedlings were obtained from the same nursery (seed

zone 522.20, lot 3245). Two-year-old A. lasiocarpa seedlings in containers were

obtained from Plants of the Wild, Tekoa, Washington (seed from south of Lewiston,

Idaho, 1,372 m). Two-year-old, bare root stock of P. engelmannii (seed source Warm

Lake, Idaho) was obtained from Lucky Peak Nursery. One-year-old container-grown

seedlings of P. menziesii were obtained from Rex Timber, Inc., Cottage Grove, Oregon

(seed zone 471, 620 m).

Effect of Ozone Exposure on Seasonal Gas Exchange of Five

Western Conifers1

1 An abbreviated version of this paper was

presented at the International Symposium on Air Pollution and Climate

Change Effects on Forest Ecosystems,

February 5-9, 1996, Riverside California.

2 Research Plant Physiologist and Research Plant Pathologist, respectively,

Pacific Southwest Research Station,

USDA Forest Service, 4955 Canyon

Crest Drive, Riverside, CA 92507.

3 Research Plant Pathologist, Southern

Research Station, USDA Forest Service,

Box 227, Stoneville, MS 38776.
230 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Seedlings (6.4 cm dia., 25.4 cm tall) were potted in a mix consisting of equal

parts Promix B1 and perlite. Before fumigation, plants were kept in an activated

charcoal-filtered, evaporative-cooled greenhouse adjacent to the fumigation site.

The greenhouse and fumigation site were installed at the Kenworthy Ranger Station (33° 37’ N, 116° 37’ W), in the San Jacinto Mountains, 22 km SE of Idyllwild,

and 80 km W of Palm Springs, California. The elevation of the site was 1,387 m. A

drip irrigation system was used to water and fertilize the plants (North Carolina

State University fertilizer mix). Plants were watered two times weekly to pot capacity. Nutrients were applied every 1-2 weeks through the irrigation system from

March through October 11, 1988.

Ozone Exposure

Seedlings were exposed to either charcoal-filtered air (CF), or charcoal-filtered air

augmented with ozone (+O3 ) under ambient conditions in open-top exposure

chambers (Heagle and others 1979). Chambers operated from June 4 (Julian day

155) through October 11, 1988. For the elevated ozone treatment, a 30 d diurnal

profile was developed from average air quality characteristics during the summer

months in the midwest (1.8 times ozone exposure level; Hogsett and others 1989;

Lefohn and others 1986a, 1986b). The 30 d base profile was repeated about four

times in the 124 d exposure period (fig. 1). The ozone exposure in each treatment

was estimated from the following equations:

CF: y = 0.278x - 1.44 (1)

O3

: y = 1.118x - 1.66 (2)

in which y was ozone exposure in ppm-h and x was number of days of exposure.

Ozone was generated, distributed, and sampled within each chamber by

using a computer-controlled program with a data acquisition system (Hewlett

Packard).4 Oxygen (O2 ) was supplied to an ozone generator (Griffin Technics, Model

GTC-0.5b), and distributed to chambers with mass flow controllers (Datametrics, Type

825). Ozone was sampled from the center of each chamber by using a sequential

sampler (Scanivalve, custom-made electronics), Blower boxes provided a chamber

exchange rate of 2 min-1 and ozone concentrations were determined by ultraviolet

photometry (Dasibi, Model 1003AH, calibrated bi-weekly on site). Fiberglass dust

filters and activated charcoal-filters were in place for all chambers. Concentrations of ozone in chambers were recorded hourly as were ambient ozone, chamber

air temperature, soil temperature at 5 cm, and total global radiation (Licor 200SZ).

Gas Exchange

Before experimentation, a typical subset of plants for physiological measurement was placed in each chamber: all had one terminal bud, appeared to be in the

same phenological stage, and were visually similar in height, numbers of side

branches, and foliage color. Bud flush had occurred before placement in chambers.

Three seedlings of each species per chamber and two chambers each for charcoal filtered air and elevated ozone treatments were available for physiological

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger

Figure 1 — Seasonal course of

daily mean, maxima, and minima

ozone concentration in the

elevated ozone chambers. The

lowest ozone concentrations

occurred in the morning from

0700 to 0900, and the highest

ozone concentrations occurred

between 1500 and 1700 hr.

4 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 231

measurements. Means of each plant were obtained from two to four sequential

runs (three to four observations averaged per run). Gas exchange from the three

seedlings of each species in one chamber were averaged together to obtain the chamber mean and standard error. The analysis of variance (ANOVA) calculations were

strictly applied to the chamber means (one degree of freedom within a treatment).

The conifers were exposed to charcoal-filtered air and elevated ozone in opentop chambers for one growing season. The conifers included Abies concolor [Gord.

& Glend.] Lindl., Abies lasiocarpa [Hook.] Nutt., Picea engelmannii Parry, Pinus ponderosa Dougl. Ex Laws, and Pseudotsuga menziesii Franco. Gas exchange of current

year foliage was measured four times during the exposure period from early June

through late September. Dark respiration rate and light response of A. concolor, P.

ponderosa, and P. menziesii were measured once at the end of the experimental exposure and the growing season. Biomass was measured twice on a separate set of

seedlings: at the end of the experiment in the fall, and in the fall after l year of

“recovery.” The seasonal pattern of gas exchange and their relative susceptibility

to ozone exposure of the five conifers was measured. For each species, gas exchange

of current year foliage from the same three seedlings from each chamber was measured at the beginning of the exposure period in early June and subsequently at 1-

month intervals through late September. CO2

 exchange rate (CER) and stomatal

conductance (gs) were measured with a Licor 6200 photosynthetic system (Lincoln, Nebraska). A 0.25 L cuvette was used for all species except P. ponderosa.

For this species, a custom cuvette (12 x 15 x 15 cm) was constructed from Acrylite

lined with Teflon film so that shoot architecture was not disturbed during measurement. Leak rates of the custom cuvette were low and comparable to the

purchased cuvette.

Gas exchange measurements were made outside fumigation chambers between

0930 and 1530 h. Plants of the same species were out of chambers approximately 1

h before measurement (Hinckley and others 1990) and were measured at the same

time of day throughout the growing season. During all measurement sessions, plants

from CF chambers were alternated with ozone chambers to nullify possible diurnal effects. In addition to biweekly watering, all plants were watered to capacity

0.5 d before measurements.

Gas exchange was measured within 1 minute. Gas exchange measurements for

the ozone experiment were taken at ambient light within a photosynthetic photon

flux density (PPFD) range of 750-1525 ␣μmol m-2 s-1, leaf temperature of 29-37 °C,

and vapor pressure deficits (VPD) of 2.75-4.50 kPa. CO2 exchange rate and stomatal

conductance appeared to have a flat response across this range of physical factors,

other than light, for which lower levels could have limited gas exchange up to 10

percent. Dark respiration rate of A. concolor, P. ponderosa, and P. menziesii was

measured near the end of the season-long experiment, approximately 0.5 h after

sunset. Measurements were taken within an air temperature range of 16-20 °C.

CER was calculated on the basis of needle surface area. Surface area was calculated from simple geometric models of leaf surfaces with consideration for surfaces with stomata. Measures of needle width (in the middle) or fascicle diameter

(just above the sheath) and length were made with a digital micrometer.

Light response was determined near the end of the summer ozone exposure

period. Four frames consisting of a light source, heat/light separation mirror, and

neutral density filters were used to control light intensity (Grulke and others 1989).

The same plants measured for seasonal response were also measured in this analysis.

Plants were allowed to equilibrate for 0.5 h at each light level before gas exchange

was measured. Gas exchange measurements were made between cuvette

temperatures of 22-26  ̊C. For light response determinations of P. ponderosa, three

fascicles were enclosed in a 0.25 L cuvette (spread to reduce self- shading). A simple

leaf model was used to define the relationship: (CER = Pmax (1-e-¢I/Pmax), in which ¢ is

apparent quantum efficiency and I is PPFD; Thornley 1976). The CO2 light

compensation point and CER with saturating light were calculated from the fitted

regression equations for each plant.

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
232 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Leaf Nitrogen

Leaf tissue was sampled for nitrogen content just after the monthly gas exchange

measure by using a standard micro-Kjeldahl digest technique on dried tissue

(65 °C, 24 h), ground to pass a 40 mesh. Digests were run on an auto-analyzer

(Technicon) with standards, blanks, replicates, and duplicate runs every 20 samples.

Samples with > 2.5 percent error ([S.D./x] x 100 percent) were re-analyzed on a

Carlo-Erba NA1500 Series 2 analyzer.

Biomass and Plant Size

Plant height (mm) and stem diameter (0.01 mm) was measured before experimental exposure for six additional seedlings per chamber. At the end of the experiment,

plant height, stem diameter, and biomass allocation to roots, stem, older needles,

and new (current year) needles were determined for the same six seedlings. The

change in seedling height and stem diameter was relativized by using the initial

measures taken before the experiment ( ∆ ht/htix 100 percent). For yet another set

of six seedlings per chamber, plant size and biomass allocation was measured after 1 full year of growth in a charcoal-filtered air glass house after experimentation.

Results

CO2 Exchange Rate

Seasonal trends in CER were different for each species: maximum CER occurred at

the first sampling date for P. ponderosa, at the second sampling date for Abies

lasiocarpa, and at the last sampling date for Abies concolor, Picea engelmannii, and

Pseudotsuga menziesii (fig. 2). The maximum observed CER varied nearly twofold

for current year foliage between the species (A. concolor, 1.68+0.42; A. lasiocarpa,

1.42+0.08; P. engelmannii, 2.40+0.11; P. ponderosa, 1.52+0.11; P. menziesii, 2.50+0.22

␣μmol CO2 m-2 s-1). Particularly low values of CER were obtained on the second

sampling date for P. menziesii because of high vapor pressure deficits.

Only P. ponderosa showed significant effects of elevated ozone, which were

measured at the second sampling date (fig. 2). Despite the few chambers used, several trends are apparent. Although initial CER in the elevated ozone treatment was

sometimes greater than that of the charcoal filtered air treatment (e.g., A. concolor, P.

engelmannii), the end of season mean CER was lower in the elevated ozone treatment for all species. After the exposure regime began, the largest differences in

CER between the elevated ozone and charcoal filtered air treatments occurred early

season for A. concolor and A. lasiocarpa, and midseason for P. ponderosa. We found

no differences attributable to treatment for both P. engelmannii and P. menziesii.

Because of high within-chamber variability due to an uneven number of

sensitive and tolerant individuals in each chamber, and because the seasonal gas

exchange patterns were different for the plants within each chamber, significant

differences were not found between treatments (except for P. ponderosa). For example,

one A. concolor seedling had a positive parabolic pattern, one had a constantly

declining seasonal gas exchange pattern, while the remaining four seedlings within

a treatment had a negative parabolic pattern of seasonal gas exchange. The net

effect of these differing seasonal patterns of gas exchange is a relatively flat seasonal

response with high standard errors (see A. concolor, fig. 2). Neither condition

(sensitivity or asynchronous seasonal patterns of gas exchange) could have been

known before the experimental exposure.

Measured at the end of the experiment, dark respiration was higher in seedlings exposed to elevated ozone than charcoal filtered air treatments for all species,

but differences were not statistically significant (table 1).

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 233

Figure 2 — Seasonal trend in

CO2

 exchange rate (CER) for

charcoal-filtered (CF) air (●) and

elevated ozone treatments (O)

for five species of western

conifers. Symbols and error bars

represent averaged means for

the two chambers + 1 S.E. Values

in parentheses are probabilities

that the two means are

significantly different in a oneway ANOVA. The relationship

between CO2

 exchange rate

(CER) and stomatal conductance

(gs) for the five species for

month 2-4 of treatment is also

given. Symbols represent average

values for 2-4 runs per plant; in

all cases, standard error was

contained within the symbol.

Values in lower right-hand

corner are probabilities that

the treatment (Tr) and stomatal

conductance (gs) were significantly different. Slopes of the

lines were not significantly

different between treatments for

any species.

Leaf Nitrogen

No seasonal trends in needle nitrogen content were apparent for any of the species.

Needle nitrogen differed significantly between the charcoal filtered air and elevated

ozone treatments averaged over the duration of the experiment for A. concolor, but

not for the other species (table 2). CER based on leaf nitrogen content did not improve

statistical separation of response to experimental treatment (fig. 1).

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
234 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

1 Mean and error represent averaged means for the two chambers and within-chamber standard error in each treatment.

2 Probability for significant differences between values (one-way ANOVA).

Table 1 — Dark respiration ( able 1 μmol CO2 m-2 s-1) at the end of the experiment for charcoal-filtered (CF air) and elevated

ozone (+ O3) treatment for the five conifer species studied.

Species CF air 1 + O3

1 p 2

Abies concolor 0.21+00 0.22+0.02 (0.52)

Abies lasiocarpa 0.16+0.01 0.17+0.02 (0.72)

Picea engelmannii 0.15+0.01 0.16+0.01 (0.14)

Pinus ponderosa 0.20+0.04 0.24+0.03 (0.50)

Pseudotsuga menziesii 0.30+0.03 0.34+0.00 (0.24)

Table 2 Table 2 — Leaf nitrogen (percent) averaged for the four sampling points throughout the duration of the experiment for Table 2 

charcoal-filtered (CF air) and elevated ozone (+ O3 ) treatment.

Species CF air 1 + O3

1 p 2

Abies concolor 1.38+0.08 0.98+0.01 (±.01)

Abies lasiocarpa 1.27+0.06 1.14+0.04 (0.11)

Picea engelmannii 1.84+0.10 1.90+0.11 (0.72)

Pinus ponderosa 1.20+0.06 1.15+0.05 (0.51)

Pseudotsuga menziesii 1.11+0.08 1.13+0.12 (0.89)

1 Mean and standard error.

2 Probability for significant differences between values (one-way ANOVA).

Stomatal Conductance

Seasonal patterns in stomatal conductance (gs) largely followed CER (fig. 1).

However, after a month of treatment exposure, gs was generally greater in elevated

ozone treatment for A. concolor, A. lasiocarpa, and P. ponderosa than in the charcoalfiltered air treatment. CER was plotted as a function of gs to further test for the

effects of elevated ozone exposure (fig. 1) for months 2 to 4 of treatment exposure.

The slopes of the regressions were not significantly different for four of the species

(Draper and Smith 1981). For P. ponderosa, CER was significantly reduced in the

elevated ozone treatment relative to the charcoal filtered air treatment at the same

gs, suggesting reduced water use efficiency. The differences in CER were greater

than differences in dark respiration (table 1).

Light Response

Light compensation points were not significantly different between treatments for

A. concolor, P. ponderosa, or P. menziesii (table 3) using a one-way ANOVA. CER at

light saturation (1,600 ␣μmol m-2s-1) was significantly different between treatments

for P. ponderosa, but not for A. concolor or P. menziesii (table 3). The apparent quantum

efficiency (the slope of the regression line between 0 and 100 ␣μmol quanta m-2 s-1)

for P. ponderosa seedlings growth in charcoal-filtered air and elevated ozone was

statistically significant (p<0.01) (Draper and Smith 1981). The difference in light

response between treatments in P. ponderosa was much greater than the difference

in dark respiration observed (table 1).

Biomass Allocation

Biomass allocation to roots, stems, old needles (produced before this experiment),

and new needles (current year) was analyzed for seedlings immediately after the

growing season, and 1 full year after experimental exposure (fig. 3). Some species

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 235

showed adverse ozone effects on biomass or growth in the year of exposure (A.

lasiocarpa, P. ponderosa), some showed only adverse effects in the year after exposure

(A. concolor, P. engelmanii, P. menziesii). There were few consistent patterns in which

plant component was affected, either in the year of exposure or the year after

exposure. For all species, the percent biomass allocation to older needles was lower

for seedlings grown in elevated ozone chambers relative to charcoal-filtered air

chambers, but the differences were not significant (table 4). Total biomass was lower

for seedlings in elevated ozone than charcoal-filtered chambers immediately after

exposure for all species except P. ponderosa, where seedlings were larger (+10.6

percent in total biomass, and 10.1 percent in dia.) at the end of the experimental

exposure. Plants in the elevated ozone treatments were also larger in May before

experimental exposure. Two species also had lower biomass allocation to new

needles (A. lasiocarpa (p=0.01), P. menziesii, p =0.57), but A. concolor, P. engelmanii,

and P. ponderosa had greater allocation to new needles.

Discussion

In this study testing the sensitivity of five western conifers to ozone, P. ponderosa

appeared to be the most sensitive to ozone on the basis of reductions in CER (in

midseason, CER versus gs responses, and in light saturation experiments at the

end of the growing season). Among all of the species, P. ponderosa had the greatest

gs early in the season when young leaf tissues were most likely to be damaged by

oxidants. In general, species of pine appear to be more sensitive to ozone exposure

than any other conifer (Bytnerowicz and Grulke 1992).

Although A. lasiocarpa had a seasonal pattern of gs similar to that of P. ponderosa, few symptoms of sensitivity to ozone developed. In this study, as well as in

their natural habitat, CER and gs of A. concolor (Conard and Radosevich 1981) and

P. engelmannii (Carter and others 1988, DeLucia and Smith 1987) increased

gradually through the growing season. Comparisons of instantaneous gas exchange measures showed that both species appear to be relatively resistant to

ozone exposure (fig. 1).

Instantaneous measures of gas exchange as well as biomass of P. menziesii did

not statistically demonstrate evidence of damage by exposure to ozone. However,

1 Mean and error represent averaged means for the two chambers and within-chamber standard error in each treatment.

2 Probability for significant differences between values (one-way ANOVA).

3 μmol quanta m-2 s-1.

4 μmol CO2 m-2 s-1.

Table 3 Table 3 — Light compensation point and CO Table 3 2

 exchange rate at the end of the exposure for charcoal-filtered air (CF air)

and elevated ozone (+ O3 ) treatments.

Species CF air 1 + O3

1 p 2

Light compensation

point 3

Abies concolor 28 + 5 28 + 4 (0.98)

Pinus ponderosa 47 + 5 49 + 6 (0.86)

Pseudotsuga menziesii 20 + 4 19 + 4 (0.89)

CO2 exchange rate at

light saturation 4

Abies concolor 3.17 + 0.34 3.21 + 0.36 (0.93)

Pinus ponderosa 2.67 + 0.19 1.21 + 0.24 (0.001)

Pseudotsuga menziesii 4.84 + 0.51 5.45 + 0.39 (0.37)

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
236 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

all species showed a trend of increased instantaneous measures of dark respiration, and subsequent decreased biomass of older needles (increased needle turnover rates) immediately after the experiment. Most species (except P. menziesii) showed

a trend of decreased biomass of new needles 1 full year after experimental exposure.

In another study of P. menziesii exposed to elevated ozone in Washington, measures of

instantaneous gas exchange did not reflect significantly reduced stem diameter and

root to shoot biomass at the end of a 2-year study (Hinckley and others 1990).

Instantaneous values are appropriate to understand what plant processes may

be sensitive to oxidants, but measuring long-term biomass changes may be a better

measure to detect damage. Modeling efforts that incorporate the small differences

in gas exchange and subsequent carbon allocation within the plant year after year

may demonstrate the consequences of subtle changes in carbon balance more clearly.

Few of the differences observed between the charcoal-filtered air and the elevated ozone treatments were significant. The lack of statistical significance was

primarily because of high within-chamber variance. Even if chamber effects were

assumed to be null and each plant were treated as a true replicate, there were few

additional statistically significant differences between treatment means. In many

pollutant exposure studies, plants are tested after the maximum cumulative exposure period, often at the end of a growing season, when gas exchange rates are

declining and the ability to statistically distinguish between means is the least powerful. In this study, the statistically significant differences between the charcoal filtered air and elevated ozone treatments occurred in the second month of exposure

for P. ponderosa (fig.1). Cumulative exposure to ozone was not necessarily correlated to peak deleterious effects on gas exchange in other studies (Byres and others

1992, Weber and others 1993). In this study, high variability in plant response within

a chamber and too few chambers significantly hindered the evaluation of the effects of exposure to elevated ozone.

To better clarify the lack of statistical significance, and assuming the same

within- chamber standard error, the difference between treatment means required for significance at the p = 0.05 level, or the number of chambers of plant

gas exchange that would have been required for the means to have significance was calculated (table 5). Untenably large numbers of chambers would

Figure 3 — Change of biomass

allocation by percent for seedling

growth in charcoal-filtered air

(first column) compared to those

in elevated ozone chambers

(second column). Two sets of

seedlings were harvested: (a)

immediately after the growing

season of experimental exposure

and (b) after 1 full year of growth

in a glass house supplied with

charcoal-filtered air. Bar lengths

represent the means of the two

chambers and + 1 S.E.

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger Biomass belowground

80

60

40

20

0

20

40

60

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

–

October, 1989 harvest

Abies

concolor

Abies

lasiocarpa

Picea

engelmanii

Pinus

ponderosa

Pseudotsuga

mensiesii

Biomass

aboveground

_

_

_

_

_

_

_

_

_

_

_

_

_

_

_

80

60

40

20

0

20

40

60

October, 1988 harvest pct

pct

–

–

–

–

–

–

Biomass

belowground

Biomass

aboveground

Roots Stems Old needles New needles

–

–

–

–

–
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 237

have been required to demonstrate statistical significance between treatments

for all species other than P. ponderosa.

One source of high population variance may be naturally occurring genotypes

that are “sensitive” or “resistant” to pollutants (Ernst and others 1985, Taylor 1978).

Evidence for differences in population sensitivity to ozone was found in different

individuals of P. jeffreyii (Patterson and Rundel 1989), different seed sources of

1 Parenthetical values are the probability that the change is significant between treatments (chamber means of six seedlings were the replicates, df=1). Bold indicates values

significant at p<0.05.

2 Greater allocation to a tissue type in elevated ozone versus charcoal filtered chambers.

Table 4 — Magnitude and direction of change of biomass allocation by percent for seedlings grown in charcoal filtered air compa Table 4 red to those in elevated ozone chambers.

Two sets of seedlings were harvested: (A) immediately after the growing season of experimental exposure and (B) after 1 full year of growth in a glasshouse supplied with

charcoal-filtered air.

A. 1988 October Harvest A. 1988 October Harvest

Abies Abies Picea Pinus Pseudotsuga

concolor lasiocarpa Engelmannii Engelmannii ponderosa menziesii menziesii

biomass 1 - 1.80 (0.91) - 15.6 (0.12) - 25.0 (0.35) + 10.1 (0.51) - 14.0 (0.48)

height + 3.3 (0.85) + 23.4 (0.78) + 23.8 (0.71) + 8.9 (0.76) - 18.6 (0.31)

diameter + 21.6 (0.25) + 8.3 (0.79) - 31.7 (0.72) 2

 + 10.1 (0.01) - 18.2 (0.27) 10.1

root + 5.3 (0.63) + 5.7 (0.47) - 13.9 (0.32) + 7.1 (0.27) - 17.3 (0.39)

stem - 9.6 (0.17) - 8.7 (0.38) + 14.1 (0.30) + 1.3 (0.89) - 14.2 (0.41)

old needles - 7.1 (0.51) - 17.3 (0.35) - 11.1 (0.67) - 50.2 (0.23) - 17.8 (0.46)

new needles + 17.0 (0.62) 2 - 15.5 (0.01) + 27.0 (0.25) + 19.5 (0.28) - 8.8 (0.57) - 15.5

B. 1989 October Harvest B. 1989 October Harvest

Abies Abies Picea Pinus Pseudotsuga

concolor lasiocarpa Engelmannii Engelmannii ponderosa menziesii menziesii

biomass 2 + 29.5 (0.01) 0.0 (0.99) - 10.3 (0.61) + 50.7 (0.10) + 10.0 (0.30) 29.5

height + 11.6 (0.10) + 12.0 (0.36) + 8.2 (0.56) + 26.9 (0.16) + 11.1 (0.23)

diameter 2

 + 14.9 (0.03) + 0.8 (0.91) - 3.2 (0.78) + 9.8 (0.09) + 19.3 (0.23) 14.9

root - 3.5 (0.77) - 13.8 (0.10) 2

 + 8.6 (0.23) + 7.2 (0.63) + 0.5 (0.96) 8.6

stem + 14.4 (0.28) + 11.4 (0.15) - 4.6 (0.37) - 7.7 (0.32) + 7.0 (0.52)

old needles - 21.9 (0.27) + 18.9 (0.29) - 14.3 (0.44) + 3.3 (0.89) 2 - 22.8 (0.02) - 22.8

new needles 2

 + 23.4 (0.40) - 59.8 (0.38) - 5.3 (0.71) - 26.3 (0.86) + 10.8 (0.81)

1 Parenthetical values are probability of significant differences between gas exchange values from figure 1.

Table 5 Table 5 — Evaluation of statistical significance for CO Table 5 2

 exchange from figure 1. The differences between treatment means

or the number of chambers that would have been required for the means to have been significantly different at the p=0.05

level have been calculated, assuming the same population variance.

Species p1 Date of Date of Difference in No. of chambers

comparison mean required required

 CO2

 exchange

Abies concolor (0.77) 9/22 1.8x 500

Abies lasiocarpa (0.99) 9/22 5.2x 330

Picea engelmannii (0.70) 9/19 4.0x 190

Pinus ponderosa (0.27) 8/10 1.1x 2

Pseudotsuga menziesii (0.75) 9/21 3.5x 140

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
238 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

P. strobus (Barnes 1972), and in different families of P. ponderosa (Beyers and others

1992). In addition to genetic differences, plant water status, leaf tissue nutrient content, and different seasonal patterns of gas exchange or phenology may alter individual plant sensitivity to ozone and contribute to the lack of statistical power to

detect treatment effects or to resolve discrepancies between published studies.

Acknowledgments

We thank Paul Dunn (planning), Anthony Gomez (installing the fumigation facility,

planting), David Herring (computer programming, data acquisition and control

systems), Susan Schilling and David Randall (statistical design and analysis), and

David Larson (maintaining plants). Douglas Pumphrey (District Ranger, San Jacinto

District, San Bernardino National Forest) kindly permitted use of land and water at

the fumigation site. This research was supported in part by the Environmental

Research Laboratory, U.S. Environmental Protection Agency, Corvallis, as a

component of the National Acid Precipitation Assessment Program.

References

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3: 133-138.

Beyers, J.L.; Riechers, G.H.; Temple, P.J. 1992. Effects of long-term ozone exposure and drought on the photosynthetic capac- Effects of long-term ozone exposure and drought on the photosynthetic capacity of ponderosa pine (Pinus ponderosa Laws). Laws). New Phytologist 122: 81-90. Laws). 

Byres, D.P.; Johnson, J.D.; Dean, T.J. 1992. Seasonal response of slash pine (Pinus elliottii Pinus elliottii var. elliottii Engelm. elliottii Engelm.) photosynthe- )

sis to long-term exposure to ozone and acidic precipitation. New Phytologist 122: 91-96

Bytnerowicz, A.; Grulke, N.E. 1992. Physiological effects of air pollutants on western trees. Physiological effects of air pollutants on western trees. In: Olson, R.K.; Binkley, D., eds. Physiological effects of air pollutants on western trees. 

Effects of air pollution on western forests. New York: Springer-Verlag; 183-233.

Carter, G.A.; Smith, W.K.; Hadley, J.L. 1988. Stomatal conductance in three conifer species at different elevations during

summer in Wyoming. summer in Wyoming. Canadian Journal of Forest Research 18: 242-246. summer in Wyoming. 

Conard, S.G.; Radosevich, S.R. 1981. Photosynthesis, xylem pressure potential, and leaf conductance of three montane chap- Photosynthesis, xylem pressure potential, and leaf conductance of three montane chaparral species in California. arral species in California. Forest Science 27(4): 627- 639. arral species in California. 

DeLucia, E.H.; Smith, W.K. 1987. Air and soil temperature limitations on photosynthesis in Engelmann spruce during sum- Air and soil temperature limitations on photosynthesis in Engelmann spruce during summer. Canadian Journal of Forest Research 17: 527-533. mer. 

Draper, N.; Smith, H. 1981. Applied regression analysis. Applied regression analysis. New York: John Wiley and Sons, Inc.; 709 p. Applied regression analysis. 

Ernst, W.H.O.; Tonneijck, A.E.C.; Pasman, F.J.M. 1985. Ecotypic response of Silene cucubalus to air pollutants (SO2

, O3).

Journal of Plant Physiology 118: 439-450.

Grulke, N.E.; Miller, P.R.; Wilborn, R.D.; Hahn, S. 1989. Photosynthetic response of giant sequoia seedlings and rooted

branchlets of mature foliage to ozone fumigation. branchlets of mature foliage to ozone fumigation. In: Olson R.K.; Lefohn A.S., eds. Effects of air pollution on western branchlets of mature foliage to ozone fumigation. 

forests. Pittsburgh, PA: Air, Waste, and Water Management Association; 429-441.

Hanson, P.J.; Samuelson, L.J.; Wullschleger, S.D.; Tabberer, T.A.; Edwards, G.S. 1994. Seasonal patterns of light saturated

photosynthesis and conductance for mature tree and seedling Quercus rubra foliage: differential sensitivity to ozone

exposure. exposure. Tree Physiology 14: 1351-1366. exposure. 

Heagle, A.S.; Philbeck, R.B.; Rogers, H.H.; Letchworth, M.B. 1979. Dispensing and monitoring ozone in open-top chambers

for plant effects studies. for plant effects studies. Phytopathology 69: 15- 20. for plant effects studies. 

Hinckley, T.M.; Schaap, W.; Wang, D. 1990. Rapid recovery of photochemical efficiency of PSII in Douglas-fir following ozone

treatment. treatment. Ecological Society of America Bulletin 71(2): 189. treatment. 

Hogsett, W.E.; Tingey, D.T.; Hendricks, C.; Rossi, D. 1989. Sensitivity of western conifers to SO2

 and seasonal interactions of and seasonal interactions of

acid fog and ozone. acid fog and ozone. In: Olson R.K.; Lefohn A.S., eds. Effects of air pollution on western forests. Pittsburgh, PA: Air, Waste, acid fog and ozone. 

and Water Management Association; 469-492.

Lefohn, A.S.; Hogsett, W.E.; Tingey, D.T. 1986a. A method for developing ozone exposures that mimic ambient conditions in

agricultural areas. agricultural areas. Atmosphere and Environment 20: 361-366. agricultural areas. 

Lefohn, A.S.; Hogsett, W.E.; Tingey, D.T. 1986b. The development of sulfur dioxide and ozone exposure profiles that mimic

ambient conditions in the rural southeastern United States. ambient conditions in the rural southeastern United States. Atmosphere and Environment 21: 659-669. ambient conditions in the rural southeastern United States. 

Patterson, M.T.; Rundel, P.W. 1989. Seasonal physiological responses of ozone-stressed Jeffrey pine in Sequoia National Park, easonal physiological responses of ozone-stressed Jeffrey pine in Sequoia National Park,

California. California. In: Olson R.K.; Lefohn A.S., eds. Effects of air pollution on western forests. Pittsburgh, PA: Air, Waste, and California. 

Water Management Association; 419-427.

Samuelson, L.J.; Kelly, J..; Mays, P.A.; Edwards, G.S. 1996. Growth and nutrition of Growth and nutrition of Quercus rubra L. Seedlings and mature

trees after three seasons of ozone exposure. trees after three seasons of ozone exposure. Environmental Pollution 91: 317-323. trees after three seasons of ozone exposure.

Taylor, G.E., Jr. 1978. Genetic analysis of ecotypic differentiation within an annual plant species, Genetic analysis of ecotypic differentiation within an annual plant species,

Geranium carolinianum L., in response to sulfur dioxide. L., in response to sulfur dioxide. Botanical Gazette 139: 326-368. L., in response to sulfur dioxide. 

Thornley, J.H.M. 1976. Mathematical models in plant physiology. Mathematical models in plant physiology. London: Academic Press; 318 p. Mathematical models in plant physiology. 

Weber, J.A.; Clark, C.S.; Hogsett, W.E. 1993. Analysis of the relationships among O3

 uptake, conductance, and photosynthesis

in needles of in needles of Pinus ponderosa. Pinus ponderosa. Tree Physiology 13: 157-172. 

Session I1I Effect of Ozone Exposure on Seasonal Gas Exchange of Five Western Conifers Grulke, Miller, Leininger
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 239

Mark A. Poth and Mark E. Fenn2

Abstract

Current-year needles from mature ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) were sampled at four sites

across the air pollution gradient in the San Bernardino Mountains in southern California. The sites, in order of

decreasing air pollution exposure, included: Sky Forest (SF), Conference Center (CC), Camp Angelus (CA) and Heart

Bar (HB). Needle nutrients measured were: nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium

(Mg), manganese (Mn), zinc (Zn), iron (Fe) and aluminum (Al). Concentrations of N in foliage (12.5 g/kg) were

significantly higher at SF, the site most exposed to air pollution. This level is similar to concentrations for containergrown seedlings fertilized with nutrient solution. Mature tree needle concentrations of K, Mg , Fe and Al were higher at

sites more exposed to air pollution. Mature trees with two annual whorls of needles (more severely injured) had

significantly higher current year needle concentrations of P, K, Zn, and Fe at SF than trees with three annual whorls of

needles (less severely injured). However, significant differences were not found in foliar nutrient concentrations between

trees with different levels of needle retention (injury) at CC, which is also a high pollution site. At Barton Flats (BF),

a low to moderate air pollution site in the San Bernardino Mountains, N, K and P were measured in current-year

needles and litter fall. The resorption efficiencies (the percent reduction in nutrient concentration upon senescence of

leaves or needles) for mature trees at BF were: 45 percent for N, 60 percent for P and 80 percent for K. N resorption

proficiency (the concentration a nutrient is reduced to before needle abscission) was high with N reduced to 4.4 g-N/

kg (considerably less than the defined threshold of 7g-N/kg). However, P resorption proficiency was very poor with P

reduced to 0.59 g-P/kg (higher than the defined threshold of 0.4 g-P/kg). A complete understanding of the changes in

tree nutrition associated with air pollution will require more research.

Introduction

Long-term chronic exposure to ozone and other associated air pollutants has a

pronounced effect on growth and mortality of ponderosa pine (Pinus ponderosa

Dougl. ex. Laws.). This has been clearly documented (Miller and others 1963, Miller

and others 1989) over the last 25 years in the San Bernardino Mountains of southern

California. The San Bernardino and San Gabriel Mountains form the transverse

ranges, which run from west to east and form the northern boundary of the South

Coast Air Basin. The Basin includes Los Angeles and the surrounding communities.

Vegetation in the San Bernardino Mountains varies with elevation and aspect. Scrub

and chaparral species dominate the lower elevations (1,200-1,600 m) on the western

and southern ends of the range, while the pinyon-juniper type is found on the

desert-facing eastern and northern extremes of the San Bernardino Mountains. The

mixed conifer forest occurs at elevations of about 1,500 to 2,400 m. Ponderosa pine

is the dominant pine species of the mixed conifer forest, except on the eastern end

of the San Bernardino Mountains where ponderosa pine is replaced by the closelyrelated Jeffrey pine (Pinus jeffreyi Grev. & Balf.). Visible indications of ozone stress

in ponderosa pine in the mixed conifer zone include a combination of reduced

needle retention, thinning of the lower crown, chlorotic mottle of foliage, the

production of shorter needles, and mortality—typically as the result of an insect

attack (Miller 1983).

A major research effort in Germany has shown that forest decline in a spruce

forest was caused by nutrient imbalances from atmospheric deposition of sulfur

(S), and nitrogen (N). (Huettl and others 1990, Schulze 1989). Magnesium (Mg)

was retranslocated from old needles to new foliage until luxury levels were

reached in the current-year foliage, and Mg deficiency levels occurred in the

older and subsequently-chlorotic foliage. Although Mg levels were in excess

Mature Ponderosa Pine Nutrient Use and Allocation

Responses to Air Pollution1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, Riverside, California, February 5-9, 1996.

2 Soil Microbiologist and Plant Pathologist, respectively, Pacific Southwest Research Station, USDA Forest

Service, 4955 Canyon Crest Drive,

Riverside, CA 92507.
240 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn

in current-year foliage, N levels in new foliage became deficient, notwithstanding

the excessive deposition of N (Schulze 1989). Similarly, given the dramatic effects

of ozone and associated air pollutants in the San Bernardino Mountains on foliar

injury, growth, and abscission in ponderosa pine (Miller 1983), it is likely that nutrient relations of these ponderosa pine trees are also significantly affected.

Despite the importance of ponderosa pine in western North America, information on ponderosa pine nutrition is scant. Available data are restricted to studies of

N fertilization or the responses of seedlings or unique site situations, some of which

are thought to define the limits of species range. Rarely is any nutrient other than N

and P examined. This is a reflection of the pattern of use for the ponderosa pine

resource. Historically, ponderosa pine has been harvested primarily from old-growth

stands. Second-growth from natural regeneration can take 80 to 100 years for trees

to reach marketable size. Although silviculturalists have successfully developed

methods for ponderosa pine regeneration, specific knowledge of nutrient use and

requirements remains poor (Powers 1981), particularly compared to the available

information for more intensively managed and shorter rotation species like Douglas fir. A search of the National Agricultural Library database yields 1,086 references with ponderosa pine in the title or as a key word and 2,344 for Douglas fir.

Further examination of the ponderosa pine citations reveals the paucity of nutritional data.

In this study we measured and compared foliar nutrients at four sites along the

ozone gradient in the San Bernardino Mountains. We present data on foliar nutrient concentrations, and relate the concentrations to visible ozone symptoms (needle

retention) in ponderosa/Jeffrey pine. Our objectives were to determine if foliar

nutrient levels might be related to the severity of ozone stress incurred by mature

trees, and to help define the range of nutrient concentrations seen in mature ponderosa/Jeffrey pine. In addition, we have examined the influence of air pollution

on nutrient use by mature and container-grown ponderosa pine by comparing nutrient concentrations in needles and litter. The nutrient resorption efficiencies (the

percent reduction in nutrient concentration upon senescence of leaves or needles)

and resorption proficiencies (the concentration a nutrient is reduced to before needle

abscission) determined for ponderosa pine are also reported.

Materials and Methods

Needle samples were collected from four plots located along a west to east ozone

concentration gradient spanning 55 km in the San Bernardino Mountains. The plots

were originally established in 1972-73 as part of a large interdisciplinary study of

the effects of air pollution on the mixed conifer forest in the San Bernardino

Mountains (Miller 1983). We sampled trees in the plots, in order of decreasing

exposure, at Sky Forest (SF), Conference Center (CC), Camp Angelus (CA), and

Heart Bar (HB). Ozone concentrations in the permanent plots were measured during

July and August from 1974-78. The respective estimated 24-hour average ozone

concentrations (μg/m3) at the sites were 196 at SF and CC, 157 at CA, and 59 at HB

(Miller and others 1989). Ozone concentrations have decreased slightly during the

1980s, but concentrations within the greater Los Angeles basin remain among the

highest in the nation. Visible symptoms of ozone injury were still prominent at SF

and CC, and to a lesser degree at CA.

We selected dominant and codominant trees for needle sampling. A minimum

of 24 mature ponderosa pine trees with a dbh of at least 30 cm were sampled at

each of the four sites, except at HB where Jeffrey pine was sampled. Trees with

mistletoe, pest problems or obvious abnormalities were avoided. Current-year

foliage was collected from exposed branches (one branch per tree) in the middle

crown in late August of 1986. For each tree sampled we also recorded the number

of annual whorls of needles and the presence of needle chlorotic mottle characteristic

of ozone damage.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 241

Sampled needles were dried and ground for chemical analysis. N and P were

determined by standard colorimetric methods following Kjeldahl digestion. Calcium (Ca), magnesium (Mg), potassium (K), manganese (Mn), zinc (Zn), iron (Fe),

and aluminum (Al) were analyzed by atomic absorption spectrometry after perchloric acid digestion of tissues. All analyses were done in the laboratories of the Institute

of Soil Science and Forest Nutrition, Albert-Ludwigs University, in Freiburg, Germany.

In a separate study at Barton Flats in the San Bernardino Mountains, needles

were collected in August of 1993 from mature ponderosa pines. This was done in

late summer/early fall when new needles were fully expanded. More than 100

trees were sampled. Needles were divided into age classes, oven dried at 70 °C and

ground to pass through a 40-mesh screen. Needles from each tree were analyzed

for Ca, Mg, K, Mn, and Zn by atomic adsorption spectroscopy after perchloric -

nitric acid digestion. Phosphorus (P) was determined from the same digest (Glaubig

and Poth 1993). Coupled combustion - gas chromatographic analysis was used to

measure total N and carbon (C). Older age classes of needles were only analyzed

for total C and N.

Pine litter was collected during the fall from litter traps at Barton Flats. After

collection, combined samples (all fall collection dates) were oven dried at 70 °C

and ground to pass through a 40 mesh screen. Pine litter samples were also analyzed for C, N, P, Ca, Mg, K, Mn, and Zn. For mobile nutrients, nutrient efficiency

(the percent reduction in nutrient concentration upon senescence of leaves or

needles) was calculated.

Results and Discussion

Foliar concentrations of K and Mg were highest at the high pollution sites SF and

CC (fig. 1). Foliar N was significantly higher at SF than at any other site (fig. 2).

Nutrients did not appear to be severely limiting for mature ponderosa pine, as was

reported in a previous study (Huettl and others 1990, Powers 1991).

Critical levels of N and P in ponderosa pine foliage (Powers 1981) are reportedly 9 and 0.8 g/kg. Below these levels the N and P supply becomes deficient for

adequate growth. Foliar concentrations of K, Mg, and Ca in this study appear to be

within the range of values for ponderosa pine reported in the literature (Clayton

and Kennedy 1980). Levels of Fe and Al were highest in the two high-pollution

sites (SF and CC), and Mn levels were very high at CC compared to the other sites

(fig. 3). Zn was highest at HB, but showed no pattern of variation across the ozone

gradient. Micronutrient levels are within the normal ranges reported for ponderosa pine (Powers 1981).

Figure 1 — Macronutrient

concentrations for current-year

foliage of mature ponderosa pine

trees in the San Bernardino

Mountains. Letters associated

with bars indicate significant

differences in foliar concentrations between the plots

according to Tukey’s multiple

comparison test (p=0.05).

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn
242 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Ozone stress is the main inciting factor leading to premature needle abscission

of ponderosa pine in the high-pollution sites in the San Bernardino Mountains.

Other factors, such as drought stress, could also reduce needle retention. However,

the high-pollution sites receive higher levels of precipitation (Miller and others

1989), yet needle retention is much greater in the drier, less-polluted sites. Thus, the

number of annual whorls of needles retained, especially when accompanied with

symptoms of chlorotic mottle, is considered a reliable measure of the level of ozone

stress affecting ponderosa pine. In some cases there are statistically significant differences in nutrient concentrations between trees retaining 2 or 3 years of foliage at

the two high pollution sites. Trees with only two annual whorls (the current-year

and the previous year) at SF have higher needle concentrations of P, K, Zn, and Fe

than trees with three annual whorls (fig. 4, 5, and 6); presumably the higher concentrations in the current-year whorls are the result of ozone-induced senescence and

nutrient retranslocation.

Figure 2 — Nitrogen concentrations for current-year foliage

of mature ponderosa pine trees

in the San Bernardino Mountains.

Letters associated with bars

indicate significant differences in

foliar concen-trations between

the plots according to Tukey’s

multiple comparison test

(p=0.05).

Figure 3 — Micronutrient

concentrations for current-year

foliage of mature ponderosa pine

trees in the San Bernardino

Mountains. Letters associated

with bars indicate significant

differences in foliar concentrations between the plots

according to Tukey’s multiple

comparison test (p=0.05).

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 243

In a previous study at Strawberry Peak, a site 4 km west of SF, current-year

foliage of ponderosa pine trees retaining only one annual whorl contained significantly higher levels of P than foliage of trees retaining two annual whorls (Fenn

1991). Phosphorus and potassium are highly mobile and are readily remobilized in

plants. Zinc is considered to be intermediate in terms of plant mobility (Epstein

1972, Salisbury and Ross 1985). Zinc remobilization is closely related to leaf senescence (Marschner 1986). Iron generally shows low to intermediate plant mobility

(Epstein 1972, Salisbury and Ross 1985). The retranslocation of micronutrients is

markedly increased during foliar senescence (Marschner 1986).

We did not detect significantly higher nutrient concentrations in current-year

foliage of trees with either two or three annual whorls at CC. Although estimated

ozone concentrations at SF and CC were reported to be equivalent based on limited data (Fenn 1991), the severity of ozone damage to ponderosa pine was greater

at SF than at CC (Miller and others 1989). This difference may explain why higher

Figure 4 — Macronutrient (K,

Ca, and Mg) concentrations for

current-year foliage of mature

ponderosa pine trees in the San

Bernardino Mountains retaining

two or three years of foliage.

Letters associated with bars

indicate significant differences in

foliar concentrations between

trees at the same plot according

to Tukey’s multiple comparison

test (p=0.05).

Figure 5 — Micronutrient

concentrations for current-year

foliage of mature ponderosa pine

trees in the San Bernardino

Mountains retaining two or three

years of foliage. Letters

associated with bars indicate

significant differences in foliar

concentrations between trees at

the same plot according to

Tukey’s multiple comparison test

(p=0.05).

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn
244 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

levels of nutrients were not found in foliage of trees with fewer annual whorls at

CC, contrary to the results at SF in this study, and in an earlier study at Strawberry

Peak in the San Bernardino Mountains (Fenn 1991). More severe ozone damage at

SF may be a result of greater water availability, resulting in greater stomatal uptake

of ozone. Available soil water in the top 90 cm (19 and 13 percent) and annual

precipitation from 1973 to 1977 (109 and 90 cm) were higher at SF than at CC (Miller

and others 1989). Ambient temperature is not expected to vary greatly between CC

and SF based on their proximity and similar elevations (1,705 m at SF and 1,610 m

at CC; Miller and others 1989).

In a previous study in the San Bernardino Mountains, N and P retranslocation

in ponderosa pine occurred just before or during needle senescence and death (Fenn

1991). In the present study we sampled foliage in August, during the summer growing season. A considerable amount of nutrient retranslocation probably occurred

later in fall. However, where the ozone stress is sufficient to result in premature

needle loss, nutrient retranslocation may occur earlier in the season. The timing of

nutrient retranslocation may also differ for the various mobile elements. Had we

collected foliage later in fall, concentrations of other mobile elements may have

also increased in current-year foliage, especially in trees severely damaged by ozone.

N and Mg were not higher in new foliage of trees with only two annual whorls.

Because of the mobility of N and Mg in plants (Salisbury and Ross 1985), they

would also be expected to retranslocate to younger foliage before abscission of older

needles. Retranslocation of N at the high-pollution sites may be reduced because of

a fertilizer effect of high N deposition (Fenn and Bytnerowicz 1993). Reduced

retranslocation of N from senescing foliage in response to fertilization has been

observed in several tree species (Flanangan and Van Cleve 1983).

Increased nutrient uptake by ozone-impacted trees could also account for the

increase in nutrient concentrations in trees retaining only two annual whorls. This

seems unlikely, however, given that ozone has been shown to reduce root growth

and root carbohydrate content in ponderosa pine (Andersen and others 1991). The

greatly reduced foliar biomass of ozone-stressed ponderosa pine is likely to result

in reduced total photosynthetic rates in the long term; although younger foliage

has been shown to compensate, at least in the short term, for the loss of older

foliage in ozone-stressed ponderosa pine seedlings (Beyers and others 1992).

Assuming that total photosynthate production is reduced in mature ozonestressed ponderosa pine trees, it is doubtful that increased nutrient uptake occurs in ozone-stressed trees.

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn

Figure 6 — Nitrogen and

phosphorus concentrations for

current-year foliage of mature

ponderosa pine trees in the San

Bernardino Mountains retaining

two or three years of foliage.

Letters associated with bars

indicate significant differences in

foliar concentrations between

trees at the same plot according

to Tukey’s multiple comparison

test (p=0.05).
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 245

Which nutrients have been conserved, presumably by retranslocation, could

be influenced by the supply of nutrients in the soil. P, K, Zn, and Fe were all found

in higher concentrations in current-year foliage of trees at SF that retained only two

annual whorls of needles. No data are available for soil concentrations of Zn or Fe

in the four plots. P levels in soil are similar in the high- and moderate-ozone sites in

the San Bernardino Mountains, but are significantly lower in the eastern low-ozone

sites (Fenn 1991). In the original studies in the 18 permanent plots established along

the ozone gradient, the soils at SF, CC and HB contained 0.49, 0.57, and 0.52

milliequivalents of K per 100 g soil (Arkley and others 1977). Soil at HB contains

only 0.88 milliequivalents of Mg per 100 g, which is also reflected in the lower Mg

concentrations in foliage at HB. It is unlikely that the apparently slight differences

in soil P and K concentrations that occur at SF and CC account for why at SF, but

not at CC, new foliage of trees with two annual whorls had higher nutrient levels

than trees with three annual whorls. Apparently ozone stress (as indicated by needle

retention), and not soil deficiency, is the primary factor that causes enhanced nutrient retranslocation in the more severely ozone-impacted trees at SF.

It may be a general phenomenon in perennial plant species sensitive to ozone

that ozone exposure results in enhanced nutrient retranslocation to younger foliage, especially under nutrient-limiting conditions. The concentration of N and S in

current-year foliage of loblolly pine (Pinus taeda L.) was higher in seedlings exposed to ozone than in plants exposed to sub-ambient ozone levels (Tjoelker and

Luxmoore 1991). Phosphorus concentrations in foliage also increased (p=0.10) in

seedlings exposed to ozone; and K tended to increase, but differences were not

significant (Tjoelker and Luxmoore 1991). Leaves from cottonwood (Populus deltoides

J. Bartram ex Marsh.) saplings exposed to ozone abscised prematurely and had

higher concentrations of N than control plants (Findlay and Jones 1990).

Higher nutrient concentrations in current-year foliage of ponderosa pine with

prematurely-abscised needles is indicative of how ponderosa pine trees respond to

ozone stress. Before total senescence and abscission of older foliage, mobile nutrients are retranslocated to younger foliage, which then assume a larger physiological role. After two seasons of ozone exposure, ponderosa pine seedlings compensated for loss of older foliage by increasing net assimilation rates and N concentration of current-year foliage (Hom and Reichers 1991). Nutrient retranslocation to

current-year foliage, and enhanced productivity of new foliage appear to be key

adaptive mechanisms in ponderosa pine for overcoming the ozone-induced loss of

productivity from older foliage.

Another important consideration of plant nutrient relations and ozone exposure is the influence of plant nutrient status on sensitivity to ozone. The co-occurrence of chronic N deposition in many areas with elevated ozone concentrations

further emphasizes the importance of investigating the interactions and relationships between atmospheric deposition, soil-plant nutrient relations, site fertility,

and plant sensitivity to ozone. For instance, Tjoelker and Luxmoore (1991) found

that growth of current-year foliage of loblolly pine was more sensitive to elevated

ozone concentrations when seedlings were grown in soil with higher levels of N.

The upper limit for needle N content can be defined by growing trees in defined nutrient-rich conditions. Results from such a study with seedlings

(Bytnerowicz and others 1990) yielded needle N concentrations of 12.8 g/kg. The

lowest needle N concentrations that can support growth are those in ponderosa

pine stands that have developed on extremely poor soils. Isolated stands growing

on nutrient depleted soils formed on hydrothermally altered parent materials in

the Sierra Nevada of California had needle N concentrations of 8.1 g/kg (Schlesinger

and others 1989). Mature needles on the trees at the Barton Flats sites had N concentrations ranging form 6.25 to 9.92 g/kg depending on the needle age. This comparison seems to demonstrate little difference between the Barton Flats site and

remote sites in California, Nevada and Arizona (Klemmedson and others 1990,

Schlesinger and others 1989, Powers 1981). However, N deposition at Barton Flats

was above historic background levels.

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn
246 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

With initial N deposition we would expect greater N availability to trees (Aber

and others 1989, Schulze and others 1989, van Dijk and Roelofs 1988). This difference would parallel the situation at more fertile sites or sites that are fertilized. As

a result trees could conserve energy by translocating less N before needles are abscised. At sites with poor soil fertility 60 percent of needle N is typically translocated before needles are abscised (Schlesinger and others 1989). Fertile volcanic

soils with associated greater N availability result in less N translocation and

readsorption (48 percent) before needle abscission (Klemmedson and others 1990).

At Barton Flats only 45 percent of N is readsorped before needle abscission. This is

the lowest resorption efficiency reported in the literature for ponderosa pine. Resorption efficiency (the percent reduction in nutrient concentration upon senescence of leaves or needles) has recently been contrasted by Killingbeck (1996) with

resorption proficiency (the nutrient concentration achieved in senesced leaves or

needles). Plants that can reduce N concentrations in senesced leaves below 7 g/kg

are highly proficient. Ponderosa and Jeffrey pines at Barton Flats could reduce the

concentration of N in senesced needles to 4.4 g/kg and so were highly proficient in

N resorption.

P and K resorption efficiencies at Barton Flats by ponderosa pine were 60 percent and 70 percent effective, respectively. This level of P readsorption matches

that found for ponderosa pine growing on very infertile soils (Schlesinger and others 1989). The P resorption proficiency for these two sites differ considerably: 0.59

g/kg at Barton Flats compared to 0.42 g/kg at the infertile soil sites in the Great

Basin (Schlesinger and others 1989). Killingbeck (1996) has defined complete resorption proficiency in evergreen species as reducing the P concentration below 0.4

g/kg and incomplete resorption as a P final concentration greater than 0.5 g/kg.

Clearly, P resorption proficiency at Barton Flats is incomplete.

The poor correlation of needle nutrient content with indicators of air pollution

damage may be the result of the plants’ ability to compensate physiologically for

some air pollution damage and the variability of response between genotypes

(Wellburn 1994) The principal air pollutant causing visible injury and thought to

be responsible for growth changes is ozone (Peterson and others 1991, Temple and

Miller 1994). The influence of ozone on nutrition is certainly indirect and chronic,

whereas the response of a needle may be integrated over the much shorter time

span (the lifetime of the needle). Consequently the possibility of developing a simple

diagnostic needle nutrient analysis to assess air pollution influences seems remote.

Ozone exposure can lead to physiological disruptions that cause changes in

the nutrient concentrations of current-year ponderosa pine needles. At SF, the site

receiving the greatest ozone exposure, concentrations of P, K, Zn, and Fe in currentyear foliage were highest in ponderosa pine trees with the most severe ozone injury, presumably due to retranslocation from older, ozone-damaged needles to

younger foliage. Further studies are needed to elucidate mechanisms whereby

ozone-stressed trees respond to, and in some cases possibly compensate for, acute

and chronic ozone exposures and the interaction of these responses with associated N deposition.

References

Aber, John D.; Nadelhoffer, K.J.; Steudler, P.; Melillo, Jerry M. 1989. Nitrogen saturation in northern forest ecosystems. Bio- Nitrogen saturation in northern forest ecosystems

science 39: 378-386.

Andersen, C.P.; Hogsett, William E.; Wessling, R.; Plocher, M. 1991. Ozone decreases spring root growth and root carbohy- Ozone decreases spring root growth and root carbohydrate content in ponderosa pine the year following exposure. Canadian Journal of Forest Research 21 (8): 1,288. drate content in ponderosa pine the year following exposure

Arkley, Robert J.; Gersper, P.J.; Glauser, R. 1977. General description of ecosystem properties: Soils. General description of ecosystem properties: Soils. In: Photochemical oxi- General description of ecosystem properties: Soils.

dant air pollution effects on a mixed conifer forest ecosystem; A progress report. U.S. EPA. Corvallis, Oregon: Environmental Research Laboratory; 19-28.

Beyers, Jan. L.; Reichers, George H.; Temple, Patrick J. 1992. Effects of long-term ozone exposure and drought on the photo- Effects of long-term ozone exposure and drought on the photosynthetic capacity of ponderosa pine (Pinus ponderosa Laws.). New Phytologist 122: 81-90. Laws.)

Bytnerowicz, Andrzej; Poth, Mark; Takemoto, Brent. 1990. Effects of photochemical smog and mineral nutrition on ponde- Effects of photochemical smog and mineral nutrition on ponderosa pine seedlings. Environmental Pollution 67: 233-248. rosa pine seedlings

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 247

Clayton, J.L.; Kennedy, D.A. 1980. A comparison of the nutrient content of Rocky Mountain Douglas-fir and ponderosa pine

trees. Res. Note. INT-281. Ogden, Utah: Intermountain Forest and Range Experiment Station, USDA Forest Service. trees

Epstein, E. 1972. Mineral nutrition of plants: principles and perspectives. New York: John Wiley and Sons, Inc.; 212-214. Mineral nutrition of plants: principles and perspectives

Findlay, S.; Jones, C.G. 1990. Exposure of cottonwood plants to ozone alters subsequent leaf decomposition. Exposure of cottonwood plants to ozone alters subsequent leaf decomposition Oecologia 82 (2): 

248-253.

Fenn, Mark. 1991. Increased site fertility and litter decomposition rate in high pollution sites in the San Bernardino Moun- Increased site fertility and litter decomposition rate in high pollution sites in the San Bernardino Mountains. Forest Science 37: 1,163-1,181. tains

Fenn, Mark E.; Bytnerowicz, Andrzej. 1993. Dry deposition of nitrogen and sulfur to ponderosa and Jeffrey pine in the San

Bernardino National Forest in southern California. Environmental Pollution 81: 277-285. Bernardino National Forest in southern California

Flanagan, P.W.; Van Cleve, Keith. 1983. Nutrient cycling in relation to decomposition and organic-matter quality in taiga

ecosystems. Canadian Journal of Forest Research 13: 795. ecosystems

Glaubig, Robert; Poth, Mark . 1993. A method for total phosphorus determinations in nitric-perchloric digests of pine needles

and roots. Communications in Soil Science and Plant Analysis 24: 2,469-2,477. and roots

Hom, John L.; Riechers, George H. 1991. Carbon assimilation, allocation and nutrient use efficiency of ponderosa pine as a

compensatory response to ozone fumigation and water stress. In: Supplement to the bulletin of the Ecological Society of compensatory response to ozone fumigation and water stress

America, program and abstracts. Volume 72, No. 2; San Antonio Texas: 76th Annual ESA Meeting; 144 p.

Huettl, Reinhart F.; Fink, S.; Lutz, Hans J.; Poth, Mark; Wisniewski, Joseph. 1990. Forest decline, nutrient supply and diagnos- Forest decline, nutrient supply and diagnostic fertilization in southwestern Germany and in southern California. Forest Ecology and Management 30: 341. tic fertilization in southwestern Germany and in southern California

Killingbeck, Keith T. 1996. Nutrients in senesced leaves: keys to the search for potential resorption and resorption profi- Nutrients in senesced leaves: keys to the search for potential resorption and resorption proficiency. Ecology 77: 1,716-1,727. ciency

Klemmedson, J. O.; Neier, C. E.; Campbell, R. E. 1990. Litter fall transfers of dry matter and nutrients in ponderosa pine

stands. Canadian Journal of Forest Research 20: 1,105-1,115. stands

Marschner, H. 1986. Mineral nutrition of higher plants. London: Academic Press; 96-98. Mineral nutrition of higher plants

Miller, Paul M. 1983. Ozone effects in the San Bernardino National Forest. In: Proceedings of the symposium—air pollution Ozone effects in the San Bernardino National Forest

and the productivity of forests. Iowa City: Izaak Walton League of America; 161-197.

Miller, Paul M.; McBride, J.R.; Schilling, Susan L.; Gomez, Anthony P. 1989. Trends in ozone damage to conifer forests be- Trends in ozone damage to conifer forests between 1974 and 1988 in the San Bernardino Mountains of southern California. In: Transactions—effects of air pollution tween 1974 and 1988 in the San Bernardino Mountains of southern California

on western forests. Air and Waste Management Association; 309-324.

Miller, Paul M.; Parmeter, John R.; Taylor, O.C.; Cardiff, E.A. 1963. Ozone injury to the foliage of Ozone injury to the foliage of Pinus ponderosa. Pinus ponderosa. Phytopathology 53: 1072.

 Peterson, David L.; Arbaugh, Michael J.; Robinson, Lindsay J. 1991. Regional growth changes in zone-stressed ponderosa

pine (Pinus ponderosa) in the Sierra Nevada, California, USA. Holocene 1:50-61. ) in the Sierra Nevada, California, USA

Powers, Robert F. 1981. Nutritional ecology of ponderosa pine (Pinus ponderosa Laws.) and associated species. Berkeley: Laws.) and associated species

University of California; 234 p. Ph.D. dissertation.

Salisbury, Frank B.; Ross, Cleon W . 1985. Plant physiology. 3rd ed. Belmont, California: Wadsworth Publishing Co.; 106-113. Plant physiology

Schlesinger, William H.; DeLuca, E. H.; Billings, W.D. 1989. Nutrient use efficiency of woody plants on contrasting soils in the

Western Great Basin, Nevada. Ecology 70: 105-113. Western Great Basin, Nevada

Schulze, E. D. 1989. Air pollution and forest decline in a spruce (Picea abies) forest. Science 244: 776. ) forest

Schulze, E.D., Vries, W. De; Hauhs, M.; Rosen, K.; Rasmussen, L.; Tamm, C.O.; Nilsson, J. 1989. Critical loads for nitrogen

deposition in forest ecosystems. deposition in forest ecosystems. Water, Air and Soil Pollution 48: 451-156. deposition in forest ecosystems.

Temple, Patrick. J.; Miller, Paul R. 1994. Foliar ozone injury and radial growth of ponderosa pine. Canadian Journal of Forest Foliar ozone injury and radial growth of ponderosa pine

Research 24: 1,877-1,882.

Tjoelker, M.G.; Luxmoore, Robert J. 1991. Soil nitrogen and chronic ozone stress influence physiology, growth and nutrient

status of status of Pinus taeda L. and L. and Liriodendron tulipifera L. seedlings. New Phytologist 119: 69. L. seedlings

van Dijk, F.G.; Roelofs, J.G.M. 1988. Effects of excessive ammonium deposition on the nutritional status and condition of Effects of excessive ammonium deposition on the nutritional status and condition of

pine needles. Physiologia Plantarum 73: 494-501. pine needles

Wellburn, Alan. 1994. Air pollution and climate change: the biological impact. 2nd ed. New York: John Wiley and Sons, Inc. Air pollution and climate change: the biological impact

Session I1I Mature Ponderosa Pine Nutrient Use and Allocation Responses to Air Pollution Poth and Fenn

SESSION IV

Forest Health Monitoring:

National Perspectives

Paul R. Miller and Oleg Blum, Chairs

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 251

Abstract

A three-stage approach was applied to assess the quality of forests in southwestern Poland, which are heavily

affected with air pollution and insect infestations. In the first stage a ground evaluation of spruce stands was

done within the selected test areas. Three main characteristics of forest quality were determined as a result of

these works: defoliation, discoloration, and vigor of trees. Next, aerial color infrared photographs were taken

over the study area. They covered stands at different stages of forest decline. Detailed analysis of these photographs enabled discrimination of four levels of damage to spruce stands, as well as various phenomena,

which accompany forest decline and transformation (loose canopy closure, windbreaks and windfalls, afforestations and regenerations). Results of analysis of aerial photographs and ground-truth information supported the main stage of the work: digital classification of Landsat Thematic Mapper (TM) satellite images.

Digital classification of Landsat TM images can be useful for evaluation of forest quality and for rapid

large-area assessment of forest changes, which accompany degradation processes. Different methods of analyzing satellite data were studied to obtain maximum information content. Supervised classification based

on the TM3, TM4 and TM5 spectral bands was then applied. As a result of classifications for the test site,

which is seriously affected with air pollution, three levels of forest damage were distinguished, as well as

clear-cuts and vegetated openings accompanying forest decline. Comparison of two classifications performed

on images collected in 1984 and 1990 revealed great changes in forest structure.

Introduction

Air pollution has been considered the main cause of damage and deforestation to

forests in Europe. Air pollution increases along the direction of the prevailing wind

from southwest to northeast so that pollution originating in southern and western

Europe accumulates in Poland. This country itself contributes significantly to total

air pollution, adding great amounts of pollutants from fuel and power industries,

means of transport, and fossil fuel-heated homes.

Poland belongs to the group of most polluted nations because of the high concentration of SO2

. Limitation of the emission of sulphur compounds has not improved the situation considerably because about 50 percent of the total amount of

SO2

 comes from neighboring countries.

The technology applied by foresters to determine forest damage has been based

on analysis of the morphological changes in needles and tree growth. In the 1980’s the

Remote Sensing and Spatial Information Centre, OPOLIS, began to investigate the

application of remote sensing to the detection and determination of damage to pine

and spruce stands. At the beginning of the work, false-color aerial photographs were

used to detect air pollution damage to pine stands. It was possible to distinguish between dead and dying trees based on changes in crown color as seen on aerial photographs. Five categories of damage to pine forest have been recognized by calculating the

percentage of dead and dying trees in a given stand. A number of maps have been elaborated and published presenting the state of health of forests in different regions of Poland.

Damage to some Polish forests has been so extensive that it was decided to use

satellite images in the investigation of the problem. The Landsat Multispectral Scanner (MSS), Thematic Mapper (TM), SPOT images, and Cosmos space photographs

have been applied in these works. After several attempts we found that the Landsat

TM images have been the best source of information for this type of investigation.

This paper discusses a three-stage approach ground evaluation, aerial

photograph analysis, and digital classification of satellite images — to assess the

extent of air pollution to Polish forests.

Assessment of Forest Quality in Southwestern Poland

with the Use of Remotely Sensed Data1

1 An abbreviated version of this manuscript was presented at the International Symposium on Air Pollution

and Climate Change Effects on Forest

Ecosystems, February 5-9, 1996, Riverside, California.

2 Photogrammetrist, Professor of Geography, Professor of Ecology, respectively, Institute of Geodesy and Cartography Remote Sensing and Spatial

Information Centre, OPOLIS Jasna 2/

4, 00-950 Warsaw, Poland.

Zbigniew Bochenek, Andrzej Ciolkosz, Maria Iracka2
252 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Study Area

The Sudety Mountains, where the damage to coniferous forests was one of the

most serious in Poland, were chosen as a test site. This region, located at border

areas of Poland, Germany, and the Czech Republic, has been highly affected by

industrial activity and insect infestations. Concentration of industrial plants and

coal mines within the region caused serious degradation of large forest areas. Extent and intensity of the damages are so significant that this area is currently treated

as a region of ecological disaster and it has been called “Black Triangle.”

The test area covers the western part of the Sudety Mountains, i.e., part of the

Karkonosze and Izerskie Ranges, with dominating spruce stands forming a large

forest complex. The most serious damage of coniferous stands, as well as different

levels of forest transformation, occur in this region.

This differentiation of stands within the study area can support optimally the

main aim of the studies; it enables one to determine, which stand characteristics,

useful for large-area forest evaluation, can be derived from satellite data.

Materials and Methods

At the first stage of the work we decided to examine the usefulness of high-resolution satellite images for classifying health conditions of the forest, its species composition, and other forest phenomena. To fulfill this task, Landsat TM data collected in July 1984 and in August 1990, covering both test areas, were selected. The

analysis was done with the use of ERDAS image processing system. This system

enables interactive analysis of satellite images: radiometric and geometric correction, location of training areas on satellite images, determination of spectral signatures, and data classification.

Source Materials — Field Work and Aerial

Photographs

To find relationships between field characteristics of stands and their signatures on

satellite images, field evaluation of spruce stands was done at a preliminary stage

of the works by using more than 500 test fields located at the Karkonosze, Izerskie,

Sowie, and Stolowe Ranges. Field appraisal was performed on the basis of instruction prepared by the Forestry Research Institute. Each test field covered a homogeneous appraisal unit (1-10 ha), which was characterized through direct measurements and observations. The following information was collected during field work:

● Description of terrain conditions (type, relief, slope, aspect, type of cover and

degree of coverage by rocks, stones, debris, etc.).

● Description of stand (structure, layer, species composition, age, form of mix

ing, closure, average breast-diameter, mean height, height of crown base,

number of trees per unit area, including dying and dead trees).

● Evaluation of damage of stand (defoliation — Df, discoloration — Dc,

quality of trees — Dm).

Defoliation and discoloration was evaluated in accordance with International

Cooperation Program on Assessment and Monitoring Pollution Effects on Forests

(ICP) recommendations, while quality of trees was assessed on the basis of the

state of assimilatory apparatus, height increments, and vigor of trees.

At first stage of the work numerous relations were studied between stand/

terrain characteristics and spatial signatures of stands, derived from Landsat TM

satellite images, including elevation/aspect information. The following conclusions

were drawn from these studies (Zawila-Niedzwiecki 1994):

● The higher elevation above sea level, the lower height of trees and height

of crown base; lower quality trees found more on a high elevation than

less damaged trees.

Session IV Assessment of Forest Quality in Southwestern Poland Bochenek, Ciolkosz, Iracka
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 253

● High correlations (R2>0.90) were found between spectral response in

Landsat TM bands and some indices of quality of stands (Df, Dc, Dm).

The strongest relations exist for defoliation and quality index, while discoloration is less correlated. Usually the relations are defined through

multiple regression, which involves use of 2 to 4 spectral TM bands (most

commonly visible near infrared and middle infrared bands).

● Loose canopy closure disturbs relationships between ground parameters

and spectral signatures of stands, so if it exists on a larger area, this

information should be taken into account in the process of supervised

classification.

Analysis of the Karkonosze — Izerskie

Ranges Test Area

Aerial false-color photographs taken in July 1984 were the supplementary source

of information on the state of forests within the study area. The map of the health

status of forests, prepared on the basis of these photographs, was the reference

material for making appropriate selection of training and test areas, which was

used further in the process of supervised classification of satellite data.

At the first stage of analysis the rectification of a satellite image was performed,

through its transformation to the Gauss-Kruger projection system, which is commonly used for medium-scale topographic maps in Poland (1942 system). Next,

correlation analysis was done for all seven bands of Landsat TM, in order to choose

the three least-correlated channels. As a result of this analysis the following TM

spectral bands were selected: TM3, TM4, and TM5. This choice is consistent with

the other studies conducted at the European and American remote sensing centres.

These studies revealed high usefulness of the mentioned spectral bands for classifying forest areas (Horler and others 1986, Kadro 1988, Rock and others 1986). According to these studies TM4 and TM5 give the best information for vegetation

analyses, while TM3 combined with the latter ones forms a color composite that

emphasizes spectral differences within vegetation classes. TM5 is especially useful

for examining damage caused by loss of water in plant tissue and characterizing

vigor of vegetation. In the case of forests at the Sudety Mountains, we found as a

result of preliminary analysis that coniferous forests can be delineated with TM3,

while TM4 differentiates grasslands, deciduous and mixed forests. TM5 allows

different levels of defoliation to be discriminated, as well as clear-cut areas.

After preliminary analysis of color composite images, eight forest categories

were selected:

1. Slightly impaired spruce stands 5. Openings with vegetation cover

2. Heavily impaired spruce stands 6. Clear-cuts

3. Dying and dead stands 7. Deciduous and mixed stands

4. Young spruce stands 8. Dwarf mountain pine.

Training areas representing these categories were transferred to the satellite

image; next, spectral separability analysis was done to determine those categories

that could be classified with the acceptable accuracy (exceeding 80 percent). We

found that all selected classes, except dwarf mountain pine, were spectrally separable. So, seven forest categories were finally selected for classification, while areas

of dwarf mountain pine were transferred from topographic map.

After selecting the classes, supervised classification of the Karkonosze-Izerskie

Ranges test area was performed by using the 1984 image. Next, the same process

was repeated for the 1990 image by using the analogue set of training areas. Results

of both classifications were printed in the form of color maps. Finally postclassification accuracy assessment was done for both classification images by using

the set of test fields distributed randomly throughout the study area.

Session IV Assessment of Forest Quality in Southwestern Poland Bochenek, Ciolkosz, Iracka
254 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Discussion

Comprehensive analysis of Landsat TM satellite images, conducted on two test

areas for two dates of image acquisition, proved that spectral differentiation of

spruce stands in this region enables the discrimination of a maximum of three

classes of forest quality. Class No. 1 includes healthy and slightly impaired stands,

with up to 30 percent loss of assimilatory apparatus. Initial phases of spruce damages cannot be separated from healthy stands with the use satellite data. So training fields located at the stands with 0-30 percent loss of assimilatory apparatus

should be treated as representatives of one forest category.

Accuracy of discriminating on satellite images the other classes of spruce damage (more than 30 percent) depends on local conditions: canopy closure, soil cover,

understory vegetation, etc. For instance, if stands with loose canopy closure contain openings with vegetation cover, spectral signature for these stands changes

towards a class of better quality. Generally, there is the possibility to distinguish

heavily impaired stands with 30-60 percent loss of assimilatory apparatus, as well

as dying and dead stands that have lost more than 60 percent of needles.

Young coniferous stands were delineated well on the satellite image from 1984.

This class includes mainly spruce stands (and partly pine stands) at ages of 7-25

years, covering 9 percent of the Karkonosze and Izerskie Ranges test area. The

other classes, which are usually hardly discernible on satellite images (i.e. grasslands and afforestations) were quite well discriminated. As afforestations cover

about 7 percent of the total classified area, this class significantly characterizes

structure of the studied forests.

Main forest categories — three levels of damage of spruce forests and deciduous/mixed forests — were classified with accuracy exceeding 80 percent. By considering complex natural phenomena that existed at the studied region, the obtained results should be treated as fully satisfactory. They characterize both structure and health conditions of forest existing at the Sudety Mountains with acceptable accuracy.

Comparative analysis of two classified images from 1984 and 1990 enabled

evaluation of qualitative changes that occurred during 6 years within the study

area. The main observed phenomena were as follows (Zawila-Niedzwiecki 1994):

● An increase of the area comprising impaired spruce stands by 30 percent.

● A decrease of the area of dying and dead trees from 15 percent to 5 percent.

● An increase of the area of openings and clear-cuts from 8 percent to

15 percent with domination of vegetated openings in 1990.

Comparison of the classified image with forest maps revealed satisfactory consistency between classes determined from satellite data and stand characteristics

contained at forest maps. Thus, characteristics of stands, obtained as a result of

classification of Landsat TM images for the Karkonosze and Izerskie Ranges region, are suitable for large-area description of health conditions and structure of

stands in this region. Landsat TM satellite images, then, enable evaluation of general characteristics of health conditions and structure of stands and thereby permit

rapid evaluation of degradation processes.

References

Ciolkosz, A.; Iracka, M.; Zawila-Niedzwiecki, T. 1987. Monitoring of forest decline in Poland by remote sensing. Proceedings . Monitoring of forest decline in Poland by remote sensing

of the seminar on remote sensing and forest pollutants; 1987 March 11-12, Laxenburg: Austria; IIASA-EPRI, EA-575.

Ciolkosz, A.; Zawila-Niedzwiecki, T. 1990. Remotely sensed data and limitation of forest productivity in Poland. Nature and Remotely sensed data and limitation of forest productivity in Poland

Resources 26 (1): 41-44.

Horler, D.N.H.; Ahern, F.J. 1986. Forestry information content of thematic mapper data. International Journal of Remote Forestry information content of thematic mapper data

Sensing 7 (3).

Kadro, A.. 1988. Use of landsat TM data for forest damage inventory. ESA Publishing. Use of landsat TM data for forest damage inventory

Rock, B.N.; Vogelmann, J.E.; Williams, D.L.; Vogelmann, A.F.; Hozhizaki, T. 1986. Remote detection of forest damage. Bio- Remote detection of forest damage

science 36: 439-445.

Zawila-Niedzwiecki, T. 1994. Evaluation of forest state in threatened ecosystems with the use of satellite images and GIS. (in Evaluation of forest state in threatened ecosystems with the use of satellite images and GIS

Polish). Proceedings of the Institute of Geodesy and Cartography; Vol. XLI, No 90.

Session IV Assessment of Forest Quality in Southwestern Poland Bochenek, Ciolkosz, Iracka
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 255

Abstract

Vitality inventories of Scots pine stands, the most common species in Poland, have been done since 1991 by

using the ICP-Forest methodology. In Scots pine stands older than 40 years, 1,040 observation plots were

established. Defoliation was used as the primary indicator of stand vitality. During 1991 to 1995, SO 2 and

NOx were measured at 1,417 forest locations using passive samplers. The interrelation between changes of

precipitation amounts during the growing season and vitality of Scots zones with different air pollution levels

was studied. The analysis was done in three zones of Poland: north Poland is considered a relatively clean

area; the central part of Poland is characterized as having an average level of pollution; and south Poland

has the highest pollution. In each of these zones, decreasing concentrations of SO2 and NOx were observed

during the past 5 years. Changes of vitality in Scots pine stands caused by the amount of precipitation and

air pollution level in the growing season were studied for each of the three zones. 

Introduction

Forest health status in Poland has been strongly influenced for many years by high

levels of air pollution. Emissions of SO2 (5,000,000 tons) and NOx (2,500,000 tons) in

the 1980’s were among the highest in Europe (Ochrona Srodowiska i Gospodarka

Wodna 1987). Vegetation condition of forest trees in Poland have been unfavorable.

The forests in Poland grow on mainly sandy soil with poor nutrient availability

and shortage of water because of low precipitation during the growing season. In

these circumstances impacts of air pollution on forest health are particularly profound and cause increases of damage reflected in high defoliation levels

(Malachowska and Wawrzoniak 1995). In the beginning of the 1990’s, rapidly declining air pollution levels were observed. Emission of SO2

 and NOx was reduced

by 30 to 50 percent (Ochrona Srodowiska 1993). In spite of this, forest health condition did not indicate signs of improvement. A second essential factor that impacted

forest health condition was the amount of precipitation during the growing season. The combination of air pollution and water deficiency determined the health

status of foliage and its photosynthetic ability. Generally, the long-term health condition of Scots pine stands on a regional scale has been determined by the level of

air pollution, but yearly changes of health condition has depended on the amount

of precipitation during the growing season (Malachowska and Wawrzoniak 1995).

If water deficiency occurs during the growing season, response of stand vitality

differs between relatively healthy stands situated in low polluted areas and more

damaged stands in highly polluted areas.

The purpose of this study was to identify the reaction of Scots pine stand vitality in various levels of air pollution in three zones of Poland impacted by a shortage of water during the growing season in 1991-1995.

Methods

Mean defoliation of Scots pine stands in specific zones was used as an indicator of

forest vitality. This parameter was calculated on the basis of defoliation assessments on 5,000 trees in each zone (fig. 1). According to the ICP-Forest Manual (United

Nation Economic Commission for Europe 1994), assessments were done by trained

staff every year between July 1 and August 30, 1991-1995.

Response of Scots Pine Stand Vitality to Changes in

Environmental Factors in Poland, 1991 - 19951

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Forester, The Forest Research Institute, Bitwy Warszawskiej 3, 00-973

Warsaw, Poland.

Jerzy Wawrzoniak2
256 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

SO2 and NOx deposition were determined by passive method with 1-month

exposure of samplers (Krochmal and Kalina 1997). Measurements were made at

600 points in each zone. Mean values of SO2

 and NOx were calculated separately

for winter and summer periods in each zone. Yearly precipitation during 1991-

1995 was calculated as the mean of values from about 60 meteorological stations.

Results and Discussion

Comparison of air pollution levels during the summer period expressed in SO2 and

NOx indicator values shows differences between the three zones (fig. 2). The highest values were above 8 mg/m2

/24h in southern Poland and below 4.5 mg/m2/

24h in north Poland (fig. 2).

Considering mean defoliation of Scots pine stands in specific zones, a distinct

difference was found between south Poland and north Poland or central Poland,

but no difference between north and central Poland (fig. 3). The low level of air

pollution in north Poland seems to be an important factor influencing the condition of pine stands that appear more healthy than in south Poland where air pollution is much higher. The lack of differences between central and north Poland is

difficult to explain, despite differences in air pollution level.

Figure 1 — The three district air

pollution zones and locations of

observation plots in Poland.

I –north Poland (low level of air

pollution), II – central Poland (medium level of air pollution),

III – south Poland (high level of

air pollution).

Figure 2 — Changes in the SO2

 indicator during the summer period

for 1991 to 1995.

Figure 3 — Changes in Scots pine defoliation (two-tailed Tukey’s

Test) from 1991 to 1995.

Session IV Response of Scots Pine Stand Vitality to Changes in Environmental Factors in Poland, 1991 - 1995 Wawrzoniak
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 257

Figure 4 — Changes in Scots pine defoliation compared to the sum

of growing season precipitation in north Poland from 1991 to 1995 .

Figure 5 — Changes in Scots pine defoliation compared to the sum

of growing season precipitation in south Poland from 1991 to 1995.

Figure 6—Changes in Scots pine defoliation compared to the sum

of growing season precipitation in central Poland from 1991 to 1995.

The most interesting question that should be considered is the response of Scots

pine stands of various health status to changes of precipitation during the growing

season. By comparing changes between mean defoliation in a specific year and

changes of precipitation in the same year, a pattern was found in the healthy stands

of the north Poland zone. In 1992 and again in 1994, decreased precipitation

(Miesieczny Preglad Agrometeorologiczny 1991-1995) was followed by increased

mean defoliation in the same year. In 1993 and in 1995, increased precipitation was

followed by decreased mean defoliation levels in the same year (fig. 4).

The response of Scots pine stand vitality, indicated by mean defoliation to changing amounts of precipitation, was different in the more polluted and more damaged stands in the south Poland zone. Decreasing precipitation in 1992 was not

followed by increasing mean defoliation in the same year, as happened in north

Poland. However, mean defoliation increased in 1993 despite increased precipitation. Improvement in stand health condition, indicated by decreased mean defoliation, was found in 1995 after 2 years of increased precipitation (fig.␣ 5).

Similar reactions of Scots pine stands to precipitation amount was also found

in central Poland (fig. 6) where the air pollution level is lower than in south Poland

(fig. 2), and the health condition of Scots pine stands is better (fig. 3).

Session IV Response of Scots Pine Stand Vitality to Changes in Environmental Factors in Poland, 1991 - 1995 Wawrzoniak
258 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The different responses of the vitality indicator (mean defoliation) to changes

in precipitation amount during the growing season seems to be connected to the

level of health condition at the beginning of the period studied.

In healthy pine stands in north Poland, the majority of trees had three or four

years of needles. A small amount of precipitation during the growing season resulted in a shortage of water and trees immediately dropped one or two of the

oldest years of needles, regardless of the consequences to buds for the next year.

Trees in this region easily regenerate foliage the following year, when precipitation

in the growing season is increased.

Scots pine stands in south Poland are in poor health condition and usually

trees have 2 years of needles. If precipitation is low during the growing season,

trees cannot drop needles without weakening the assimilation processes and endangering their survival for the next year. The only possible response in this condition is limiting bud formation in the same year and maintaining needle retention at

the same level in the following year. If precipitation is adequate during the growing season, Scots pine trees show more effective bud formation in the current year

and the improved vitality of stands is reflected by increased needle retention in the

next year.

Conclusions

The response of Scots pine stand vitality to a shortage of water during the growing

season is different in high polluted areas and low polluted areas. The health

condition of Scots pine stands at the beginning of drought periods is crucial to the

response of the stands. Scots pine stands in good health condition respond to drought

during the growing season in the same year by dropping needles. Scots pine stands

in poor health condition respond to drought in the growing season by changes in

the bud formation process and increased defoliation in the following years.

Acknowledgments

I thank Laurie Dunn for technical editing of this manuscript.

References

Krochmal, D.; Kalina, A. [In press]. Measurements of nitrogen dioxide and sulphur dioxide concentrations in urban and rural Measurements of nitrogen dioxide and sulphur dioxide concentrations in urban and rural

areas of Poland using passive sampling method. areas of Poland using passive sampling method. Environmental Pollution. areas of Poland using passive sampling method. 

Malachowska, J.; Wawrzoniak, J. 1955. Stan uszkodzenia lasow w Polsce na podstawie badan monitoringowych. Stan uszkodzenia lasow w Polsce na podstawie badan monitoringowych. Warszawa: Stan uszkodzenia lasow w Polsce na podstawie badan monitoringowych. 

Biblioteka Monitoringu Srodowiska.

Miesieczny Przeglad Agrometeorologiczny. 1991-1995. Warszawa: Instytut Meteorologii i Gospodarki Wodnej. Warszawa: Instytut Meteorologii i Gospodarki Wodnej.

Ochrona Srodowiska i Gospodarka Wodna. 1987. Warszawa: Glowny Urzad Statystyczny. Warszawa: Glowny Urzad Statystyczny.

Ochrona Srodowiska. 1993. Warszawa: Glowny Urzad Statystyczny. Warszawa: Glowny Urzad Statystyczny.

United Nations Economic Commission for Europe. Manual on methods and criteria for harmonized sampling, assessment, . Manual on methods and criteria for harmonized sampling, assessment,

monitoring and analysis of the effects of air pollution on forests. 1994. monitoring and analysis of the effects of air pollution on forests. 1994. Hamburg and Prague: Convention on Long-Range monitoring and analysis of the effects of air pollution on forests. 1994. 

Transboundary Air Pollution.

Session IV Response of Scots Pine Stand Vitality to Changes in Environmental Factors in Poland, 1991 - 1995 Wawrzoniak
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 259

Air Pollution and Forest Health Studies along a

South-North Transect in Poland1

Stefan Godzik, Jerzy Szdzuj, Tomasz Staszewski, Wlodzimierz Lukasik2

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Professor of Natural Sciences, Physicist, Biologist, and Biologist, respectively, Institute for Ecology of Industrial Areas, Kossutha 6, 40-833

Katowice, Poland.

Abstract

Air pollution, bulk deposition and throughfall, soil characteristics, needle chemistry, and forest injury were

studied on six permanent plots from the south (Brenna and Salmopol in the Beskidy Mountains) to the north

(Gac, the Baltic Sea coastal area) in Poland. The concentrations of sulfur dioxide and nitrogen dioxide were

the highest at the Katowice location and the lowest at Gac. Annual mean concentrations of sulfur dioxide at

the Brenna, Salmopol, Katowice, and Puszczykowo sites, have exceeded the critical level of 20 μg m-3 proposed by United Nations- Economical Committee for Europe (UN-ECE) bodies, and the Polish national

standard (32 μg m-3 for the whole country, and 11 μg m-3 for special protected areas, e.g. National Parks).

Nitrogen dioxide concentration has neither exceeded the UN-ECE nor the Polish national standards for the

whole country (30 and 50 μg m-3, respectively). Bulk deposition from all locations was acidic. The pH of

throughfall in spruce stands was lower than the bulk deposition by about 0.5 of a unit. A larger decrease of

pH has been found occasionally. Concentration of all elements measured was higher in throughfall as compared to bulk deposition. However, the differences in concentration of elements between bulk deposition and

throughfall varied. The differences were from three to five times for sulfates, but for nitrogen compounds, such

as nitrates and ammonium, differences were smaller or did not exist at all. The soil pH of all plots ranged

from 2.9 to 3.6 (H2O) in the organic layer. The highest concentrations of lead and zinc were found in the

organic layer of the sites in southern Poland. The greatest needle loss (crown transparency) was found at the

highest elevation site (Salmopol) for Norway spruce (Picea abies) and in the most polluted site (Katowice)

for Scots pine (Pinus sylvestris). The smallest crown transparency of spruce was found for the Gac location.

Discoloration of needles was the highest at Gac, followed by Katowice and Salmopol. Insects contributed to

the observed higher crown transparency on plots at Brenna and needle discoloration at Gac.

Introduction

Sulfur dioxide (SO2) air pollution acting directly or indirectly has been one of the

major causes for poor forest health status and forest decline in Poland (Ashmore

and others 1990, Forschungsbeirat Waldschaeden/Luftverunreinigungen 1989,

Godzik 1984, Godzik and Sienkiewicz 1990, Narodowy Program 1988, Koth-Jahr

and Kollner 1993, Paluch 1968). According to model calculations, a gradient of both

concentrations and depositions of sulfur compounds was found from the south to

the north of Poland (Narodowy Program 1988) (fig.1). The reason of such uneven

distribution was mostly emission from sources located in the south of Poland, the

Czech Republic, and Germany (former German Democratic Republic). Results of

forest injury inventories in Poland have followed a similar pattern. The most severe injuries (class three) were noted in southern Poland along the border with the

Czech Republic. Severe injury was also observed on a local scale in other parts of

the country (Godzik and others 1996, GUS 1995, Malachowska and Wawrzoniak

1995). Although emission of air pollutants in recent years has decreased, the pattern of its distribution has remained similar.

The total area of forest in Poland is 873,200 hectares, comprising 27.9 percent of

the total area of the country (GUS 1995). Scots pine (Pinus sylvestris) cover about 70

percent of the total forest area. Norway spruce (Picea abies) cover about 6 percent of

the forest (GUS 1995).

On the basis of data concerning the gradient of sulfur dioxide concentration in

Poland, six permanent plots in coniferous forests have been established. Climatic
260 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

differences are represented by plots at different locations above the sea level (asl) in

the Brenna region and at the Baltic Sea coast. Differentiation of air pollution level is

represented along the line from the south to the north of Poland (fig. 1).

This paper reports a study of the concentrations of sulfur dioxide and nitrogen

dioxide in the air, deposition of air pollutants and mineral nutrients, and their relationship to forest stand parameters, such as needles and soil characteristics, on six

permanent plots in Poland.

Site Description, Material, and Methods

Permanent forest plots were established in southern Poland (two plots in the Brenna

area and one plot in the Katowice region), central Poland (Puszczykowo —

Wielkopolski National Park (WNP)), and two plots in northern Poland (Gac —

Slowinski National Park (SNP) — at the Baltic Sea coast) (fig.1). The size of the plots

was 50 by 50 m. All plots met standards required by the measurement program for

sites for Integrated Monitoring for Forest Ecosystems within the Long Range

Transboundary Air Pollution (LTRAP) UN-ECE protocol (Environmental Data Centre 1989).

The two plots in the Brenna area, the Beskidy Mountains, were located at 1,000

m asl (Salmopol) and 650 m asl (Brenna). The main tree species in those sites were

Norway spruce with additions of white fir (Abies alba) or beech trees (Fagus sylvatica).

The plot in the Katowice region was basically a Scots pine stand with admixture of

broad leaf species in the lower layer. This region was the most polluted and has

been influenced by air pollution for more than a century. The only tree species at

the Puszczykowo plot was Scots pine. In Gac, at the Baltic Sea coast, two plots were

established in spruce and pine stands. On every plot 15 throughfall collectors were

randomly distributed. In an open area, collectors for bulk precipitation were placed.

Ceramic vacuum cup lysimeters were installed at 25 and 50 cm depth on each of

plots in a regular net. All samples were collected and analyzed monthly.

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik

Figure 1 — Locations of six forest permanent plots in Poland

with isolines showing SO2 annual

mean concentration in the air. SNP

= Slowinski National Park; WNP

= Wielkopolski National Park.

Gac (SNP)

Puszczykowo (WNP)

SO concentration < 32 2 μg/m3

SO concentration 32 - 64 2 μg/m3

SO concentration > 64 2 μg/m3

Warszawa

Katowice

Brenna

Salmopl
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 261

45

40

35

30

25

20

15

10

5

0

[ ug/m-3 ]

Brenna Salmopol Katowice Puszczykowo (WNP) Gac (SNP)

–

–

–

–

–

–

–

–

–

–––––

NO2 SO2

Ions were determined with ion chromatograph DX-100 with column AS4A.3

Metallic cations were determined with atomic absorption spectroscope Varian Spectra AA 300. Concentration of NH4

 was determined by using the Nessler method

(Hermanowicz and others 1976). Soil samples were collected from each horizon

and analyzed separately. For the chemical soil analysis, standard methods recommended by the Soil Service Laboratories were applied (Litynski and others 1976).

Needles for chemical analysis were taken in late October. Forest class injury

was assessed based on degree of crown transparency and discoloration from methods recommended by the UN-ECE ICP Forest working group. Concentrations of

sulfur dioxide and nitrogen dioxide in Brenna, Salmopol, Katowice, and

Puszczykowo were determined as 24-hour means. Sulfur dioxide was absorbed in

H2O2, producing sulfates that were determined with ion chromatography. Nitrogen dioxide was determined by methods recommended by EMEP: absorbed on a

sintered glass filter impregnated with KI-NaAsO2 mixture, and colorimetrically

determined by using sulfonamide and N-(1-Naphtyl)ethylenediamine dichloride

(NEDA) reagents (Environmental Data Centre 1989). Data for the Baltic coast area

is from the national base network for air pollution measurements (PIOS 1995).

Results and Discussion

Air Pollution

Differences in concentration of sulfur dioxide and nitrogen dioxide were not found

between two elevations in the Beskidy Mountains (fig. 2). The data show that

monthly and yearly mean concentrations of sulfur dioxide were higher than for the

Rhine-Westphalen area in 1987 (Koeth-Jahr and Koellner 1993); and the values for

nitrogen dioxide were lower than in the same area of Germany (Koeth-Jahr and

Koellner 1993). For the Schwarzwald area, the values did not exceed 29 μg m-3, and

for the growing season were lower by 50 percent (Forschungsbeirat Waldschaeden/

Luftverunreinigungen 1989). The concentration pattern of sulfur dioxide on the

plots showed similar differences between winter and summer. The mean values

were rather high, given that sulfur dioxide is not the only air pollutant hazardous

for trees. Concentrations of this air pollutant are lower than proposed by the International Union of Forest Research Organizations (IUFRO) for not extreme environmental conditions (50 μg m-3). However, except for the Gac area, values of sulfur

dioxide were in the range or higher than proposed as critical levels by the UN-ECE

convention on Long Range Transboundary Air Pollution (20␣μg m-3 and 30␣μg m-3

for SO2 and NO2, respectively; Ashmore and Wilson 1994). Concentration of sulfur

3 Mention of trade names or products

is for information only and does not

imply endorsement by the U.S. Department of Agriculture.

Figure 2 — Annual mean concentration of SO2

 and NO2 (μg

m-3) in the air over permanent

plots.

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik
262 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

dioxide has exceeded or was very close to the Polish national standard (32 μg m-3 as

yearly mean value) for the whole country. For the WNP, as a “special protected

area,” the standard is 11␣μg m-3. Concentration of sulfur dioxide was found to be

several times higher.

The most probable direct effects on the forests caused by air pollutants (only

SO2) were the sites in the Katowice area, and to a lesser extent in the WNP. Direct

effects of nitrogen dioxide seems to be less probable. For the plots located in the

Beskidy Mountains, ozone should be considered as a possible factor contributing

to the forest health status (Godzik, this volume).

Deposition

As expected from the model calculations (Narodowy Program 1988), the bulk deposition concentrations of most elements were higher in the polluted locations:

Katowice and Puszczykowo (WNP) (see Staszewski 1996 for results of all six locations). The most interesting data for loads of sulfur and nitrogen are the differences

between pine and spruce stands at the Gac site (table 1). This difference may be the

result of species-specific characteristics and/or number of trees for the same area.

No similar comparison could be made for the southern Poland stands. Higher loads

of sulfur and nitrogen for the spruce stands in Brenna and Salmopol compared to

the Katowice area can be explained by rainfall that is almost two times higher in

the Brenna region. If spruce stands were compared, loads for spruce in the Katowice

region would likely be even higher. We may also consider that the extent of the

earlier and more severe deterioration of spruce stands compared to pine stands in

the Silesia Industrial Region could be attributed to differences in loads (Godzik

and Harabin 1968). Amount (load) of sulfur and nitrogen deposited to all forest

ecosystems under study exceeded, except SNP sites, the critical loads recommended

by the UN-ECE.

1 The values are a sum of data from separate sampling periods (sampling intervals). During the whole sampling period,

concentration of elements in throughfall was always higher if compared to the same sampling time for the bulk deposition.

Table 1 Table 1 — Annual loads of sulfur and nitrogen calculated for bulk deposition and throughfall (kg ha Table 1 -1 a-1).1

Location and species Sulfur Nitrogen

Salmopol: spruce

Bulk deposition 15.1 26.6

Throughfall 36.3 30.3

Brenna: spruce

Bulk deposition 13.6 17.2

Throughfall 41.0 28.0

Katowice: pine

Bulk deposition 12.7 10.4

Throughfall 27.3 13.0

WNP: pine

Bulk deposition 13.1 13.7

Throughfall 27.8 15.5

SNP

Bulk deposition 8.4 8.8

Throughfall(pine) 10.0 8.0

Throughfall(spruce) 17.0 12.5

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 263

Soil Characteristics

Soils of all but one location (spruce site in Gac) were of the podzolic type and of

low pH (Godzik and others 1994). Soil profiles represented typical handbook

descriptions of distributions of genetic horizons. Low amounts of calcium and

magnesium that are mostly responsible for the soil pH were found. In all soils

potassium content was higher than calcium. The highest concentrations of lead

and cadmium were found in the organic soil layer. The concentration of lead in

samples from southern Poland exceeded the concentration recognized as “normal”

for soils of this region.

Soil investigation data showed that soils on all but one plot were comparable.

Thus, this data shows that climate and air pollution were the major variables in our

studies. Differences in forest injury class at plots under investigation could not be

explained by differences in soil properties. This conclusion is based on the comparison of soil data from two plots in the Brenna area for spruce, and KatowiceWNP for pine. Investigations performed earlier for the Izerskie-Beskidy Mountains showed similar conclusions (Godzik and Szdzuj 1994).

Needle Chemistry

The highest concentration of sulfur in needles of spruce has been found for the

coastal area (Gac), followed by that found in the Salmopol and Brenna needles

(fig. 3). The highest injury class has been assessed at the Salmopol location, and the

least at the coastal area (SNP). Further investigations are needed to explain the

highest concentration of sulfur in spruce needles from the Gac location, where the

air pollution was the lowest. One possible explanation is the difference in soil properties at this site, as compared to the locations at Brenna and Salmopol. The influence of the nearby Baltic Sea seems to be less likely when looking for the results for

sulfur concentration in pine needles. For the pine stands, the sulfur content of needles

is higher in forests of higher injury class: Katowice had the highest pine needle

injury, followed by WNP and SNP.

Forest Injury

The highest mean defoliation (crown transparency) in spruce stands was found in

Salmopol, followed by Brenna and the Gac (SNP) (fig. 4). At the Salmopol location,

100 percent of trees were in injury classes 2 and 3. At the Brenna and Gac locations

60 and 30 percent, respectively, were in these classes. Only trees at the Brenna location were found with no injury. At the Brenna and Gac locations, insect injuries

were noted.

Figure 3 — Content of sulfur

(percent dry weight) in 1-year-old

spruce and pine needles.

0.2

0.13

0.16

0.14

0.12

0.1

0.08

0.06

0.04

0.02

0

[ pct dry weight ]

Brenna Salmopol Gac Katowice Puszczykowo Gac 

 (SPN) (WNP) (SNP) 

–

–

–

–

–

–

–

–

–

–

–

Spruce Pine

––––––

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik
264 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Most severe defoliation of pine trees has been found at the Katowice plot, where

80 percent of trees were in the injury class 3. The numbers for the WNP and SNP

plots are 20 and 25 percent, respectively.

Mean values for discoloration of spruce trees are the highest for the Gac, followed by Salmopol and Brenna (fig. 4). However, at the Salmopol plot 35 percent of

trees are in the discoloration classes 2 and 3. For the two remaining plots the numbers are 10 percent for Gac and 5 percent for Brenna. No discoloration was found

on 35 percent of trees at the Brenna plot, followed by Gac with 25 percent, and

Salmopol with 5 percent.

The highest mean discoloration of pine trees was found at the Katowice

plot. No difference in discoloration was found between the WNP and SNP locations (fig. 4). All trees at the Katowice plot are in the discoloration class 1 (60

percent) and 2 (40 percent). For the WNP plots they are 30 percent and 10 percent; and 55 percent and 0 percent for the SNP plots — class 1 and 2, respectively. Similar patterns of sulfur concentration in pine needles have been found.

Because soil properties at all plots did not differ distinctly, the discoloration

differences may be more related to concentration of sulfur dioxide and load of

sulfur deposited to these ecosystems.

Conclusions

Concentrations of sulfur dioxide were the highest in the Katowice region, followed

by the Wielkopolski National Park (WNP), the Brenna (Beskidy Mountains) and

Gac (Slowinski National Park) locations. Concentrations of nitrogen dioxide followed a similar pattern. Load of sulfur and nitrogen to the forest floor differs depending on localization and tree species. Higher loads were measured on spruce

stands as compared to the pine (Gac). Higher loads have been found for spruce

stands in Salmopol and Brenna when compared to pine stands in the Katowice

region (where concentration of air pollutants is higher). The result is caused by

rainfall that is almost two times higher in the Beskidy location. However, other

factors may be involved as well.

Forest soils at all sites were acidic. Except for the soil at the spruce stand at Gac,

they were also of similar physico-chemical characteristics. The highest levels of

lead and cadmium were found in an organic layer and in sites in the southern part

of the country.

Figure 4 — Mean values (percentage) of defoliation and discoloration of pine and spruce stands

on permanent plots.

45

40

35

30

25

20

15

10

5

0

[ pct ]

Salmopol Brenna Gac Katowice Puszczykowo Gac 

 (SNP) (WNP) (SNP) 

–

–

–

–

–

–

–

–

–

–

Defoliation Discoloration

–––––––

Spruce Pine

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 265

No clear relationships were found between air pollution — concentration and

load — and forest health on plots under investigation. However, injury of spruce

and pine trees were higher at locations where air pollution levels were also higher.

The comparison of results from Brenna and Salmopol suggests that climatic parameters are an important factor in forest injury. However, a lack of measurements

resulted in the inability to assess the possible contribution of ozone to the more

severe forest injury at the higher elevations.

Acknowledgments

We thank F. Andersson, Peringe Grennfelt, and J. Bergholm of Sweden, for

advice and help. We are also grateful to reviewers for very valuable comments,

suggestions, and correction of the language. These studies started within a

frame agreement on scientific cooperation between the Polish and Swedish

Academies of Science.

References

Ashmore, M.R; Bell, J.N.B; Brown, I.J., eds. 1990. Air pollution and forest ecosystems in the European community. Air pollution and forest ecosystems in the European community. Air Pollu- Air pollution and forest ecosystems in the European community.

tion Research Report 29. Brussels: Commission of the European Communities.

Ashmore, M.R.; Wilson, R.B., eds. 1994. Critical levels of air pollutants for Europe. London: Department of Environment. Critical levels of air pollutants for Europe. London: Department of Environment.

Environmental Data Centre 1989. Field and laboratory manual. International Cooperative Program on Integrated

Monitoring. Monitoring. Helsinki: National Board of Waters and the Environment. Monitoring. 

Forschungsbeirat Waldschaeden/Luftverunreinigungen. 1989. Dritter Bericht. Dritter Bericht. Kernforschungszentrum Karlsruhe GmbH. Dritter Bericht. 

Godzik, S. 1984. Air pollution problems in some central European countries: Czechoslovakia, German Democratic Republic

and Poland. and Poland. In: Koziol, M.J.; Whatley, F.R., eds. Gaseous air pollutants and plant metabolism. London: and Poland. 

Butterworth; 25-34.

Godzik, S.; Harabin, Z. 1968. Waldbestaende im Raume Katowice und Myslowice zu Ende des XIX Jahrhubderts und

Gegenwaertig. Gegenwaertig. In: Paluch J.; Godzik S., eds: Katowice: Materialy VI Miedzynarodowej Konferencji Wplyw Zanieczyszczen Gegenwaertig. 

Powietrza na Lasy. Zaklad Badan Naukowych GOP PAN; 355- 370.

Godzik, S.; Lukasik, W.; Poborski, P.S.; Staszewski, T.; Szdzuj, J.; Korlacka, I. 1994. Oddzialywanie i obieg zwiazkow siarki i Oddzialywanie i obieg zwiazkow siarki i

azotu zawartych w powietrzu i opadach w eksosystemach lesnych - badania w gradiencie stezen i klimatu (Effects and azotu zawartych w powietrzu i opadach w eksosystemach lesnych - badania w gradiencie stezen i klimatu

turnover of sulfur and nitrogen compounds from the air and deposits in forest ecosystems - investigation in gradients of

concentration and climate). Raport 324/DN/93. Katowice: Instytut Ekologii Terenow Uprzemylowionych.

Godzik S.; Sienkiewicz, J. 1990. Air pollution and forest health in central Europe: Poland, Czechoslovakia, and the German

Democratic Republic. Democratic Republic. In: Grodzinski W.; Cowling, E. B.; Breymeyer, A.I., eds. Ecological risks. Perspectives from Poland Democratic Republic.

and the United States. Washington, D.C.: National Academy Press; 155-170.

Godzik S.; Szdzuj J. 1994. A comparison of air, soil and spruce needle chemistry in The Izerskie and Beskidy Mountains. A comparison of air, soil and spruce needle chemistry in The Izerskie and Beskidy Mountains. In: A comparison of air, soil and spruce needle chemistry in The Izerskie and Beskidy Mountains.

Solon, J.; Roo-Zielinska, E.; Bytnerowicz, eds. Climate and atmospheric deposition studies in forests. Conference Papers

19, Warszawa: IGSO PAS; 187-195.

Godzik, S.; Szdzuj, J.; Staszewski, T. [In press]. Research on air pollution and its effects to forests in Poland. Research on air pollution and its effects to forests in Poland. NATO ARW Research on air pollution and its effects to forests in Poland. 

Workshop. Spindleruv Mlyn.

GUS 1995. (Glowny Urzad Statystyczny 1995). Ochrona Srodowiska 1995 (Environment Protection 1995). Warszawa 1995. Ochrona Srodowiska 1995

Hermanowicz, W.; Dozanska, W.; Dojlido J.; Koziorowski, B. 1976. Fizyczno-chemiczne badania wody i sciekow. Fizyczno-chemiczne badania wody i sciekow.

Arkady. Warszawa.

Koth-Jahr,I.; Koellner, B., eds. 1993. Forschungsschwerpunkt Luftverunreinigungen und Walsdschäden. Ziele - Ergebnisse - Forschungsschwerpunkt Luftverunreinigungen und Walsdschäden. Ziele - Ergebnisse -

Schlussfolgerungen. Schlussfolgerungen. Ministerium f. Umwelt, Raumordnung und Landwirtschaft des Landes Nordrhein-Westfalen. Schlussfolgerungen. 

Litynski, T.; Jurkowska, H.; Gorlach, E. 1976. Analiza chemiczno-rolnicza. Analiza chemiczno-rolnicza. PWN Warszawa. Analiza chemiczno-rolnicza. 

Malachowska, J.; Wawrzoniak, J. 1995. Stan uszkodzenia lasow w Polsce w 1994 r. na podstawie badan monitoringowych

(Forest injury in Poland in 1994 according to monitoring). Panstwowa Inspekcja Ochrony Srodowiska. Warszawa:

Biblioteka Monitoringu Srodowiska.

Narodowy Program Ochrony Srodowiska do roku 2010. 1988 (National program of natural environment protection till the

year 2010). year 2010). Draft. Ministerstwo Ochrony Srodowiska. Warszawa. year 2010). 

Paluch, J. 1968. Einige Probleme der Luftverunreinigung in Hochindustrialisierten Gebieten. Einige Probleme der Luftverunreinigung in Hochindustrialisierten Gebieten. In: Paluch, J; Godzik, S., eds. Einige Probleme der Luftverunreinigung in Hochindustrialisierten Gebieten. 

Materialy VI Miedzynarodowej Konferencji Wplyw Zanieczyszczen Powietrza na Lasy. Katowice: Zaklad Badan Naukowych

GOP PAN; 26 p.

PIOS (Panstwowa Inspekcja Ochrony Srodowiska) 1995. (Zanieczyszczenia powietrza w Polsce w 1994 roku. (Zanieczyszczenia powietrza w Polsce w 1994 roku. (Air pollution in (Zanieczyszczenia powietrza w Polsce w 1994 roku. 

Poland for the year 1994). Warszawa: Praca zespolowa. Biblioteka Monitoringu Srodowiska.

Staszewski, T. 1996. Depozycja i obieg skladnikow zanieczyszczen powietrza i mineralnych skladnikow odzywczych na szesciu

lesnych powierzchniach badawczych. lesnych powierzchniach badawczych. Report (Deposition and turnover of air pollution components and mineral nutri- lesnych powierzchniach badawczych. 

ents on six permanent forest plots). Raport 110/DN/96. Katowice: Instytut Ekologii Terenow Uprzemyslowionych.

Session IV Air Pollution and Forest Health Studies Godzik, Szdzuj, Staszewski, Lukasik

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 267

Abstract

About 886.5 Mha in Russia is occupied by forests, including 763.5 Mha of tree stands and 123 Mha of nonstocked lands. The Russian forests comprise about 22 percent of the earth’s forest area or 43 percent of the

earth’s temperate and boreal forests. Main forest- forming species are Larix sp. (32 percent of the growing

stock), Pinus sylvestris (20 percent), Picea sp. (15 percent), Betula sp. (13 percent), Pinus sibirica (10

percent), Populus tremula (5 percent), Abies sp. (3 percent), Quercus sp. (1 percent) and other species (1

percent). About one-third of tree stands and two-thirds of forest ecosystems in Russia are disturbed by natural and anthropogenic stresses. Most prevalent are natural causes of disturbance (climate, aging of stands,

fires, pests) that affect 200-250 Mha of forest stands. About 45 Mha of forest stands (6 percent of stocked

area) are under the impact of anthropogenic influence. Atmospheric pollution is the most dangerous form of

anthropogenic stress in the Russian forests.

Introduction

According to national inventory data from 1993, lands classified from the Forest

Fund of Russia (1995) occupy 1,180.9 Mha. These lands include 294.4 Mha of nonforest land (mires, water, rocks, wastelands, etc.) and 886.5 Mha of forest land, including 763.5 Mha of tree stands and 123.0 Mha of non-stocked lands (woodlands,

clear-cuttings, burned stands, etc.). These forest lands comprise about 22 percent of

the Earth’s forest area or 43 percent of the earth’s temperate and boreal forests

(United Nations 1992).

Forests are distributed in plain forest-tundra and subarctic mountains (14.1

percent), boreal plain and mountain regions (66.0 percent), zone of mixed forests

(1.7 percent), zones of forest-steppes, steppes, and subboreal mountains (17.9 percent), and zone of semi-deserts and subarid mountains (0.3 percent). Natural forests comprise 98 percent and plantations 2 percent of the total forested area. About

350 Mha of forests are in permafrost territory, and about 340 Mha in mountains

(mainly in subarctic and boreal regions). The average yield of the Russian forests is

not high: the growing stock of mature and overmature forests is 136.7 m3/ha

(table 1). Main forest-forming species are Larix sp. (32 percent of the total growing

stock), Pinus sylvestris (20 percent), Picea sp. (15 percent), Betula sp. (13 percent),

Pinus sibirica (10 percent), Populus tremula (5 percent), Abies sp. (3 percent), Quercus

sp. (1 percent), and other species (1 percent). Forests are considerably damaged by

natural and anthropogenic stresses.

This paper presents the current health status of the Russian forests and predicts possible trends caused by global climate change.

Forest Health Status and Factors of Disturbance

Climate

One of the primary disturbance factors of trees in the frontier forests of the foresttundra and subarctical mountains on permafrost territories of Siberia and eastern

Russian is severe climatic conditions (table 2). Trees in open, sparse unproductive

forests and woodlands of these regions have crowns heavily damaged by frost and

wind. Leaf area index (LAI) of trees is three to five times less than LAI of trees

in more southern territories. Healthy stands and healthy trees in stands and

Forest Health Status in Russia1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Professor of Ecology, Russian Academy of Sciences, St.-Petersburg Research Institute of Forestry, 21

Institutskiy Ave., St. Petersburg

194021, Russia.

Vladislav A. Alexeyev2
268 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

woodlands of permafrost territory are a rare phenomenon. The area of such naturally disturbed forests is estimated to exceed 150-200 Mha.

Long-term monitoring of several Siberian stands showed no change for the

worse of the health status of such forests. Moreover, data indicate improvement of

health condition of these forests. However, the improved condition of forests has

not yet been statistically proven.

As a result of global warming in the next 100 years, an improvement of forest

health on the present permafrost territory and decrease of damaged area of stands

can be expected.

Session IV Forest Health Status in Russia Alexeyev

1 The data presented in this table are the author’s expert estimates of disturbed forest areas (except the total area of burnt stands and stands damaged by pests) and may

not agree with point of view of Federal Forest Service of Russia.

2 Percent of total stocked area.

3 Sum subtotal and total do not equal to mathematical sum of items because different factors can act simultaneously in the same territory.

Table 2 Table 2 — Present and forecasted forest area of Russia disturbed by natural and anthropogenic factors. Table 2 1

The primary Present status Forecasted

 reason of damage status in

year 2100

Stands Mha pct 2 Ecosystems Mha pct 2 Stands Ecosystems

Natural factors

Climate 150-200 20-26 150-200 20-26 decrease decrease

Fires 10 1 50 7 increase increase

Pest 10 1 20 3 increase increase

Aging of trees 100-150 13-20 100-150 13-20 decrease decrease

Subtotal pct3 200-250 26-33 250-300 33-40 decrease decrease

Anthropogenic factors

Harvesting 20 3 100 13 the same the same

Pollution 5 1 15 2 decrease decrease

Fires 20 3 100 13 the same the same

Subtotal 3 45 6 215 28 the same the same

Total3 240-290 31-38 460-510 60-67 decrease decrease

Table 1 — Age structure of area and growing stock in Russian forests (Forest Fund of Russia 1995). Table 1

Age classes of stands

 Statistic data Young Middle-aged Premature Mature and Total

overmature

Area,

million ha 134.4 201.6 74.8 352.7 763.5

percent 17.6 26.4 9.8 46.2 100

Growing stock,

billion m3 3.7 20.2 10.5 46.3 80.7

percent 4.6 25.0 13.0 57.4 100
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 269

Aging and Mortality of Forests Stands

Aging of trees is a genetically determined process of weakening and disturbing

plants and their communities. As a rule, deterioration of old trees’ condition reveals itself through action of various factors such as climatic stresses, insect pests,

rots, etc. The age structure of stands in the ecoregions and administrative territories of Russia is extremely non-uniform (table 2). The forests of the European part of

the country are strongly depleted by logging, and the areas of the old-growth forests

in some regions are very small. The territories of Siberia and eastern Russian, on the

contrary, have a large portion ( > 50percent ) of mature and overmature forests. The forests

of the overmatured and weakened stands categories amount not less than 150 Mha.

As a result of succession processes, during the next century these forests will

be a strong source of carbon dioxide.

Fires

According to the national inventory data from 1988, the total area of burnt stands

in Russia is 27 Mha (Forest Fund of the USSR 1990). About 17 to 20 Mha of these

stands have burnt as a result of negligence of people. Statistical data do not include

the area of forests burnt by surface fires. This type of forest fire damages tree stands;

but more importantly, other compartments of ecosystems are also damaged, such

as understory, litter, and part of organic substances of soils. Natural reforestation of

young forest stands requires about 10 to 20 years. However, for recovery of other

components of ecosystems, for example lichen cover, 50 to 100 years or more are

necessary (Gorshkov 1994). This difference explains why the area of disturbed forest ecosystems is larger than the area of damaged tree stands. Main territory of

wild fires occurs in the Asian part of Russia, especially Yakutia and eastern Russian. We can expect that global climate warming will increase the number and areas of fires, burnt stands, and disturbed forest ecosystems (table 2).

Pests

Russia has 10 Mha of forests damaged by different pests (table 2; Isaev, pers. comm.).

In 1995 in Middle Siberia about 1 to 2 Mha of conifers stands were damaged by the

Asian gypsy moth alone.

Based on unfavorable age structure of Russian forests, namely, large areas of

overmature forests (table 1) we can predict increasing outbreaks of insects, fungi,

and other pathogens in the future.

Harvesting

As a result of harvest activity, Russia has about 25 Mha of disturbed tree stands and

about 100 Mha of recovering forest ecosystems (table 2). It is very difficult to predict

the rate of clearcutting, other types of fellings, and future damage of forests. This is

mostly because Russia is presently undergoing various, often unpredictable, political and economic changes.

Air Pollution

Area of stands damaged by air pollutants equals about 5 Mha (0.6 percent of total

stocked area) (table 2). About 15 Mha of forest ecosystems (2 percent of total stocked

area) have injured lichen cover. This relative damage is much better than in other

industrialized countries.

After Russia became a new independent country, atmospheric deposition of

air contaminants in Russia decreased during the last 4 years. This decrease is not

the result of improved industrial technology, but an economic depression and a

decrease in industrial production. Table 3 shows the dynamics of sulfur dioxide

emission from two industrial giant copper-nickel smelters in the European and

Asian parts of Russia.

Session IV Forest Health Status in Russia Alexeyev
270 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

It is well known that the weakening of plants and their communities by pollutants changes the consortive interrelations in ecosystems. In particular, it may stimulate outbreaks of forest insects. On the other hand, the combined damaging effects

of pollutants on the ecosystems already weakened by other anthropogenic or natural stresses are much stronger and faster than air pollution effects alone (Alexeyev

1995). This consideration is extremely important for the Russian forests because a

significant part of them are damaged by multiple stresses.

Hopefully, the impact of atmospheric pollution in the next century will be less

pronounced because of changing social awareness (Green Peace, and other ecological movements).

Conclusion

About one-third of tree stands and two-thirds of forest ecosystems of Russia are

disturbed by natural and anthropogenic factors. Most prevalent are natural causes

of disturbance; their impact affects 200 to 250 Mha of stands and 250 to 300 Mha of

forest ecosystems. Natural disturbance processes are an integral part of natural

forest life. Nevertheless, determinations of the size of damaged areas and trends of

the observed changes must take into account management strategies of the Federal

Forest Service of Russia. Not less than 45 Mha stands (6 percent of stocked area) are

under the impact of anthropogenic influence. Atmospheric pollution is the most

dangerous form of the anthropogenic stresses in forests in Russia.

Acknowledgments

I thank Laurie Dunn for technical editing of this manuscript.

References

Alexeyev, V.A. 1995. Impact of air pollution on far north forest vegetation. The Science of the Total Environment 160/161: Impact of air pollution on far north forest vegetation

605-617.

Forest Fund of Russia. 1995. Handbook. Moscow: Federal Forest Service of Russia, VNIIT Cresours; 280 p. [in Russian]. Handbook. Moscow: Federal Forest Service of Russia

Forest Fund of the USSR. 1990. Statistical handbook. Moscow: State Forestry Committee. Vol. 1; 1006 p. [in Russian]. Statistical handbook. Moscow: State Forestry Committee

Gorshkov, V.V. 1995. Characterization of recovery of forest ecosystems after fires. Reports of the Academy of Sciences in Characterization of recovery of forest ecosystems after fires

General Biology 333 (6): 811-813 [in Russian].

Idimechev, V.F.; Magdalina, V.V.; Shevchenko, T.F. 1995. Conditions of environment in Krasnoyarsk kray in 1994. Krasnoyarsk Conditions of environment in Krasnoyarsk kray in 1994

Report of Krasnoyarsk Environmental Committee; 92 p. [in Russian].

Poznyakov, V.Y. 1993. The Severonikel smelter complex: history of development. In: Kozlov, M. V.; Haukoja, E.; Yarmishko,V.T., The Severonikel smelter complex: history of development

eds. Aeral pollution in the Kola Peninsula: Proceedings of the international workshop; 1992 April 14-16; St. Petersburg:

Apatity; 16-19.

Vlasova, T.M.; Kovalev, B.I.; Filipchuk, A.N. 1991. Effects of point source atmospheric pollution on boreal forest vegetation of Effects of point source atmospheric pollution on boreal forest vegetation of

Northwestern Siberia. Northwestern Siberia. In: Weller, G.; Wilson, C.L.; Northwestern Siberia. Severin, B.A.B., eds. Proceedings of a conference on the role of the 

polar regions in global change. 1990 June 11-15; Fairbanks: University of Alaska. Vol.II; 423-428.

1 No data available.

Table 3 — Table 3 — Emissions of sulfur dioxide from Norilsk and Monchegorsk smelters (1,000 t/year) (Barkan, pers. comm.; Idimichev an Table 3 — d others 1995; Pozniakov 1993; Vlasova

and others 1991).

Smelters Years

1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994

Norilsk 2648 2724 2325 2244 2242 2216 2180 ND 1 ND 1863 1860

Monchegorsk 257 254 243 224 212 200 287 196 182 137 135

Session IV Forest Health Status in Russia Alexeyev
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 271

Abstract

Forests in the Ukraine are affected by environmental pollution, intensive forestry practice, and recreational

uses. These factors make them sensitive to impacts of climate change. Since 1989 Ukraine has participated in

the International Cooperative Program on Assessment and Monitoring of Air Pollution Effects on Forests

(ICP-Forests). A network of monitoring plots has been established in 14 administrative regions and in the

Crimea. From 1989 to 1995 the primary estimation criteria by the ICP Level 1 Program — defoliation and

discoloration of forest stands — have been observed on 215 plots. Data obtained by the ICP-Forests protocol

show that the recent decrease in air pollution was not followed by reduced forest defoliation. Thus, the estimation parameters established for that monitoring program are inadequate to determine accurate causeeffect relationships between forest health and external impacts. Defoliation and discoloration extended and

expanded with the support of the USDA Forest Service’s Forest Health Monitoring Program (FHM). More

intensive and more informative monitoring has been implemented, complementing activities on the primary

extensive monitoring network. Climate change impact may indicate pest damage, air pollution, fungi, or

weather anomalies. These causes are difficult to identify at the extensive monitoring level. In summer 1995,

climate change impact on the Ukrainian forests was estimated using the FHM protocols for forest vulnerability and adaptations assessment. Potential forests, main tree species distribution, and forest communities biodiversity was made estimated by using a geographical information system (GIS).

Introduction

Ukrainian forests cover 10 million hectares and about 14.3 percent of the total country. Forest distribution pattern, growth, and health are connected with temperate,

continental climate conditions, except for a narrow strip of subtropical coastal forest in the southern Crimea. Ukrainian forests are composed of about 54 percent

coniferous trees and 46 percent deciduous trees. The average annual growth increment is about 4 cubic meters per hectar. Managed plantations are about half of the

forested lands. Pinus sylvestris L. (about 35 percent of forest stands), Picea abies Dietr.

(16 percent), and Abies alba Mill. (3 percent) are the main forest- forming coniferous

species. Quercus spp. (22 percent), Fagus sylvatica L. (13 percent), and Carpinus

betularius L. (2 percent) are dominant hardwood species. Softwood genuses are Betula

spp., Populus spp., Alnus spp., and Tilia spp. (9 percent).

Ukrainian forests are considered valuable environment-forming factors. Among

the countries of the former Soviet Union, Ukrainian forests have been strongly affected by anthropogenic sources such as the high levels of air pollution. Overall

industrial emission of pollutants was about 17 million tons per year. The main

sources are ferrous metallurgical plants and the coal industry. Industrial emissions

consist of 21.4 percent particulate pollution, 30.2 percent SO2

, 8.0 percent NO and

NO2, 33.5 percent CO2, and 6.9 percent hydrocarbons. It is estimated that motor

vehicles add about 40 percent to the overall air pollution: more than 51 percent

CO2, 46 percent hydrocarbons, and 22 percent NO2 of the total volume of emissions

are caused by motor vehicles.

This paper discusses the spatial distribution and degree of anthropogenic damage to Ukrainian forests as determined by using the monitoring protocol of the

ICP-Forests and the FHM programs.

Air Pollution and Climate Change Effects on Health of the

Ukrainian Forests: Monitoring and Evaluation1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Forester, Biologist, Ecologist, and

Applied Mathematician, respectively,

Scientific Research Institute of

Forestry and Afforestation

(UkrSRIFA), Pushkinska Str. 86,

Ukraine 310024 Kharkiv E-mail:

buksha@ufri.kharkov.ua

Igor F. Buksha, Valentina L. Meshkova, Oleg M. Radchenko, Alexander S. Sidorov2
272 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Air Pollution

To evaluate the intensity of air pollution, we used generalized data on industrial

pollutant emissions from a network of plots (grid size 16x16 kilometers) in 18 administrative regions and big industrial cities in the Crimea Republic from 1985 to

1993 (fig. 1). Since 1985 until 1993, the total amount of pollutant emissions from

industrial sources decreased by 52.8 percent. Air pollution levels are expected to

continue decreasing with the industry recession.

To determine geographical zones that are homogenous in relation to air pollution, we used the cluster analysis method (Duran and Odell 1974). By using

Chebyshev’s metrics, data on the total amount of pollutant emissions, their sources,

and dynamics were plotted on a dendrogram reflecting the degree of similarity

between the territorial units (fig. 2). The dendrogram shows that all the administrative regions can be divided into two classes that differ by total intensity and composition of emission. Further K-means clustering (Duran and Odell 1974) was based

on the preliminary hypothesis of only two classes in the sample. The analysis shows

that the first class includes the Donetskaya region and parts of Luganskaya,

Zaporozhskaya, and Dnepropetrovskaya region (territorial units 3, 5, 7, 8, and 11).

The second class includes the remainder units. The first class shows a higher density level of pollution as indicated by emissions within that area and the impact of

stationary sources (figs. 3, 4). This resulting classification scheme is used to study

relationships between changes in forest health and atmospheric pollution.

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov

Figure 1 — International Cooperative Program on Assessment

and Monitoring of Air Pollution

Effects on Forests (ICP-Forests)

and Forest Health Monitoring

(FHM) plots.

Figure 2 — Differences between

geographical units by pollution.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 273

Assessment of Dynamics and Distribution

of Forest Damage in the Ukraine

In the Ukraine network, 215 forest monitoring plots have been established (fig. 1),

but some plots were not observed annually. There were 53 plots in 1989 (552 coniferous and 736 deciduous trees) and 134 in 1995 (1,467 coniferous and 1,743 deciduous trees). The monitoring is designed to identify zones of forest decline and to

study cause-effect relationships between changes in forest health and atmospheric

pollution.

Degree of crown defoliation in the stands is a common indicator of monitoring

that characterizes the forest condition. For a comparative analysis all data of observations are divisible by the following classes according to degree of crown defoliation: not defoliated, 0-10 percent; slightly defoliated, 11-25 percent; moderately

defoliated, 26-60 percent; severely defoliated, more than 61 percent. In accordance

with ICP Forests assessment criteria, defoliation less than 25 percent is not considered an indicator of worsening forest condition, since such values may be within

the range of natural fluctuation of tree phytomass (tables 1-4).

Figure 3 — Emissions by regions

with intensive air pollution

(Group 1).

Figure 4 — Emissions by regions

with low air pollution (Group 2).

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov Quantity of polutants, ths.t/sq.km

180.0

160.0

140.0

120.0

100.0

80.0

60.0

40.0

20.0

0.0

1985 1990 1993

Year

Mobile sources

Fixed sources

Quantity of polutants, ths.t/sq.km

20.0

18.0

16.0

14.0

12.0

10.0

8.0

6.0

4.0

2.0

0.0

1985 1990 1993

Year

Mobile sources

Fixed sources
274 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The results showed an increase in tree defoliation during the observational

period from 1990 to 1995 (figs. 5-7). Fraction of trees that defoliated to 25 percent

increased in 1990 to 1995 from 4 to 27 percent. Average yearly defoliation of the

forest stands increased from 9.5 to 22.6 percent. Maximum defoliation as high as 44

percent was found in the Luganskaya region in 1995; the minimum was about 5

percent in the Ternopol’skaya region in 1990. Average defoliation for the whole

monitoring period was highest in the Donetskaya (36 percent) and the Luganskaya

(28 percent) regions probably because of air pollutants emitted in the Kharkovskaya

and Rovenskaya industrial areas (fig. 8). The lowest defoliation levels for the whole

period were in the Ternopol’skaya (8 percent) and in the L’vovskaya (9 percent)

regions. The greatest defoliation was seen in 1994.

Comparative analysis of tree species was accomplished by assessing differences in defoliation levels by years. Neuman-Keuls post-hoc test (Murphy and others 1988) was used to obtain the matrix of the differences reliability. Further analysis related only species with reliable differences from other ones (P-level less than

0.05). Then species’ sets were divided into classes so that within a class there is no

species with reliable differences in defoliation levels.

Table 2 Table 2 — Defoliation of all species by 10 percent classes (1995). Table 2 

Species Total Tree quantity (pct) in 10 percent defoliation classes

trees 0-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100

All 3,210 23.6 1.9 3.6 13.0 3.7 1.8 1.0 0.6 0.5 0.3

Conifers 1,467 28.1 0.3 5.6 9.3 3.4 1.4 0.9 0.5 0.5 0.0

Broadleaves 1,743 19.8 3.3 1.9 16.1 3.9 2.2 1.0 0.7 0.5 0.6

Conifers

age to 59 years age equal or more than 60 years

Defolia Defolia- Defolia- Overall Overall

-tion -tion

class (pct) Pinus Juniperus Abies Picea Total Pinus Juniperus Picea Abies Other Total

sylvestris communis alba abies sylvestris comm- abies alba

unisabies

0 0-10 14.3 100 33.3 12.2 50.5 21.4 100 95.5 28.8 100 24.2 28.1

1 11-25 71.4 0.0 50.0 61.1 36.1 51.6 0.0 4.5 27.3 0.0 48.1 46.2

2 26-60 10.7 0.0 16.7 25.6 12.5 25.3 0.0 0.0 43.9 0.0 25.8 23.9

3 61-99 3.6 0.0 0.0 1.1 0.9 1.7 0.0 0.0 0.0 0.0 1.9 1.8

4 100 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Total 100 100 100 100 100 100 100 100 100 100 100 100

Table 3 Table 3 — Defoliation of conifers by age group and species. Table 3 

Table 1 Table 1 — Defoliation of all species by classes (1995). Table 1 

Percent of defoliation

 Total Total Class 0 Class 1 Class 2 Class 3 Class 4 Classes 2-4 Classes 1-4

 plots trees 0-10 11-25 26-60 61-99 100

134 3,210 23.6 46.8 28.1 1.5 0.0 29.6 59.3

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 275

Figure 5 — Defoliation of conifers from 1989-1995.

Figure 6 — Defoliation of

broadleaves from 1989-1995.

Figure 7 — Defoliation of all

species from 1989-1995.

Percentage

100

90

80

70

60

50

40

30

20

10

0 1989 1990 1991 1992 1993 1994 1995

Year

Defoliation:

more that 60 pct

26 - 60 pct

11 -25 pct

less than 11 pct –

–––––

Percentage

100

90

80

70

60

50

40

30

20

10

0 1989 1990 1991 1992 1993 1994 1995

Year

Defoliation:

more that 60 pct

26 - 60 pct

11 -25 pct

less than 11 pct –

–––––

Percentage

100

90

80

70

60

50

40

30

20

10

0 1989 1990 1991 1992 1993 1994 1995

Year

Defoliation:

more that 60 pct

26 - 60 pct

11 -25 pct

less than 11 pct –

–––––

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
276 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. Table 4 Table 4 — Defoliation of broadleaves by age group and species. Table 4 Broadleaves Defoliation Defoliation Age to 59 years Age equal or more then 60 years class percent percent Quercus Fagus Caprinus Other Total Quercus Quercus Caprinus Fagussylv Caprinus Other Total Overall Overall robur sylvatica betularius robur petrea betularius atica betularius 0 0-10 20.3 4.2 24.4 44.4 18.1 9.1 12.6 20.9 34.7 20.0 30.2 20.0 19.8 1 11-25 29.0 50.0 24.4 44.4 34.5 49.5 50.3 59.7 33.8 64.0 46.2 48.6 47.2 2 26-60 40.6 43.7 51.2 11.2 42.7 39.5 35.2 19.4 31.5 16.0 23.6 30.5 31.7 3 61-99 10.1 2.1 0.0 0.0 4.7 1.9 1.9 0.0 0.0 0.0 0.0 0.9 1.3 4 100 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Total 100 100 100 100 100 100 100 100 100 100 100 100 100

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 277

The data collected were used to group species by defoliation change rate by

using cluster analysis of average defoliation values:

● Group 1—Betula verrucosa, Ulmus spp., Quercus robur, Alnus incana,

Pinus sylvestris;

● Group 2—Fagus sylvatica, Quercus petrea, Acer tataricus, Acer

pseudoplatanus, Tilia cordata, Juniperus communis, Abies alba,

Fraxinus exelsior;

● Group 3—Betula pubescens, Fagus orientalis, Carpinus orientalis,

Carpinus betularius, Acer platanoides, Alnus glutinosa, Populus

tremula, Pinus pallasiana, Padus racemosa;

● Group 4—Quercus pubescens, Picea abies, Juniperus oxycedrus.

Group 1 had a relatively uniform defoliation growth rate (fig. 9). Its average

defoliation rate was greater than for any of the other groups (except for group 4 in

1993). Group 3 had the same pattern. However, the average defoliation rates for

group 3 was reliably lower than the rest. The defoliation rate of group 2 steadily

increased from 1989 to 1994, then noticeably decreased in 1995. Group 3 was

different from others due to the high defoliation peak in 1993. We expect the

cause of the peak was the particular climate conditions in 1993 and decreased

defoliation in 1994 and 1995. The measured pattern of defoliation changes in

the Ukraine forests showed that their photosynthetic function continues to

decline. Defoliation of group 1—consisting of main forest forming species,

Quercus robur and Pinus sylvestris—tended to steadily increase. Its correlation

coefficient for linear trend D=3.7279xT+2.4141 was 0.8989. To isolate causes of

defoliation change patterns will require longer time-series observations and

more adequate and complete data sets of measurements.

Figure 8 — The mean defoliation by regions in the Ukraine

from 1989-1995.

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
278 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The expected decrease of general defoliation has not occurred in spite of the

considerable decrease of air pollutant emissions since 1992 in the Ukraine. The possible causes of this phenomenon include:

● Affect of accumulation of pollutants in soil from previous years;

● Global climate changes;

● General increase of anthropogenic factors on forest ecosystems

(lumbercamp, pesticides and herbicides, recreation, drainage, and

irrigation, etc.);

● Advection of pollutants to Ukrainian forests from upwind industrial

countries.

In 1989, 82.9 percent of coniferous trees were not defoliated, 15.9 percent were

slightly defoliated, and 1.4 percent were moderately defoliated. In 1995 the percent

of non-defoliated trees fell (24.7 percent), and the percent of slightly and moderately defoliated trees increased (48.7 and 24.6 percent, respectively). In 1989, 84

percent of deciduous trees were not defoliated, 12.9 percent slightly, 3 percent moderately, and 0.1 percent were severely defoliated. In 1995 the portion of non-defoliated trees was the least (19.8 percent), and the portion of slightly, moderately, and

severely defoliated trees increased (49.3, 30, and 0.9 percent, respectively).

Among the coniferous trees, Pinus and Picea (Juniperus in Crimea) had the highest defoliation, while Quercus, Fagus, Caprinus, and Betula were the most defoliated

among the deciduous trees. However, the observed defoliation and discoloration

may be induced by pests, diseases, air pollution, or weather anomalies. In order to

isolate a real cause-effect relationship between forest conditions and external impacts, we used complex methods of analysis.

GIS Mapping

Sustainable forest management requires the development of methods for forest

health and productivity forecasting. Assessment of the forests’ vulnerability to global climate change is one of the generally recognized problems that needs such

forecasting. The research team from the Ukrainian Forestry Research Institute has

gained experience related to forests’ vulnerability assessment to global climate

change as a participant in the USDA Forest Service’s Forest Health Monitoring Program (FHM). The computer geographic information system (GIS) mapping of potential vegetation land cover was the primary approach to assessing the impact of

climate change on forests. Vegetation cover prognosis was based on Holdrige (1967)

Figure 9 — Defoliation of species groups from 1989-1995.

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 279

life zones and on Ukrainian forest land classifications (Holdrige 1967, Vorobjov

1960). Climate change scenarios GFDL, GISS, UKMO, and CCCM (IPCC 1990) from

general circulation models were used for projection with the classification models,

and maps of potential forest distribution were used to assess the general pattern of

climate change impacts. Large scale changes in biological productivity of forest

stands were estimated with climate-related productivity indexes (Anuchin 1977).

As an initial approach to describe future forest composition, the ranges of climatic

species were modelled to determine dominant tree species. Incorporation of forest

ecosystem models (gap-type models) provided more detailed forecasts for the most

vulnerable forest sites. However, gap-models are inadequate to assess conditions

in the Ukraine, and serious problems within the use of gap-models developed in

other regions. Ukrainian silviculture uses particular assessment indices for plant

response to climate change that are different from the gap-models requirements.

The methodological void in the assessment approach is between the large-scale

level of climate conditions and the “locally specific” gap-models that are difficult

to spatially extrapolate. Thus, methods for mid-scale impacts assessment were proposed by using the Ukrainian school of forest typology (Alekseyev 1929, Lavrinenko

1954, Pogrebniak 1968, Vorobjov 1960), which synthesizes forest-typological maps

from mid-scale maps of soils, hydrology, and relief (1:100,000 - 1:1,000,000).

There are numerous forest-typological classification schemes developed by the

Ukrainian school (Ostapenko 1978). They mainly differ by forest land classification

criteria used to determine soil humidity and trophic conditions. The latest research

involves the improvement of quantitative indices for the criteria and extension of

climatic and edaphic grid models onto representation of trophically poor, more

warm, and more dry sites. To make mid-scale maps of forest growth conditions

(e.g.,forest land maps, forest edaphic maps), forest typology schemes require synthesis of two maps: forest trophic conditions and soil moisture availability. Forest

trophic conditions may be derived from available soil maps: tables of correspondence can be established between soil types and trophic indexes compiled from

empirical data. However, a preferable for trophic index evaluation method is to

estimate the content of phosphorus and potassium in the root inhabited layer of

soil. The territorial units of the resulting mid-scale edaphic maps describe edaphic

conditions with some degree of generalization, depending on the scale and accuracy of source maps. However, the forest-typological meaning of such an edatope

is subject to study. Actually, adjustments of forest estimation methods to the “midscale” edatope are required. This methodology is applicable to the broad scale assessment of global climate change and damage caused by air pollution.

Discussion

This assessment of the relative changes of tree foliage suggests a progressive deterioration of crown condition from 1989 to 1994 followed by a slight improvement

in 1995. The highest level of defoliation was found among the main conifers, Pinus

spp. and Picea spp., and among the main broadleaves Quercus spp., Fagus spp.,

Carpinus spp., and Betula spp. Forests of the southeast region were the most affected, and the most damaged forests are located in the regions with higher pollution density. However, air pollution is not the only influence on forest health because climate conditions of this region are also not favorable to forest growth.

The set of estimation parameters used for the initial forest monitoring program

are not adequate for the determination of real cause-effect relationships between

forest conditions and external impacts in the Ukraine. These impacts are air pollution, pests, fungi diseases, unfavorable weather conditions, and intensification of

anthropogenic pressure on forest ecosystems. Study of the global climate change

influence on forests shows the attributive data composition that is necessary to

resolve this problem.

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
280 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

To obtain adequate forest ecosystem data, we must develop a multilevel monitoring system that includes extensive and intensive items. Since 1995 an extensive

monitoring system designed according to the ICP-Forests approach was extended

with the support of USDA Forest Service’s Forest Health Monitoring Program

(FHM). This observation program can provide data collection methods that may

be used for gap-model development and forestry statistics interpretation. In summer 1995 the Ukraine implemented these methods with the support of the FHM

program. A data base has been established to gather data on 32 plots. In addition,

more intensive programs to observe Ukrainian forest ecosystems have also been

prepared.

To effectively use these methods, they must include forest typology classification of forest growth conditions. This will ensure proper interpretation of the monitoring results and adequate assessment of forest condition dynamics affected by

various external factors, including global climate change.

Acknowledgments

We thank the reviewers for their help with the preparation of this manuscript. We

also thank Laurie Dunn for technical editing of this manuscript.

References

Alekseyev, Y.V. 1929. Silvics. Silvics. Kiev. (Russian). Silvics.

Anuchin, N.P. 1977. Forest inventory. Forest inventory. Moscow, “Lesnaya promishlennost”; 511 p. (Russian). Forest inventory. 

Duran, B. S.; Odell, P. 1974. Cluster analysis. Cluster analysis. New York: Springer-Verlag. Cluster analysis. 

Holdrige, L.R. Life zone ecology 1967. Life zone ecology 1967. San Jose, Costa Rica: Tropical Science Center. Life zone ecology 1967. 

IPCC (Intergovernmental Panel on Climate Change). 1990. Climate change: the IPCC scientific assessment. Climate change: the IPCC scientific assessment. Houghton eds., Climate change: the IPCC scientific assessment. 

New York: Cambridge University Press.

Lavrinenko, D.D. 1954. Forest types in Ukraine, Moscow, Lesbumizdat; 92 p. (Russian). Forest types in Ukraine, Moscow, Lesbumizdat; 92 p. (Russian).

Murphy, J.; Ridout, B.; McShane B. 1988. Numerical analysis, algorithms, and computation. Numerical analysis, algorithms, and computation. John Wiley and Sons; 146 p. Numerical analysis, algorithms, and computation. 

Ostapenko, B.F. 1978. Forestry-ecological typology and its classification. Forestry-ecological typology and its classification. In: Proceedings of the Kharkov Agriculture Institute. Forestry-ecological typology and its classification. 

Kharkov. (Russian).

Pogrebniak, P.S. 1968. Common sylviculture. Common sylviculture. Moscow, Colos; 439 p. (Russian). Common sylviculture. 

Vorobjov, D.V. 1960. The climate classification by forest types. The climate classification by forest types. In: Proceedings of the Kharkov Agriculture Institute. The climate classification by forest types. 

Kharkov. (Russian).

Session IV Air Pollution and Climate Change Effects on Health Buksha, Meshkova, Radchenko, Sidorov
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 281

Abstract

Systematic assessment and observation (survey, inventory) of forests in Latvia has been underway since the

1700’s. Latvia’s forests are in the boreal/temperate forest zone and cover 44 percent of the country. Forest

growing conditions are subdivided into five site class types: forests on dry mineral, wet mineral, wet peat,

drained mineral, drained peat soils. The dominant tree species in Latvia are Scots pine (Pinus sylvestris

L.), 39.7 percent; Norway spruce (Picea abies Karst.), 20.6 percent; and common birch (Betula pendula

Roth.), 28.4 percent. Systematic assessment of air pollution impacts on forest health in Latvia was started in

1990 by the International Cooperative Program on Assessment and Monitoring of Air Pollution Effects on

Forests - ICP Forests. Assessment methods were changed to the USDA Forest Service’s Forest Health Monitoring Program in 1994. Assessments of forest ecosystems in Latvia show the alteration and disappearance

of stand structure of typical, common forest ecosystems (especially Scots pine stands) , and the invasion of

plant species non-typical to these forests, which has been caused by human activities and air pollution

(sinantropization). It has also been found that Norway spruce stands will be endangered in the near future

because of their sensitivity to environmental factors, and that currently forest stand health is mainly affected

by localized entomological damage (insect and fungal diseases). In addition, environmental pollution has

caused biological destabilization in forest ecosystems.

Introduction

Latvia is one of three small Baltic countries transitioning to a free market economy.

Latvia occupies 64,000 km2, and 44 percent is covered by forests, which are of great

importance for the stability of the economy, landscape, environment, and flora and

fauna of Latvia. The first forest inventory in Latvia was carried out in 1788 for the

Lencu estate. Inventories of all the biggest estate forests were done until the beginning of the 1900’s. Then the process was periodically repeated after intervals of 10

to 15 years, including measurement and assessment of a forest stand and its health

condition. On this basis long-term forest changes, even for 100 years, were assessed.

During the late 1900’s, when environmental pollution significantly increased,

there was a necessity for more comprehensive forest assessment methods. In 1990

the European program, “International Cooperative Program on Assessment and

Monitoring of Air Pollution Effects on Forests (ICP Forests),” was implemented all

over the territory of Latvia. In 1994 establishment of sample plots for the Forest

Health Monitoring (FHM) program was started. This program is a component of

the United States’ Environmental Monitoring and Assessment Program (EMAP).

This paper discusses the current status of forests in Latvia as determined by

the newly implemented Forest Health Monitoring Program.

Forest Composition

The forests in Latvia are in the boreal/temperate forest zone. The total area covered

by forests has changed substantially over time. Agriculture lands freed from forests have been reforested as a result of various events such as wars or plague. The

productive forest area has changed from 25 percent in 1923 to 27 percent in 1949, 38

percent in 1973, and 44 percent 1995 (fig. 1).

Forest Condition in Latvia1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Chief Engineer, Latvian Institute of

Forest Inventory, Riga Street 113,

Salaspils, Latvia LV-2169.

Madis Sipols2
282 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Session IV Forest Condition in Latvia Sipols

Figure 1 — Forest cover in

Latvia by percent.

Forest growing conditions in Latvia are varied and include dry mineral soils

(58 percent), wet mineral (10.4 percent), wet peat (12.0 percent), drained mineral

(9.6 percent), and drained peat (10 percent). The main tree species in Latvia are

Scots pine (Pinus sylvestris) (39.7 percent), Norway spruce (Picea abies) (20.6 percent), common birch (28.4 percent), common alder (2.4 percent), aspen (2.5 percent), and grey alder (5.5 percent) (fig. 2). All forests are divided into three categories according to their functions and importance (ecological, economical, etc.):

● Protected forests (9.6 percent)

Nature reserves 38,700 ha

National park forests 51,600 ha

Nature park forests 15,000 ha

Nature protected forests 87,600 ha

Soil protected forests 72,200 ha

● Restricted management forests (16.3 percent)

Landscape forests 55,100 ha

Green zone forests 244,000 ha

Environment protection forests 225,700 ha

● Commercial forests (74.1 percent).

Scots pine occupies the greatest part of the forests in all three categories. Scots

pine is also widely represented through all soil types — from sandy to fertile soils,

as well as in peat soils. Scots pine is a biologically stable tree species, stable in

windstorms, and relatively less damaged by insects and fungi diseases. Compared

with Norway spruce, Scots pine is more resistant against various stresses and often

reaches 150 to 300 years. However, for the last 20 years the condition of pine

stands have been deteriorating. When the European system of forest monitoring was carried out from 1990 to 1995, Scots pine had a high level of defoliation for all age classes — 30.0 percent in 1990, 33.2 percent in 1992, and 24.1

percent in 1995 (table 1).

Forest Damage

The methodology of the FHM system is different from the previous systems and

uses more detailed condition indicators. About 40 percent of the planned sample

plots are established. The first results of this research show that the crown density

-<30 -30-40 -41-50

-51< -forestry
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 283

of conifers on average is less than the broadleaved species (figs. 3,4). Usually crown

damages are associated with different insects. For example, in 1995 high levels of

defoliation of Scots pine in the vicinity of Riga was associated with Gilpinia pallida

Kl. and Lymantria monacha L. The total damaged area there was about 10,000 ha. In

1992/93 Pissodes piniphilus Herbst caused serious crown damage on Scots pine.

At the same time invasion of new uncommon plant species in the forests on

dry sandy soils have been observed during the last 10 years. At several levels these

new species have become dominant. Scots pine stands undergo the process of

sinantropization, and the natural characteristics of these forests have changed and

disappeared.

The second most important tree species is Norway spruce, which is biologically unstable and very sensitive towards secondary factors such as insect damage

and fungus diseases. Currently, Norway spruce is seriously endangered. ICP Forests monitoring data from 1995 shows that 8 percent of all observed spruce are

damaged, and 70 percent are damaged by Ips typographus L.

Forest health is closely related to environmental changes (air pollution, climate,

urbanization, etc.). An important environmental quality indicator is the national

emission of SOx , NOx , CO, particulate pollution, and other substances. The total

emission amount of gases and particulate matter from 1991 to 1995 has decreased.

In 1991 the total emission of gases and solid substances was 144,100 tons/year; in

1992 — 92,310 tons/year; in 1993 — 87,130 tons/year; in 1994 — 103,840 tons/year;

and in 1995 — 86,440 tons/year (fig. 5).

Figure 2 — Distribution of Scots

pine (Pinus sylvestris L.) in Latvia.

Table 1— Changes in defoliation by most represented tree species (1990-1995). Table 1

Percentages of defoliation by mean age classes (years) ercentages of defoliation by mean age classes (years)

 Tree species 1990 1991 1992 1993 1994 1995

<60 >60 <60 >60 <60 >60 <60 >60 <60 >60 <60 >60

Scots pine 27.0 32.2 28.9 33.0 31.6 34.1 29.1 30.4 27.1 27.2 24.1 24.1

Norway spruce 12.9 23.8 14.8 20.5 15.6 21.1 15.3 20.3 14.3 19.4 13.6 18.3

Common birch 16.3 17.9 18.7 18.8 19.5 21.1 18.1 19.5 14.5 18.3 13.1 16.3

Common alder 10.6 7.7 14.1 9.0 14.3 11.5 14.1 12.2 11.6 10.3 13.4 9.6

Aspen 13.2 15.1 15.0 18.2 18.3 20.3 18.3 18.4 17.5 16.5 14.5 15.0

Session IV Forest Condition in Latvia Sipols

LEGEND

Average percentage share of pine stands

20 25 30 35 40 45 50 55 60 65 

TALSI

KULDIGA TUKUMS

SALDUS

DOBELE

RIGA

OGRE

BAUSKA

LIMBAZI

VALMIERA VALKA

AIZKRAUKLE

IBKABPILS

CBSIS

OULBENE

MADONA

BALVI

PREILI

DAUGAVPILS

REZEKNE LUDZA

KRASLAVA

ALUKSNE

VENTSPILS

TELGAVA

LIEPÁJA

'
284 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

LEGEND

DENSITY (pct)

26 - 35

36 - 45

46 - 55

56 - 65

66 - 75

pine isn't represented

(or is represented only with

intermediate, overtopped trees)

LEGEND

DENSITY (pct)

26 - 35

36 - 45

46 - 55

56 - 65

66 - 75

spruce isn't represented

(or is represented only with

intermediate, overtopped trees)

Figure 3 — Crown density of

Scots pine in Latvian forests (FHM

plots established in 1994, 1995).

Figure 4 — Crown density of

Norway spruce in Latvian forests

(FHM plots established in 1994,

1995).

Session IV Forest Condition in Latvia Sipols
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 285

Summary

The area covered by forests in Latvia has increased in recent years. Health of trees

is mainly affected by local entomological damages including cyclical repetition of

insect damages. During recent years intensity of insect damage has not reached the

catastrophic levels observed in earlier years. Environmental pollution impact on

ecosystems has also changed forest stand structure (especially in Scots pine stands)

and a process of biological destabilization has been observed

Acknowledgments

I thank Laurie Dunn for technical editing of this manuscript.

Figure 5 — Total emissions of

gases, solids, and other substances

(tons/year) in Latvia in 1995.

> 1500 

2600 - 3500 

districts 

 1600 - 2500

< 3500

Session IV Forest Condition in Latvia Sipols

USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 287

Abstract

In recent years, a forest monitoring program in the Czech Republic was extended into more detailed monitoring that aimed to describe the extent of changes in forest vitality and identify the nature and the main causes

of these changes on local and regional scales. Studies were undertaken in six mountain areas in the Czech

Republic. The program of regional forest monitoring is divided into three levels according to the extent of

evaluation of the parameters of forest stand health and other components of the forest ecosystem. Level 1 is

large scale monitoring in a 1 by 1 km grid of permanent plots. The total number of plots in a single regional

study varies from 60 to more than 500. The monitoring at level 1 plots includes a visual assessment of a

broad set of features of the health state of individual trees, repeated yearly. Assessment of health includes

measurement of tree diameter and height and a basic description of growing conditions. At monitoring level

2 the research assessment is extended to other parameters that characterize the forest stand and environment. The number of plots is usually 5-10 percent of level 1 plots. Monitoring level 3 includes analysis of the

processes of nutrient cycling. Detailed analysis of stand structure is done at the plots, including biomass

measurements. Results of field measurement are recorded into a database which allows a logical organization of a large amount of data and effective processing of them. Results of monitoring are analyzed using

statistical methods and modeling. A geographical information system (GIS) is used for further analyses and

for a final interpretation of results. From some studies, 4-5 years of results are now available. The studied

regions cover a broad range of conditions, making it possible to assess global trends in the health of Czech

forests.

Introduction

Forests in the Czech Republic cover an area of 2,642,064 ha. This corresponds to

33.4 percent of the total area of the Republic. Mean timber volume of forests is 225

m3/ha and mean annual increment is 6.91 m3/ha. Conifer species occupy 79 percent of forest area, and the main tree species is Norway spruce, occupying 54 percent of forest area. The primary forest regions of the Czech Republic are mountains

located mainly along the mountainous border (fig. 1) areas.

Forest Health Monitoring and Forestry Implications in the

Czech Republic1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Foresters, Institute of Forest Ecosystem Research Ltd., 254 01 Jilove u

Prahy 1544, Czech Republic.

Figure 1 — Forest coverage of

the Czech Republic, and location

of main mountain ranges (interpretation of LANDSAT TM image done by Stoklasa TECH,

Prague).

Martin Cerny, Pavel Moravcik2

Jizerské Mts.

Krkonose Mts.

Jeseniky Mts.

Beskydy Mts. Sumava Mts.

Brdy Mts.

Ore Mts.

 ́

 ́
288 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

The Czech Republic, particularly the north-western part (Ore Mountains) has been

strongly influenced by air pollution for the past several decades. A well-known “ecological disaster” began in the Ore Mountains in the 1950’s and 1960’s. Ten years later

severe problems started in the Jizerske Mountains and Krkonose Mountains. The significant decrease of land covered by forests is visible in these previously forested areas

(fig. 1); however, almost all deforested areas are now covered by young stands.

At the beginning of the 1980’s the symptoms of forest decline were also observed in northern Moravia (Beskydy and Jeseniky Mountains), and by the mid1980’s symptoms were observed in western and southern Bohemia (Brdy Mountains, Sumava Mountains, etc.).

At the same time the forests have been impacted by air pollution in some

areas, increased forest increment has been recorded in the Czech Republic based

on the data from regular forest inventories and data from permanent research plots.

Thus, it has been proved that during the past 100 years, the current height increment

of Norway spruce increased from 25 to 35 percent. On the other hand, the clear

differentiation of tree growth within individual forest stands related to the symptoms

of forest decline has been documented by data of tree ring analyses (fig. 2): the diameter

increment decreases with increase of health damage symptoms (e.g.,tree

defoliation). This example of the contradiction between forest decline and the

continuous increase of height increment of Norway Spruce in the mountains of

the Czech Republic clearly indicates the need for additional research.

This paper discusses the various forest health monitoring programs implemented in the Czech Republic that provide current data to forest policy makers for

the improvement of forest health.

Initial Conditions and

Forest Health Monitoring Programs

Forests of the Czech Republic are endangered by the continuous influence of a

range of stress factors. These factors disrupt forest ecosystems and cause the forest decline. The significance of potential stress factors is time and area specific.

Among the stress factors, sulfur dioxide is still important, although it has gradually decreased over the past years. Ozone, nitrogen oxides, and the influence of

potential climatic changes should also be considered. The impacts of complex stress

factors have not been sufficiently described.

The pan-European forest monitoring program (ICP Forests) has been implemented in most of the European countries to provide policy makers with basic

information on forest health. Within this program a standardized methodology

has been developed, and data exchange is supported. The Czech Republic has participated in the ICP Forest program since its establishment in 1986.

Session IV Forest Health Monitoring and Forestry Implications in the Czech Republic Cerny, Moravcik

Figure 2 — Relationship between tree damage (classes of

tree defoliation) and diameter increment. Index of diameter increment compares the current increment of the past 5 years to the

previous 5-year period.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 289

However, in the Czech Republic several systems of forest monitoring or research monitoring have been established over the past few decades. These systems

have similar scientific aims as the ICP Forest programs, but were designed independently with more or less different methodologies.

In addition to the forest monitoring program, the regular forest inventory is

used to estimate forest health status for practical forest management. Forest inventory data describing the whole area of the Czech Republic has been collected every

10 years. A planned national forest inventory program using a regular grid of permanent inventory plots will contribute additional information on forest health.

Several other monitoring programs addressing different environmental components

(air pollution, agricultural land, etc.) have also been implemented in the Czech

Republic under the direction of the Czech Ministry of Agriculture and the Czech

Ministry of Environment.

During the past two decades, remote sensing technology has improved significantly. With the support of terrestrial monitoring, remote sensing can serve as

a fast and efficient source of information on forests health status. However, because the level of available information is limited to descriptive information on

several stand characteristics, it cannot be used to determine causal relationships.

Forest management practice lacks information that is necessary for ecosystem

oriented management on local and regional levels. Forest monitoring should serve

as a permanent source of continuous information enabling forest managers to evaluate results of management procedures on a regional level and evaluate and reorient forestry policy on the national level.

A system of forest monitoring should be developed to both follow changes

in forests, and to identify main potential stress factors. On the basis of the correlative analysis of forest state parameters and characteristics of the growing

environment, hypotheses can be derived and then tested by the detailed research on ecosystem mechanisms.

Objectives of Forest Monitoring

Forest monitoring is defined as the long-term investigation of the state of the forest

and growing environment by using a set of selected parameters that allow evaluation

of forest changes caused by environmental changes and forest management practices.

The primary aims of forest monitoring programs are to clarify the extent of

changes in forests in the Czech Republic, describe the character of these changes,

and estimate the main causes of changes. The forest monitoring program should

also be a unified and internationally compatible system.

Structure of Forest Monitoring

The forest monitoring program in the Czech Republic is differentiated into separate subprograms according to the topics addressed within the program (table 1).

● Permanent Sample Plots: The set of permanent sample plots was established in the 1960’s to collect data on the growth of stands of the main tree

species found in the Czech Republic. These data were collected to develop regional growth and yield tables. Most of the plots are still active and produce

valuable information on the development of the forest ecosystem.

● Regional Forest Monitoring: Regional forest monitoring is derived from

the European program of forest monitoring. For the purposes of local forest

managers, and regional authorities in forestry and the environment, the information from the sparse European grid of monitoring plots (16 x 16 km) is not

satisfactory. For that reason a dense grid, 1 x 1 km, has been established in

selected regions of the Czech Republic. Currently, there are five such regions,

and several additional smaller areas are also covered by using this approach.

Session IV Forest Health Monitoring and Forestry Implications in the Czech Republic Cerny, Moravcik
290 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

● Large Scale Forest Inventory: Large scale forest inventory by using a terrestrial survey in the grid of permanent inventory plots is one of the main tools

for collecting data about forests and forest production. Recent developments

in large scale forest inventory methodology allow the evaluation of forest ecosystem status including biodiversity.

● ICP Forests: The Czech Republic has participated in the program of European monitoring (ICP Forests) since its beginning in 1986. The data collected in

the 16 x 16 km grid are used for the purposes of European forest policy.

● Remote Sensing: Satellite imagery from LANDSAT TM data is widely

used for the estimation of forest health status.

Forest Monitoring Projects in the Czech

Republic

The forest monitoring program in the Czech Republic is coordinated by the Czech

Ministry of Agriculture and Ministry of Environment (table 2). Except for ICP Forests, all monitoring projects are funded by grants.

Methodology of Terrestrial Forest Monitoring

The methodology of terrestrial forest monitoring has been derived from European

unified methodology for ICP Forests (table 3). The methodology is split into three

levels. Level one is applied to all monitoring plots. Level two is used in about 10

percent of the monitoring plots, and separation into level two is defined by the

additional cost of methodological procedures.

Table 1 Table 1 — Different programs of forest monitoring in the Czech Republic. Table 1 

Size of Size of Intensity of Intensity of Interval Interval Application

 Program Subject Subject Characteristics area unit area unit research (years) (years) of results

Permanent forest forest health, - medium to 1-5 cause-effects studies,

sample plots. ecosystem. environment, high modeling, support of

production, decisions on conceptual level.

forest structure.

Regional site, forest health, small high 1 detailed characteristics of

monitoring. region. environment, selected region, planning

production. on the regional level,

forest improvement

measures.

Large scale region, structure of medium medium 5-10 large scale forest

inventory. country. forests, to small inventory, forestry

production, planning, management

ownership, politics.

quality of

management,

forest health.

ICP Forests. country. forest health, large medium 1 current state of forest

environment, health on country level.

production.

Remote country, forest health, small low 1 and current information on

sensing. region, changes in forest more state of forest.

site. coverage.

Session IV Forest Health Monitoring and Forestry Implications in the Czech Republic Cerny, Moravcik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 291

Level three includes procedures that evaluate the ecosystem nutrient cycle and

is performed to characterize growing conditions of the forests. The data of field

assessments are stored in the MONitoring dataBASE. Version 5 of MONBASE allows efficient computer storage of data as well as implementation of the set of basic

processing routines. Pre-processed data are further analyzed and evaluated by using GIS techniques. Point layers of terrestrial monitoring data are combined with

interpretation layers (e.g., interpreted satellite images, forest soil maps, forest types

maps, and climate maps). By using GIS techniques like co-kriging and multicriteria

analysis, the monitoring data are processed and interpreted for further use in forest management practice.

Conclusions

The forest monitoring program in the Czech Republic has significantly developed

during the past 5 years. The primary features of the program include a general

monitoring approach that is performed in cooperation with other research programs on permanent research plots in the Czech Republic (forest production research plots, forest inventory). Forest monitoring data are interpreted for use by

forest managers, and the emphasis is on developing cause-effect relationships.

The unified methodology of the program uses three levels of research monitoring (up to ecosystem level) and extends to cause-effect relationships. The regional

forest monitoring network uses permanent research plots (including sites that are

less common but important ecologically) and has been extended to several new

regions.

The database used for forest monitoring is MONBASE version 5, which is a

flexible tool for handling monitoring data. Progressive methods of data processing

and evaluation include statistics, GIS, and modeling. Forest damage and production data are interpreted by decision makers and forest policy makers at both national and regional levels. Forests are then zoned according to the level of risk (as a

result of multicriteria analysis), and they are managed on the basis of nutrition and

species composition. By understanding these forest ecosystem processes, policy

makers can apply the monitoring results in forest production research for better

management of the Czech forests.

1 Institute of Forest Ecosystem Research, Jilove u Prahy

2 Forestry and Game Management Research Institute, Jiloviste-Strnady

3 Planned for 1999.

4 Development of methodology phase

Table 2 Table 2 — Terrestrial forest monitoring network in the Czech Republic. Table 2 

Forest area Grid Number of Number of Level of Level of Time of project Time of project Run by

 Project Project covered, ha km x km plots monitoring execution

National monitoring network

Permanent sample plots 2,650,000 non regular 900 I, II 1980+ IFER 1

ICP Forests 2,650,000 16 x 16 126 I, (II) 1986+ FGMRI 2

Large scale forest inventory 2,650,000 1.41 x 1.41 13,000 I, (II) ( 3 ) (IFER) 4

Regional monitoring network

Brdy Mts. 23,000 1 x 1 220 I, II, III 1989+ IFER

Sumava Mts. 60,000 1 x 1 528 I, II, III 1990+ IFER

Krkonose Mts. 30,000 1 x 1 262 I, II, III 1991+ IFER

Beskydy Mts. 45,000 1 x 1, 4 x 4 176 I, II, III 1992+ IFER

Jizerske Mts. 20,000 1 x 1 120 I 1992-93 IFER

Other local projects 25,000 1 x 1 55 I 1992+ IFER

Session IV Forest Health Monitoring and Forestry Implications in the Czech Republic Cerny, Moravcik
292 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Table 3 Table 3 — Methodological elements for different terrestrial forest monitoring levels. Table 3 

Level of Level of

 monitoring Element of methodology Extent Extent Interval Interval

I Level one: Level one:

basic descriptive data plot once

humus forms plot 6-12 yrs

soil chemistry plot 12 yrs

needle/leaf chemistry plot 6 yrs

phytocenological description plot 10 yrs

natural afforestation plot 3 yrs

occurrence of lichens plot 1 yr

tree species trees (all) 1 yr

tree class trees (all) 1 yr

mortality of trees trees (all) 1 yr

occurrence of crown/stem break trees (all) 1 yr

defoliation of whole crown trees (all) 1 yr

defoliation of upper third of crown trees (all) 1 yr

vitality of the crown tip trees (all) 1 yr

occurrence of dead branches trees (all) 1 yr

mechanical injury to the stem trees (all) 1 yr

type of color changes (discoloration) trees (all) 1 yr

intensity of color changes trees (all) 1 yr

diameter at breast height trees (all) 3 yrs

tree height trees (10-15) 3 yrs

occurrence of secondary shoots trees (5-10) 1 yr

needle retention trees (5-10) 1 yr

occurrence of combs trees (5-10) 1 yr

crown ratio trees (5-10) 1 yr

angle of branching trees (5-10) 1 yr

type of branching trees (5-10) 1 yr

injury caused by insects trees (all) 1 yr

II Level one plus: Level one plus:

humus forms plot 3 yrs

soil chemistry plot 6 yrs

soil profile description & chemistry plot 12 yrs

needle/leaf chemistry plot 3 yrs

small root vitality plot 6 yrs

core analyses trees (10) once

LAI measurement plot 1 yr

III Level two plus: Level two plus:

soil solution chemistry plot 2 weeks

wet/dry deposition chemistry plot 2 weeks

amount & chemistry of litter plot 1 month

air pollution/passive samplers plot 2 weeks

cont. measure. of tree diameter incr. trees (10) 1 month

detail stand structure description plot 10 yrs

Session IV Forest Health Monitoring and Forestry Implications in the Czech Republic Cerny, Moravcik
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 293

Abstract

Slovakia is a mountainous and forested country (40.6 percent forest cover) in central Europe and has a large

variety of vegetation zones, forest types, and a rich diversity of forest tree species. The most important tree

species are beech (Fagus sylvatica L.), Norway spruce (Picea abies Karst.), oak species (Quercus sp.), Scots

pine (Pinus sylvestris L.), silver fir (Abies alba Mill.), European hornbeam (Carpinus betulus L.), European larch (Larix decidua Mill.), poplars and willows (Populus L. sp., Salix L. sp.), and other hardwood

broadleaves. On the basis of results of the Forest Health Monitoring System from 1987 to 1994, the scientific

information is presented for the following parameters: defoliation, discoloration, percentage of tree number

in classes of damage, and percentage of salvage cut in the total annual cut. The percentage of trees in the

defoliation classes 0, 1, 2, 3, and 4 were 14.7, 43.5, 36.2, 4.3, and 1.3, respectively (class 0 means healthy

trees; class 4 dead trees). The discoloration classes (0, 1, 2, 3, and 4) were represented by 98.0, 1.7, 0.3, 0, and

0 percent for all tree species together. The average defoliation percentage from 1987 to 1994 decreased with

time and is expected to further decrease in the following years. However, the percentage of annual salvage cut

in the total annual cut increased between 1987 and 1994. Various biotic and abiotic factors influence forest

health in Slovakia, such as air pollution, wind and snow, damages by beetles, sucking insects, and game.

Global climate change seems to be the most important among them.

Introduction

Forest health status is a phenomenon studied and monitored in central and eastern

Europe, where the natural vegetation cover consists of forests and where forestry is

an important part of the economy. Numerous scientific and scientific-technical

projects have been involved in the process to describe the main factors influencing

the health status of forest trees and to propose measures that could partially solve

the problem of forest decline.

Because Slovakia is a mountainous and forested country (about 1.94 million

ha. of forest, i.e., 40.6 percent of the total area), it is important to study changes in

forest health status. Because of the mountainous character of the country, forested

habitats in Slovakia encompass a wide range of environmental conditions and

elevations (table 1). Trees are subjected to many natural stresses, including those

induced by changes in water and nutrient relations, light, temperature, and biotic

factors. Because of their longevity, trees usually react to the changes abruptly, and

very often they are not able to adapt themselves to the changed growing conditions.

The dominant forest types in Slovakia are the beech (Fagetum pauer, Fagetum

typicum), beech-oak (Fageto-Quercetum), and oak-beech (Querceto-Fagetum, Fagetum

dealpinum) types, which together cover about 52 percent of the total forest area.

Beech-fir (Fageto-Abietinum), fir-beech (Abieto-Fagetum), and fir-beech with spruce

(Fagetum abietum - piceosum) forests cover an additional 25 percent. Other locally

important forest types include pine-oak and oak-pine (Pineto-Quercetum and

Querceto-Pinetum, 1.96 percent), Cornel oak forests (Corneto-Quercetum, 0.91 percent),

oak-ash forests (Querceto-Fraxinetum, 0.41 percent), elm-ash forest (Ulmeto-Fraxinetum

populeum and Ulmeto-Fraxinetum carpineum, 1.07 percent), hornbeam-oak forest

(Carpineto-Quercetum, 6.99 percent), acid oak-beech forest (Fagetum quercinum, 2.69

percent), linden-maple forest (Tilieto-Aceretum, 0.63 percent), ash-maple forest

(Fraxineto-Aceretum, 2.28 percent), dealpine pine forest (Pinetum dealpinum, 0.96

percent), fir-spruce forest (Abieto Piceetum, 2.84 percent), mountain ash-spruce forest

Dynamics of Forest Health Status in Slovakia from

1987 to 19941

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Forestry Ecologist, Institute of Landscape Ecology, Slovak Academy of Sciences, P.O. Box 254, Stefanikova St.

No. 3, 814 99 Bratislava, Slovakia.

Julius Oszlanyi2
294 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

(1.02 percent), maple-spruce forest (Acereto-Piceetum, 0.53 percent), and dwarf pine

forest (Mughetum, 0.60 percent).

According to the newest forest inventory results (Novotny and others 1994),

the tree species percentage in the Slovakian forests include:

Species Percent

Norway spruce (Picea abies Karst.) 27.2

European larch (Larix decidua Mill.) 2.0

Silver fir (Abies alba Mill.) 5.0

Scots pine (Pinus sylvestris L.) 7.0

Oaks (Quercus L. sp.) 14.3

Beech (Fagus sylvatica L.) 29.8

European hornbeam (Carpinus betulus L.) 5.6

Other hardwood broadleaves 4.5

Poplars and willows (Populus L. sp., Salix L. sp.) 3.5

Slovakia is situated in the central European region where prevailing northwest

winds bring pollutants from the industrial areas in the Czech Republic, Poland,

and Germany. Air pollution effects on forests have become critical in the past decades in Slovakia. Other causal factors affecting forest health in Slovakia include

soil pollution, wind damage, snow damage, damage by bark-beetles, damage by

wood-destroying beetles, damage by phytophagous and sucking insects, browsing

damage, and bark peeling (game) damage.

Like most other European countries, Slovakia began a forest tree health monitoring program, which started in 1987 and continued through summer 1995. The

ninth monitoring cycle concentrated on the permanent network of forest health

monitoring plots (16 by 16 km grid). According to the methods used in the European forest health monitoring system, the permanent monitoring plots, which have

no specific size, contain 15 trees close to the center of the plot and are used for

measurement and observations.

This paper discusses the results of the forest health monitoring system in

Slovakia that studied these parameters: tree stem circumference at the height of

1.3 m above the soil surface (the diameter at breast height calculated—dbh);

biosociological position of trees (Kraft tree classes scale); loss of assimilation apparatus (defoliation); heights of three trees (by Blume-Leiss hypsometer); fertility of

trees; mechanical damage on trees; fungi; and insects.

Session IV Dynamics of Forest Health Status in Slovakia from 1987 to 1994 Oszlanyi

Table 1— General survey of forested habitats divided into vegetation zones. Table 1

Vegetation Altitude Mean annual Mean annual Annual Vegetation

 zone (m a.s.l.) (m a.s.l.) temp. ( temp. (°C) precip. (mm) precip. (mm) period (days) period (days)

Oak <350 >8.0 <600 >165

Oak beech 350-400 7.5-8.0 600-650 160-165

Beech-oak 400-550 6.5-7.5 650-700 150-160

Beech 550-600 6.0-6.5 700-800 140-150

Fir-beech 600-700 5.5-6.0 800-900 130-140

Spruce-beech 700-800 4.5-5.5 900-1,500 115-130

Beech-spruce 900-1,050 4.0-4.5 1,050-1,200 100-115

Spruce 1,050-1,350 2.5-4.0 1,200-1,500 60-100

Dwarf pine >1,350 <2.5 >1,500 <60
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 295

Table 2 Table 2 — Tree species percentage in defoliation degrees (1994) (Novotny and others 1995). Table 2 

Degree of Defoliation

0 1 2 3 4 1+2+3+4 2+3+4

Defoliation (percent) Defoliation (percent)

Tree species 0-10 11-25 26-60 61-95 100

Fagus sylvatica L. 27.5 47.8 23.6 1.0 1.0 72.5 24.7

Quercus sp. L. 1.5 53.9 40.0 2.3 2.3 98.5 44.6

Carpinus betulus L. 30.8 45.7 22.9 0.3 0.3 69.2 23.5

Other broadleaf 3.2 23.0 52.9 16.9 4.0 96.8 73.8

Broadleaf total 19.3 45.1 31.0 3.5 1.1 80.7 35.6

Picea abies Karst. 8.4 41.9 43.6 4.9 1.2 91.6 49.7

Abies alba Mill. 15.6 36.0 37.6 8.6 2.2 84.4 48.4

Pinus sylvestris L. 4.6 40.5 48.1 4.8 2.0 95.4 54.9

Larix decidua Mill. 5.1 57.6 28.8 6.8 1.7 94.9 37.3

Coniferous total 8.2 41.5 43.5 5.3 1.5 91.8 50.3

Total 14.7 43.5 36.2 4.3 1.3 85.3 41.8

Methods

The numerous data and information obtained from the network of forest health

monitoring plots (monitoring was performed August 11-20) provide excellent material for studies. Results that concentrated on the most visible changes on the forest trees document a more complicated system of causal factors responsible for the

health of the forests. The variety of anthropogenic causal factor complexes have

interacted with natural events and processes that induce stresses in forests and can

be observed as ecological response.

Numerous papers (Lapin and others 1994, Lapin and others 1995, Lapin 1995)

give information on trends of means of temperature and precipitation in different

parts of Slovakia, starting in the year 1870. Results from localities situated in the

lowland of south Slovakia and in the mountains in the north document a

positive tendency in mean annual temperature and negative tendency in

precipitation totals and in relative air humidity.

Results

Defoliation and discoloration of foliage are considered the best and most significant indicators of the health status of forest trees (tables 2-6). Both scientists and

forestry managers are convinced that defoliation and discoloration are visible responses of the internal and external influences on the life of the individual tree

(Novotny and others 1995).

For the evaluation we used the 5 degree scale of defoliation (Novotny and others 1995) to maintain consistency with data from other European countries. But in

the Slovak Republic, narrower defoliation classes (each class is 10 percent) were

used because this provides greater precision for the ecological evaluation of forest

health and its dynamics in a longer period.

The internationally accepted scale denotes degree of damage (0 to 4) in terms

of percent of foliage loss (0 to 100 percent):

0 - healthy trees (without symptoms), 0-10 percent

1 - partially ill trees, 11-25 percent

2 - ill trees, 26-60 percent

3 - very ill trees, 61-95 percent

4 - dead trees, 100 percent.

Session IV Dynamics of Forest Health Status in Slovakia from 1987 to 1994 Oszlanyi
296 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

When the data on defoliation are put into a regression, the negative relationship is obvious. Decrease of defoliation percentage from 1987 to 1994 is related to

the changes in the economy of the central European countries: the decrease in industrial activity has resulted in decreased pollution. Air and soil pollution has been

responsible for the defoliation and discoloration of trees.

Defoliation is not the only and not the most significant indicator of health state

of the forest trees; the salvage cut percentage is also a dynamic indicator. Despite of

the fact, that the defoliation has the negative tendency in the studied years, the

salvage cut percentage has a positive tendency (table 7). There are different causal

factors responsible for the salvage cut amount in different years. Among them, the

most important are the climate changes that seriously influence the growing conditions at each forest site and subsequently forest health and the stability of forest

ecosystems.

The data concerning the degree of defoliation show a trend that is controversial to the trends of forest health status in Slovakia. The salvage cut and its percentage of the total cut is the most appropriate and useful indicator of the forest health

status in Slovakia. The salvage cut quantifies the amount of the trees (wood) that

had to be cut because of biological and abiological damage. Salvage cut quantifies

the ecological instability of part of the forest and the bad health status of the forests. It also signifies the decline of forest health status because the salvage cut percentage is increasing permanently.

The annual cut in Slovakia has been relatively stable (average annual cut

from 1991 to 1992 was 4.24 million m3

), but the increasing salvage cut documents the necessity of cutting dead, weakened , and damaged trees in increasing volume (table 7).

Discussion and Conclusion

As one of the most forested European countries, Slovakia is characterized by forests containing richness of tree species, ecosystems, habitats, and site diversity.

However, the data from the monitoring plots network give us very interesting scientific-technical information that documents the worsening state of forest health,

with negative prognosis for the future. Although the importance of the causal factors varies in the respective years, the tendency is clear, and it is negative.

Table 3 Table 3 — Tree species percentage in discoloration degrees (1994) (Novotny and others 1995). Table 3 

Degree of discoloration

0 1 2 3 4 1+2+3+4 2+3+4

Discoloration (percent) Discoloration (percent)

Tree species 0-10 11-25 26-60 61-95 100

Fagus sylvatica L. 99.6 0.3 0.0 0.1 0 0.4 0.1

Quercus sp.L. 97.1 1.9 0.6 0.2 0.2 2.9 1.0

Carpinus betulus L. 96.5 3.2 0.3 0 0 3.5 0.3

Other broadleaf 97.1 2.4 0.5 0 0 1.9 0.5

Broadleaf total 98.4 1.3 0.2 0 0.10 1.6 0.3

Picea abies Karst. 99.3 0.3 0.4 0 0 0.7 0.4

Abies alba Mill. 100.0 0 0 0 0 0.0 0.0

Pinus sylvestris L. 90.6 8.9 0.5 0 0 9.4 0.5

Larix decidua Mill. 100.0 0 0 0 0 0.0 0.0

Coniferous total 97.5 2.1 0.4 0 0 2.5 0.4

Total 98.0 1.7 0.3 0 0 2.0 0.3

Session IV Dynamics of Forest Health Status in Slovakia from 1987 to 1994 Oszlanyi
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 297

Table 4 Table 4 — Tree species percentage in defoliation classes (1994) (Novotny and others 1995). Table 4 

Defoliation (percent) Defoliation (percent)

 Tree species 0-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100

Fagus sylvatica L. 27.5 28.6 31.9 6.3 3.4 1.3 0.7 0.2 0.0 0.1

Quercus sp. L. 1.5 24.3 52.7 12.8 3.1 1.0 1.3 0.2 0.2 2.9

Carpinus betulus L. 30.8 25.5 37.1 5.7 0.3 0.0 0.3 0.0 0.0 0.3

Other broadleaf 3.2 10.7 26.9 20.1 12.7 5.5 6.9 6.1 3.7 4.2

Broadleaf total 19.3 24.6 35.8 9.5 4.4 1.7 1.7 1.1 0.6 1.3

Picea abies Karst. 8.4 24.8 30.7 16.1 9.2 4.7 2.8 1.3 0.7 1.3

Abies alba Mill. 15.6 25.8 18.8 12.9 10.2 5.9 4.8 2.2 1.1 2.7

Pinus sylvestris L. 4.6 23.7 37.7 18.3 6.1 2.8 1.8 2.0 1.0 2.0

Larix decidua Mill. 5.1 28.7 39.0 11.9 6.8 0.0 3.4 3.4 0.0 1.7

Coniferous total 8.2 24.7 31.3 16.1 8.6 4.3 2.8 1.6 0.8 1.6

Total 14.7 24.6 33.9 12.3 6.1 2.8 2.2 1.3 0.7 1.4

Defoliation and discoloration are not the only and the most significant indicators of forest health. The tendency of defoliation is negative because the trees lose

fewer and fewer leaves by August 15 (their “health state” is getting better), but the

other important indicators show the opposite tendencies. For instance, the percentage of the annual salvage cut in the total annual cut is increasing (i.e., the stability

of the forest is decreasing).

Among other causal factors, partially influencing the health state are pollution,

wind and snow damage, damage by beetles, sucking insects, and game. The most

important factor is climate change observed in Slovakia.

To evaluate forest health status, defoliation and discoloration of foliage must

be always combined with other parameters, especially the salvage cut percentage,

which is the most complete indicator of health of forest trees and of ecological stability of forests.

Table 5 Table 5 — Average defoliation percentage of tree species (1987-1994) (Novotny and others 1995). Table 5 

Tree species Average defoliation (percent) Average defoliation (percent)

1987 1988 1989 1990 1991 1992 1993 1994

Fagus sylvatica L. 23 19 23 17 13 17 17 21

Quercus sp. L. 24 30 35 31 25 27 27 30

Carpinus betulus L. 18 14 20 18 13 18 25 20

Fraxinus excelsior L. 29 23 29 38 40 38 30 40

Acer sp. L. 39 35 46 39 33 30 29 32

Robinia pseudoacacia L. 32 37 38 74 46 61 51 57

Populus sp. L. 26 40 37 38 45 50 32 36

Broadleaf total 24 23 27 25 19 23 23 26

Picea abies Karst. 34 28 31 29 25 27 29 32

Abies alba Mill. 52 31 39 37 31 33 32 33

Pinus sylvestris L. 40 45 44 44 33 42 29 32

Larix decidua Mill. 24 20 33 33 17 26 27 30

Coniferous total 35 32 35 33 27 29 29 32

Total 30 27 30 28 23 26 26 28

Session IV Dynamics of Forest Health Status in Slovakia from 1987 to 1994 Oszlanyi
298 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

References

Lapin, M., 1995. Assessment of the Slovak Republic. Vulnerability to climate change and adaptive strategies design. Journal Assessment of the Slovak Republic. Vulnerability to climate change and adaptive strategies design

of Hydrology and Hydromechanics 43 (4-5): 354-370.

Lapin, M.; Fasko, P.; Zeman, V. 1994. Contribution to analysis of possible global warming impact upon climate change in

Slovakia. National Climate Program of the Slovak Republic, I (2): 37-77 (in Slovak). Slovakia. National Climate Program of the Slovak Republic

Lapin, M.; Nieplova, E.; Fasko, P. 1995. Climate change scenarios for air temperature and precipitation change in Slovakia. Climate change scenarios for air temperature and precipitation change in Slovakia

National Climate Programme of the Slovak Republic, II (3): 19-57 (in Slovak).

Novotny, J.; Fillo, E.; Gregus, C.; Hell, P.; Hoffmann, J.; Ilavsky, J.; Kolenka, I.; Konopka, J.; Korpel, S.; Paule, L.; Paulenka, J.;

Racko, J.; Stroffek, O. 1994. Forests of Slovakia. Poly Kontakt; 89 p. (in Slovak). Forests of Slovakia

Novotny, J.; Brutovsky, D.; Findo, S.; Mankovska, B.; Paulenka, J.; Leontovyc, R.; Pajtik, J.; Turcani, M. 1995. Occurrence of Occurrence of

damages in forests of Slovakia in 1994 and their prognosis for 1995. Forestry Research Institute Zvolen; 148 p. damages in forests of Slovakia in 1994 and their prognosis for 1995

Report of the Forestry Research Institute Zvolen. Research Station Banska Stiavnica; 148 p. (in Slovak). Report of the Forestry Research Institute Zvolen

Table 6 — Percentage of all tree species in respective damage degrees from 1987 to 1994 in the Slovak Republic (Novotny Table 6

and others 1995).

Year and Tree Species Percentage of tree number in damage degrees

0 1 2 3 4 1-4 2-4 3-4

1987 Coniferous 11 36 41 11 1 89 53 12

Broadleaf 26 47 22 5 0 74 27 5

Total 19 42 32 7 0 81 39 7

1988 Coniferous 14 33 43 9 1 86 53 10

Broadleaf 33 39 23 5 0 67 28 5

Total 25 36 32 6 1 75 39 7

1989 Coniferous 9 32 49 9 1 91 59 10

Broadleaf 20 38 37 4 1 80 42 5

Total 15 36 42 6 1 85 49 7

 1990 Coniferous 14 30 47 8 1 86 56 9

Broadleaf 23 45 25 5 2 77 32 7

Total 20 29 34 6 1 80 41 7

1991 Coniferous 14 47 34 4 1 86 39 5

 Broadleaf 41 38 17 3 1 59 21 4

 Total 30 42 24 3 1 70 28 4

1992 Coniferous 15 44 33 7 1 85 41 8

Broadleaf 31 40 23 5 1 69 29 6

Total 24 42 27 6 1 76 34 7

1993 Coniferous 8 42 46 3 1 92 50 4

Broadleaf 28 43 25 3 1 72 28 4

Total 20 43 33 3 1 80 37 4

1994 Coniferous 8 41 44 5 2 92 51 7

Broadleaf 20 45 31 4 1 80 36 5

Total 15 43 36 5 1 85 42 6

Table 7 Table 7 — Percentage of the annual salvage cut in the total annual cut in 1985 (Novotny and others 1995). Table 7 

Year 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994

Percent of

salvage cut 29.7 31.1 31.9 35.9 41.5 49.2 42.8 43.2 51.7 59.6

Session IV Dynamics of Forest Health Status in Slovakia from 1987 to 1994 Oszlanyi
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 299

Abstract

Because Hungary has about 18 percent forest area, it is not as densely forested as most of the countries in

Europe. Forests are dominated by native species such as oaks, beech, hornbeam, and other broadleaves. As a

result of an intensive afforestation in the last 50 years, introduced species, such as black locust, Scots pine,

and improved poplars were widely planted and forest area increased by 50 percent — to 18,000 km2. Several

observations in Hungary from the early 1980’s detected an unusual amount and type of forest damage affecting native and introduced species. A shocking observation was the oak decline sweeping through the country

with a dieback rate of 10 to 70 percent within a decade. New methods had to be developed to detect and

explain changes of forest health status and forest ecosystems. In close cooperation with the International

Cooperative Program on the Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests), a

national multi-level monitoring and research program was launched in 1986. The first level of the program

— large scale monitoring — accepted defoliation and discoloration as key parameters, but a detailed survey

of each sample tree was added. On the basis of the annual survey of about 22,000 trees, a significant increase

of defoliation was observed. Although 79 percent of the trees were healthy in 1988, only 44 percent were

healthy in 1995. Proportion of damaged trees (defoliation greater than 25 percent) seems to have stabilized

at about 20 percent in the last 4 years, close to the average of the European survey results ranging from 6 to 60

percent. Oaks, black locust and Scots pine were particularly heavily defoliated, while beech and hornbeam

were only lightly affected. The disturbed balance of forest ecosystems is closely related to the lack of water —

a result of river regulation since the 18th century and especially the 12-year period of dry and hot weather in

the Carpathian Basin. Although SO2

 emission has been reduced by 50 percent since 1980 and NOx by 30

percent since 1987, and direct damages have not been observed, air pollution can be a predisposing factor of

forest damages. Additional intensive research is necessary to clarify the role of climatic factors and air pollution effects on forest ecosystems, including direct effects, carbon and nutrient cycling at different sites, and

damaging biotic factors.

Introduction

Because Hungary has about 18 percent forest area, it is not as densely forested as

most of the countries in Europe. Forests are dominated by native species, such as

oaks (35 percent), beech (Fagus sylvatica L) (7 percent), and other broadleaves (13

percent). As a result of an intensive afforestation in the last 50 years, introduced

species, such as black locust (Robinia pseudo-acacia L), Scots pine (Pinus silvestris L),

and improved poplars were widely planted and forest area increased by 50 percent

to 18,000 km2. Although the total area of the country is only 93,000 km2, about 150

tree species and subspecies occupy the landscape as a result of diverse site conditions with a variety of different forest types, ranging from the nearly treeless steppe

to the mountainous beech forests.

Different Sources of Information about

Forest Damage

Forest health has been a major concern of forest scientists and practitioners and has

been continually observed. In the last 40 years different systems have been developed to detect or forecast forest damage.

Forest Health Status in Hungary1

1 An abbreviated version of this paper

was presented at the International Symposium on Air Pollution and Climate

Change Effects on Foret Ecosystems,

February 5-9, 1996, Riverside, California.

2 Group Leader, Forest Management

Planning Service, H-1054 Budapest,

Szechenyi u. 14, Hungary.

András Szepesi2
300 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Local Damage Report

The traditional and most widely used method was based on the damage reports of

forest owners that were collected and evaluated by the Forest Research Institute

(FRI). This method is still in use and can provide fairly good results for forestry

practice, but it cannot be considered as representative. Thus, its role is limited.

Light Traps

Another program, a network of 25 light traps, has been run for 32 years. This monitoring provides reliable information on insect migrations and gradations. The results are collected and evaluated by the FRI and published annually with special

focus on forecasts and recommendations.

Monitoring of Oak Decline

Because of the oak decline in the early 1980’s, a specific monitoring program was

established in 1987 to follow the development of the disease. Although the decline

recently reached its peak, the network is still in use. Similar specific monitoring

started at the end of the 1980’s in beech forests.

Data are collected by the University of Forestry, the Forest Research Institute,

and the crews of the Forest Management Planning Service (FMPS). Sample plots

are located in selected stands. The data collection is completed by using visual

assessments that apply specific codes describing the different stages of the decline.

Reports are published annually.

Large Scale Surveys

Several observations in Hungary from the early 1980’s detected unusual amount

and type of forest damage affecting native and introduced species. In close cooperation with the International Cooperative Program on the Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) — a European program initiated and run by the Economic Commission for Europe (ECE) of the United Nations under the Convention on Long-range Transboundary Air Pollution — a national multi-level monitoring and research program was launched in 1985. With

special regard to the site and climatic conditions in Hungary, the priority of air

pollution effects of the international program was accepted, but special emphasis

was put on all kinds of different damaging factors.

The forest management plans cover the entire forest area, with a 10-year renewal period. The methodology was extended by the FMPS for the collection of

more detailed data on health status at the stand level, which was a specific part of

the national program in 1985. Results have been evaluated on the national and

regional level, and were the basis of the national health condition reports until

1988, when the first survey of the systematic monitoring plots was done. The main

goal was to inform forest owners on the health status of the forests and to adjust

management practices according to these findings.

The national forest protection program followed the outline initiated by the

ICP Forests Manual (1986) and distinguished three different intensity levels:

● Level I: Large scale survey of forest condition to follow the spatial

and temporal development of damages.

● Level II: Intensive monitoring of forest condition to recognize the

key factors and processes.

● Level III: Special forest ecosystem analysis aimed at gaining a

deeper insight into cause-effect relationships.

In the first level of the national program — large scale monitoring run by the

FMPS — recommendations of the ECE Manual were accepted, especially defoliation

and discoloration as key parameters. A detailed list of all kinds of visible abiotic or

biotic symptoms was also added for a better and more detailed description of the

Session IV Forest Health Status in Hungary Szepesi
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 301

sample trees. With the establishment of 1,027 sample plots (24 trees/plot) in a 4 by

4 km grid (1987) and the first survey in 1988, the first forest health status data

from a representative sampling became available for decision makers, which

was an effective tool to identify the problems and concentrate the efforts to the

critical areas.

Air Pollutants

The ICP Forests cooperation is based on a hypothesis that new types of forest damage are closely related to air pollution effects. In the recent ICP Forests report (U.N.

Economic Commission for Europe, European Commission 1995) more than half of

the participating countries mentioned air pollution in their national reports as possible predisposing, accompanying, or triggering factor of forest damage. A clear

advantage of the transitional period in Hungary was the reduction of air pollutants

as a result of the recession of heavy industry and the restructuring of the whole

economy. In addition, the environment protection authorities became more effective during the last decade and implemented the European air pollution protocols.

Therefore, an improvement of the air quality was a result of two substantially different factors.

However, numerous disadvantages of the transitional period, include the lack

of an updated and comprehensive traditional statistical system. Consequently, the

emission charts show 1992 as the last reliable publicly available data. (According to

different sources, the levels have been stabilized or continued to sink since then.)

Sulfur Dioxide

Sulfur dioxide emission is still rather high in Hungary, mainly because of brown

coal combustion. The 1.6 million tons/year emitted in 1980 was reduced to 830,000

tons/year in 1992 (fig. 1). This reduction corresponded with the Helsinki (Finland)

protocol (1985) that targeted 30 percent sulfur dioxide reduction between 1980-

1993 in Europe. (All the 20 signatories fulfilled the commitments.)

Nitrogen Oxides

The nitrogen oxides protocol signed in Sofia, Bulgaria in 1988 required that by 1994

the emissions of all signatories did not exceed the 1987 levels. The majority of the

26 signatories fulfilled the commitments, including Hungary with 31 percent reduction. Total emission of nitrogen oxides was reduced from 270,000 tons/year in

1980 to 180,000 tons/year in 1992 (fig. 2).

Concentration of Gaseous Compounds

Emission is only one factor; it is more important to follow the changes of concentrations and depositions of pollutants in the forests. Concentration of gaseous pollutants — sulfur dioxide (SO2

) and nitrogen oxides (NOx) — were considerably

lower in 1994 than at the beginning of the observations in 1988 (fig. 3). Charts are

based on data from the three measuring stations continuously operating since 1988

in the western, northern, and central forested region of Hungary. Reduction of concentrations was unexpectedly fast, not clearly explained, but related to the Europewide efforts to reduce the emission of these air pollutants (for instance, neighboring Austria also reduced SO2

 emission to one-fifth within the last 10 years.)

Despite these reductions, there are some areas where clear evidence of local air

pollution impact has been observed on forests. The general level of these frequently

cited air pollutants seems to be rather low compared to central and western Europe. There is a real danger, however, that the reconstruction of the industry may

cause an emission increase.

Session IV Forest Health Status in Hungary Szepesi
302 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Figure 2 — Nitrogen oxide

(NOx) emission in Hungary, 1980-

1992.

Figure 3 — Sulfur dioxide and

nitrogen dioxide concentrations

in Hungary’s forest area, 1988-

1994.

Figure 1 — Sulfur dioxide (SO2

)

emission in Hungary, 1980-1992.

80 81 82 83 84 85 86 87 88 89 90 91 92

year

Session IV Forest Health Status in Hungary Szepesi

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USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 303

Soil Acidification

Acidification of the soil is a real threat and a detected ongoing process in large

forest areas of Europe (Germany, Czech and Slovak Republics, Poland). The reduction rate of hydrogen ion deposition is much less than that of the gaseous compounds in the forest area (fig. 4). It has also been measured since 1988; however,

because the majority of the soils in Hungary are calcareous or well-buffered, the

risk of acidification is moderate in general. To detect any kind of acidification may

require more time, especially on soils with good buffer capacity. Measurements on

research sites have shown that acidic precipitation is neutralized on the leaves in

broadleaved forests, which diminishes the effects of acid deposition.

Clear evidence of acidification on agricultural lands was detected in the last

two decades, but the main reason was the intensive fertilization and the selection

of fertilizers. Role of air pollution was not measured or estimated.

In forested areas resampling of old soil pits and changes of the grass vegetation

may indicate the acidification process, but there are not enough reliable data available. A special sampling method was applied with nine subsamples by using a

5- or 10- year resampling period to detect the changes of main soil characteristics

quantitatively.

Climate

Climate change and its possible effects on forests are debated. Hungary lies in the

Carpathian Basin and is a meeting point of three main climatic regions: the Atlantic from west, the Mediterranean from the south, and the continental from the east.

Even a little shift of the main climatic regions may cause considerably different

weather conditions in the country dominated by lowlands.

According to the recent assumptions on climate change, a drier and milder

climate is predicted for the Carpathian Basin. Because one-third of the country is

already characterized by a continental climate, productive closed forests are not

expected without a surplus of ground water. Deterioration of large forest areas and

decrease of productivity may occur as a result of drier climate. Foresters did not

pay much attention to the climate change scenarios in Hungary because of the uncertainties, but the dry and mild climate of the last 12 years put the issues in the

focus of the experts. Abatement strategies, such as the reduction of carbon dioxide

were not effective enough, with about 25 percent reduction of the annual emission

within the last 15 years. This left the emission on a high level of 65 million tons/

Figure 4 — Hydrogen ion deposition in Hungary’s forest area,

1988-1994.

year

1988 1989 1990 1991 1992 1993 1994 1995

Source: Meteoroiogical Srevice

Session IV Forest Health Status in Hungary Szepesi
304 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

year (fig. 5). The increase of the growing stock, including afforestation and better

use of the different assortments, has also been calculated as capturing about 10

percent of the carbon dioxide emitted recently.

Precipitation has an outstanding importance in the life of forests in Hungary.

The stress of high temperature and lack of water is characterized by a drought

index that depends on the precipitation in October to August and on the monthly

average temperature in April to August ( additional factors, like hot days, rainless

periods, and groundwater level are included as well) (fig. 6). The index is wellknown in agriculture and with some modifications, it can also be informative in

forestry. Main categories of the index include:

● Moderate drought 5 - 6 ● Strong drought 7 - 8

● Medium drought 6 - 7 ● Very strong drought 8 +

Although the level of air pollutants decreased in the last 15 years, the weather

conditions became more unfavourable for the forests in general. This statement

does not refer necessarily to climate change effect, since the meteorological data are

within the natural variation of the long-term observation results, rather, this phenomenon is an indication of the possible causes regarding the health status observed from the large scale damage survey since 1988.

Figure 5 — Carbon dioxide

(CO2) emission in Hungary, 1980-

1992.

Figure 6 — Drought index in

Hungary, 1978-1993.

year

1978 1980 1982 1984 1986 1988 1990 1992 1994

Source: Meteoroiogical Srevice

Session IV Forest Health Status in Hungary Szepesi
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 305

Results of the Large Scale Forest Damage Survey

Comprehensive Observations

Defoliation, discoloration and about 44 different symptoms (classified according to

the main parts of the tree where they can appear or cause damage) have been assessed by individual sample trees annually since 1988. In 1995 about 1,104 sample

plots and 23,000 sample trees were assessed. In 1991-1993 a soil survey was performed based on the national methodology, soil pits, and sampling by genetic horizons. In 1994 additional soil samples were collected with a special sampling design that detected changes of the most important soil characteristics on 67 plots

within the 10-year resampling period.

Increment measurements started in 1993, based on 2.83 by 2.83 km net and 200

by 200 m tracts, including all plots of the 4 by 4 km damage survey network. All

data collected are important for a detailed and reliable inventory of forest health

status and for a scientific interpretation of the observations.

Defoliation in Europe

Defoliation is considered a general indicator of the health status. Despite the common European Manual and the continuous efforts on better harmonization, international comparability is still biased by the differences among the national methods. However, we can compare the forest condition in some European countries to

highlight the Hungarian situation. Because about 22 percent of the trees are considerably defoliated (defoliation above 25 percent), Hungary is close to the average

defoliation observed in Europe in 1994 (fig. 7). The highest damage was observed

in the Czech Republic, where 60 percent of the trees were considerably defoliated,

while the rate of damaged trees was only 6 percent in Portugal.

National Results

Defoliation

All types of damages are assessed in 10 percent classes, but interpretation is often

combined into 5 classes to follow a classification that is accepted internationally.

The following classes are used in the charts:

● Not damaged 0 - 10 percent

● Slightly damaged 11 - 25 percent

● Moderately damaged 26 - 60 percent

● Severely damaged, dead 61 -100 percent

Figure 7 — Defoliation (greater

than 25 percent) in Europe in

1994. Key:

CZ=Czech Republic;

PL=Poland;

SK=Slovak Republic;

DK=Denmark;

UA=Ukraine;

LV=Latvia;

N=Norway;

LT=Lithuania;

D=Germany;

CH=Switzerland;

H=Hungary;

NL=Netherlands;

B=Belgium;

EST=Estonia;

GB=United Kingdom;

SF=Finland;

F=France;

A=Austria;

P=Portugal.

pct

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Two different categories of defoilations were used in Hungary. The first is ‘total

defoliation’ including all kinds of leaf losses regardless of the reasons of defoliation;

and the second is simply ‘defoliation’, which is the international terminology that

excludes clearly identified reasons of defoliation, such as leaf eaters, fungi, or breaks.

According to the total defoliation category (fig. 8), 33 percent of the trees were

classified healthy, while 27 percent showed considerable defoliation (above 25 percent leaf loss). Noble oaks (European oak and sessile oak) were in the worst condition with about 50 percent of the trees defoliated considerably. Although clear

evidence has not yet been found, oak stands may have been affected by an increment loss because of the intensive reduction of the leaf area. Primary damage factors include foliage insects (mainly caterpillars) and some fungi that attack the

native species like oaks or hornbeam (Carbines beetles L), while conifers or black

locust have been attacked only scarcely and regionally.

The second category, defoliation (fig. 9), had a better relationship to air pollution or drought effects. Oaks and especially hornbeam were more healthy than on

the chart of total defoliation, but noble oaks were still in the worst condition. With

at least moderate defoliation (above 25 percent) European oak (Quarks robber L)

(32 percent), sessile oak (Quarks petard L) (28 percent), and black locust (26 percent) had the highest defoliation, while beech and hornbeam were the least affected (7 and 9 percent, respectively).

Figure 8 — Total defoliation

(23,000 sample trees) in Hungary,

1995. Key:

Q. rob = Quercus robur;

Q. pet = Quercus petraea;

oth. Q = other quercus species;

Q.cer = Quercus cerris;

Fagus = Fagus sylvatica;

Carp. = Carpinus betulus;

Rob. = Robinia pseudoacacia;

o. hard = other hard broadleaves;

Pop. = poplars;

o. soft = other soft broadleaves;

conif. = conifers

Figure 9 — Defoliation (23,000

sample trees) in 1995.

Session IV Forest Health Status in Hungary Szepesi

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USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 307

Defoliation change was alarming in the first 3 years of the survey, showing a

rapid increase. But since 1991 the situation seems to have stabilized (fig. 10). The

proportion of healthy trees decreased from 80 percent to 44 percent and trees with

considerable defoliation (class 2-4) increased from 7 percent to 20 percent within

the survey period of 1988-1995.

Mortality is a complex problem in managed forests where the intensity of sanitary fellings and thinning may considerably change the results collected on the

permanent plots. In 1995 about 2.9 percent of the sample trees were dead with

great differences among the tree species (spruce —12 percent, beech — 1 percent).

The mortality rate doubled during the survey period because of the health condition, and the privatization of 40 percent of the forests, in which proper management was not always performed by a responsible owner.

While the results of the total sample are slightly distorted by the replacements

and the new sample plots, a comparison was also made for the common sample

trees (common trees in 1989-1995) (fig. 11). To reduce the possible errors of the first

survey, when the staff had only moderate experience, the 1988 data was excluded.

Defoliation of the common sample trees was faster than the total sample and the

defoliation in 1995 is higher — only 38 percent of the sample trees are classified as

healthy, and 25 percent defoliated considerably. Defoliation of the most important

Figure 10 — Defoliation in 1988-

95, total sample (17,100 to 23,300

trees).

Figure 11 — Defoliation in

1989-95, common sample trees

(13,100 trees).

Session IV Forest Health Status in Hungary Szepesi
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tree species was predetermined by site conditions and annually modified by

weather conditions as well as other damaging factors.

The oak decline, affecting sessile oak, peaked in 1988, and on a lower level still

exists. Oak stands on shallow soils were severely affected by the long dry period of

the last 12 years. In the northern mountainous region, third and fourth generation

coppice stands were the most sensitive. High defoliation is not only related to the

oak decline; the water reserves in the soil also became exhausted and resistance of

the stands to various damages decreased (fig. 12).

Dieback of European oak started at the beginning of the survey (fig. 12)

and was closely related to the traditional water management practice, which

lead the water out of the Carpathian Basin, an area frequently threatened by

floods, that diminished surface waters in springtime on agricultural areas. Roots

of middle-aged and older stands were not able to follow the sinking water,

while water deficiency and possible uptake disturbances reduced the resistance of the oak. Thus, the trees also became more sensitive to insect (Tortrix

sp, Lymantria dispar L., Melolontha melolontha L.) and fungus (Microsphaera

quercina Schw. Burr., Armillaria sp.) attacks.

Turkey oak (Quercus cerris L.) is less sensitive to drought (Mediterranean species,

but native in Hungary), often growing on shallow soils or on the drier southern or

western slopes. Even the increase of slight defoliation (fig. 12) to 40 percent (similar

to noble oaks) reduced the canopy closure so that dense grass vegetation became

an important obstacle of natural regeneration. Unexplained diebacks were reported

in some localities of the northern mountainous region.

Natural and artificial regeneration was more difficult in this dry period and

frequency of good seed crops decreased (oaks, beech). Moreover, the regeneration

areas are to be protected by fences to reduce game damages. In several areas the

grass vegetation became a dangerous competitor of the seedlings because in the

spring only the upper 10-20 cm of the soil is saturated properly with water.

Beech was the healthiest species (fig. 13). Unlike other tree species in Hungary, beech seldom grew out of its ecological niche. In the humid and cool

climate, where the 7 percent beech forests are located, effects of drought were

less severe. However, local foresters and pathologists have focused on these

species for the past 10 years and have recorded various types of damage related to climatic or biotic factors. Higher defoliation in 1994 was connected

with high seed production throughout the country, a phenomenon that reduced

the amount of leaves, and increased defoliation.

Figure 12 — Defoliation of

European oak, sessile oak, and

turkey oak in 1988-1995.

Session IV Forest Health Status in Hungary Szepesi
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Black locust was introduced in the beginning of the 19th century and intensively planted in the country by small owners for fuelwood and later by the cooperatives. Because only a few fungi and insects attacked black locust in Hungary, the

species appeared healthy until the mid 1980’s, but large scale observations showed

rapid deterioration afterwards (fig. 13), especially on the sandy and shallow soils.

Although the oaks could not respond to the higher precipitation of 1994-1995, the

black locust responded, and defoliation in these years decreased. Because black

locust was mainly planted close to or on the edges of the villages, resulting in numerous scattered small forest lots, and dead trees have been continuously exploited

by local people, the number of dead trees has remained at the same level since 1988

in spite of the increasing mortality.

The group of poplars is heterogeneous, dominated by the improved poplars in

plantations (15-30 m3 annual increment) regularly suffering from specific fungi

(Chryptodiaporthe populea Sacc. Butin., Marssonina brunnea E. et E., Magn.), or the

white and black poplar, native in the country. Despite the dominance of the sensitive improved poplars, the health condition of the poplar group has only slightly

deteriorated in the last 4 years (fig. 13). It is characterized by a high variation of

spatial distribution depending on soils and different clones. The stabilized health

condition is also attributed to the focus on the selection of sites suitable for new

plantations, as well as new more resistant clones.

Although 15 percent of the forest area is covered by conifers, they are not native in Hungary. Even spruce that was planted in the humid sites does not receive

sufficiently high precipitation and permanently high air humidity. Consequently,

rapid and alarming deterioration was observed within the survey period (fig. 14)

attributed to low precipitation. Although pines dropped only the eldest needle curls

in the dry years, spruces lost more and more needles annually. In 1995 about 12

percent of the sample trees were dead, partially because trunks were attacked by

root decaying tinder (Heterobasidion annosum Fr. Bref.) in several regions where the

replanting of conifers required special technologies or they would not be economical any longer. Intensive attack of bark beetles in the weakened stands with dead

trees may result in the collapse of the forest within 2-3 years.

The problem is the same in Scots pine stands (about 8 percent of the forests)

where defoliation has been rather stable in the last 5 years (fig. 14), but the level was

constantly high. Trees were more defoliated in the western border region and in

higher elevations — within more favourable climatic conditions — than that in the

lowlands, where the existence of closed forest is uncertain (there are some different

hypotheses, but no clear explanation about the phenomenon). In these dry sandy

Figure 13 — Defoliation of

beech, black locust, and poplars

in 1988-1995.

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regions, Scots pine has no deciduous alternative if attacked by root decaying tinder

or killed by the lack of water so that replanting of conifers (species already introduced to the country) becomes impossible. The original vegetation — steppe and

scattered small groups of trees and bushes — can only be restored within a long

period, depending on the weather conditions.

Austrian black pine (Pinus nigra Arn.) is very tolerant to drought, so it has been

planted in the poor sites where soil protection has primary importance. In addition

to the lack of water, new pathogens, like Diplodia pinea, native in the Balkan peninsula, attacked the stands and increased the high defoliation attributed to drought

— especially in 1992. Since then a definitely higher mortality was observed (fig. 14).

Despite the unfavorable conditions, Austrian black pine has a relatively low defoliation, but a prolonged dry period or additional fungus attacks may rapidly increase diebacks and deforestation is also possible in some regions.

Discoloration

Although discoloration caused by air pollution or disturbed nutrient balance, such

as lack of calcium or magnesium, has been described in some European countries,

it has not been observed in Hungary. In the dry years or after unusually long hot

and dry summer periods early fall of leaves was reported, starting with yellowing

in the second half of the survey period (in the first half of August). Discoloration of

black locust, hornbeam, and Scots pine have always been high, with outstanding

results in 1990 when about 20 percent of the respective tree species were considerably discolored (more than 25 percent discoloration); but only 3 percent was found

in 1995 (figs. 15,16). While 39 percent of the sample trees were at least slightly

discoloured in 1990, less than 12 percent were discolored in 1995. Presence of fungi

or insects is another common reason of discoloration observed on oaks, poplars,

and conifers. However, decrease of discoloration was not in correlation with the

decrease of fungus and insect damage; on the contrary, the frequency of identified

crown damage increased gradually within the survey period.

Other Damage

Methodology of the field survey was extended by a detailed list of 44 various biotic

and abiotic damage factors or symptoms identified on the tree. The damage factors

were grouped according to the main parts of the tree where they appear or cause

damage (fig. 17). The intensity of the damage is also recorded in 10 percent classes.

Some damage was typical to certain tree species during the whole survey period, like foliage insects on European oak and hornbeam, and frost ribs on Turkey

Figure 14 — Defoliation of Scots

pine, Austrian black pine, and Norway spruce in 1988-1995.

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oak and some improved poplar clones, and bark injuries on beech. Other damage

from game and drought affected nearly all tree species and has been more frequently identified recently. It is difficult to clarify the role of damaging factors based

on the defoliation and discoloration results that show a general worsening of the

condition of forest health. Unfavourable weather conditions and some suspected

effects of air pollution might have reduced the resistance of the trees or caused

deterioration directly, and these weather conditions may have also resulted in an

abundance of various insects and fungi on forest trees.

Future Outlook

Despite the monitoring and research efforts, it is difficult to predict the development of forest damage in Hungary. The large-scale survey highlighted the alarming development and extent of forest damage and encouraged the forest authorities as well as the public to pay more attention to the protection of our forests. The

forest protection program is supported by the government, and as a second level of

the national implications of the ICP Forests cooperation, 14 intensive monitoring

plots were selected and equipped according to the standards of the ECE Manual.

The government provided a special fund available for forest owners to diminish

forest damages and encourage regeneration.

Figure 16 — Discoloration of

17,100 to 23,300 sample trees in

1988-1995.

Figure 15 — Discoloration of

23,300 sample trees in 1995.

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Figure 17 — Primary damage

symptoms of 23,300 sample trees

in 1995.

Effective air pollution control may keep emissions on a constant level after the

recession, or even further decrease can be expected due to the acceptance of the

European Union’s air pollution standards. Better cooperation is necessary with water

management authorities to keep more water in the Carpathian Basin and ensure

the permanent feedback to the ground water without creating new swampy areas.

Additional intensive research is necessary to clarify the role of climatic factors

and air pollution effects on forest ecosystems, including direct effects, carbon and

nutrient cycling at different sites, and damaging biotic factors.

Acknowledgments

I thank Laurie Dunn for technical editing of this manuscript.

References

Csoka, Peter.; Szepesi, Andras. 1988-1995. Large scale forest damage survey in Hungary. Budapest, Hungary: Forest Manage- Large scale forest damage survey in Hungary

ment Planning Service; 22 p.

Forest Management Planning Service. 1995. Manual of the large scale forest damage survey. 2nd ed. Budapest, Hungary; 32 p. Manual of the large scale forest damage survey

Fuhrer, Erno. 1995. Effects of weather changing to the wood productivity and health condition of forests. In: Proceedings of Effects of weather changing to the wood productivity and health condition of forests

the changing health condition of the forests; 1995 March 2; Budapest, Hungary: Hungarian Academy of Sciences, Forestry Council; 38-45.

Juhasz, Istvan, ed. 1994. Environmental indicators of Hungary. Budapest: Ministry of Environment and Regional Policy; 67 p. Environmental indicators of Hungary

Program Coordinating Centres of the ICP Forests. 1994. Manual on methods and criteria of harmonized sampling, assess- Manual on methods and criteria of harmonized sampling, assessment, monitoring and analysis of the effects of air pollution on forests. 3rd ed. Hamburg, Germany: International Coop- ment, monitoring and analysis of the effects of air pollution on forests

erative Program on the Assessment and Monitoring of Air Pollution Effects on Forests; 177 p.

Szepesi, Andras. 1995. Site exploration at the points of the 4x4 km forest protection network. In: Proceedings of the changing Site exploration at the points of the 4x4 km forest protection network

health condition of the forests; 1995 March 2; Budapest, Hungary: Hungarian Academy of Sciences, Forestry

Council; 138-150.

UN Economic Commission for Europe, European Commission. 1995. Forest condition in Europe, results of the 1994 survey. Forest condition in Europe, results of the 1994 survey

Brussels, Geneva: UN Economic Commission for Europe, European Commission; 106 p.

Session IV Forest Health Status in Hungary Szepesi
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 313

Abstract

Growth and productivity of forests are important indicators for understanding the general condition and

health of forests. It is very important that indicators detected during monitoring procedures afford an opportunity for direct or indirect evaluation of forest productivity and its natural and anthropogenic changes.

Analysis of the U.S. Forest Health Monitoring (FHM) crown indicators showed that primary crown indicators are not enough for biological interpretation of collected data and estimation of potential tree growth. The

length of crown is necessary for calculating meaningful crown indicators; consequently, measurements of tree

height should be included into a FHM sampling design. Statistically reliable tree height measurements as

well as basal area estimation are necessary for evaluation of tree volume and biomass. Results of a pilot

study on relationships between tree growth and crown indicators, and a multiple regression model for assessment of potential productivity of trees based on primary and composite crown indicators are presented in

this paper. New composite crown indicators are developed to increase interpretability of FHM data. A modification of FHM tree height, basal area measurement and estimation procedures is proposed. 

Introduction

Increased environmental pollution, especially from anthropogenic sources, emphasizes the importance of monitoring the environmental role of forests in Lithuania.

Forest vegetation, especially trees, are a basic component of forest ecosystems and

are characterized not only by their impressive dimensions and age, but also by

their high sensitivity to environmental pollution. Main coniferous species (Scots

pine and Norway spruce), which make up to about 60 percent of Lithuanian forests, are especially sensitive to air pollution. As the health and structure of forest

ecosystems worsens, biodiversity and productivity decreases, and aging processes

accelerate in this polluted environment.

Because of their longevity, forests have an important effect on various processes in the entire biosphere. Their sensitivity to anthropogenic influence integrates past and current environmental situations and is one of the most suitable

objects for environmental monitoring. Long range transboundary pollution and

environmental acidification affects are considered a main reason for the wide range

of forest decline during the last decade. Lithuania has participated in the European

Forest Monitoring program (International Cooperative Program on Monitoring and

Assessment of Air Pollution Effects on Forests) since 1988. The main forest condition indicator in this program is the visually assessed crown defoliation. According to data of the Lithuanian National survey (Ozolincius 1996), defoliation of the main tree species has affected more than 80 percent of Lithuanian

forests (table 1).

Defoliation of different tree species has increased rather substantially during

the last several years. A significant increase in defoliation was registered for Norway spruce. Although there has been a significant decrease in air pollution in

Lithuania and in Europe during the last 3-4 years (Juknys 1995), the increased defoliation can be explained as a result of the integrated effects of anthropogenic and

natural stressors. Unfavorable climatic conditions of past years, especially extremely

hot and dry summers in 1992 and 1994, and the invasion of insects have contributed to this negative process. Some stabilization and even decreased defoliation

Forest Health Monitoring Indicators and

Their Interpretability: A Lithuanian Case Study1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and Climate Change Effects on Forest Ecosystems, February 5-9, 1996, Riverside,

California.

2 Professor of Forestry, Vytautas

Magnus University, S. Daukanto St.

28, 3000 Kaunas, Lithuania.

3 Forester, Lithuanian Agricultural University, 2434 Kaunas region,

Lithuania.

Romualdas Juknys,2 Algirdas Augustaitis3
314 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

for pine and birch recorded in 1994-1995 (table 1) can hopefully be considered a first

sign of positive biological reaction to the improved environmental situation.

Forest Health Monitoring activities in Lithuania, as well as in other Baltic countries, were started in 1993 when field training with the participation of specialists

from the U.S.A. and all three Baltic countries was begun in Birstonas (Lithuania).

By following FHM protocol in the Forest Health Monitoring Field Method Guide

(FHM 1994), field work started in 1994.

This paper discusses the beginning of the FHM methodology implementation

in Lithuanian environmental monitoring procedures and the biological interpretation of the collected data; we also analyzed the necessity to change the field work

methodology to enhance the interpretability of the data.

Objectives and Methods

Growth and productivity of forests are one of the main indicators reflecting the

general condition and health of monitored forests. Therefore, it is very important

that indicators used during monitoring procedures allow for direct or indirect evaluation of forest productivity and its natural and anthropogenic changes.

Because trees are main component of forest ecosystems, growth, productivity,

and mortality of trees are very important indicators for forest monitoring. Because

direct measurements and estimation of tree growth and productivity is very time

consuming and usually requires destructive sampling procedures, indirect methods based on easily detected external indicators are preferred.

Different crown indicators are usually used to evaluate potential growth of

forest trees. Many investigations were performed during the last decade to evaluate relations between crown indicators (mainly defoliation of trees) and tree growth

(Braker and Gaggen 1987, Juknys 1991, Kontic and Winkler-Seifert 1987, Kramer

1986, Schweingruber 1985). Different results were obtained, but the main conclusions were that correlation between crown defoliation and growth of trees (usually

width of annual rings) is rather weak and the coefficient of negative correlation

usually doesn’t exceed 0.3-0.5.

Our investigation of relations between growth of trees and FHM crown indicators was performed during 1994-1995 using the European forest monitoring system. Crown defoliation was assessed at the same time. Primary proposals to improve interpretability of FHM data were presented during the Baltic countries-U.S.A.

FHM meeting, which was held in Kaunas (Lithuania) in December 1994.

Investigations were performed in Scots pine forests surrounding the

Jonava mineral fertilizer plant where nitrogen and sulfur compounds are

the primary emissions.

The main objectives of this research were:

● To elaborate methodology and models for assessment of potential

productivity of trees on the basis of primary and composite crown

indicators.

● To define the most suitable tree growth indicator as a response

function to crown quantity and quality indicators.

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis

Table 1 Table 1 — Average defoliation of different tree species. Table 1 

Average defoliation (percent) Average defoliation (percent)

 Tree species 1989 1990 1991 1992 1993 1994 1995

Scots pine 22.8 23.3 25.5 23.6 25.2 24.2 24.0

Norway spruce 15.6 16.5 17.9 17.5 23.1 21.7 28.6

Birch 19.3 21.7 23.3 19.7 23.8 20.7 20.5

Total 18.8 19.7 21.3 21.0 23.4 23.0 24.2
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 315

● To develop new indicators and proposals to improve field work

methodology in order to enhance interpretability of FHM data.

During the investigation, all tree crown and stem indicators were separated a

priori into primary indicators (measured or evaluated during field work) and composite indicators (secondary or tertiary, calculated on the basis of primary indicators). On other hand, for better understanding and interpretation, all (primary and

composite) indicators were defined as quantitative or qualitative indicators.

Primary analysis of FHM crown indicators have already shown that the list of

quantitative indicators measured during FHM field work is not sufficient for

biological interpretation of collected data and estimation of potential growth of

trees. Length of crown is absolutely necessary for calculation of meaningful

composite crown indicators. For live crown ratio evaluated according to the FHM

field protocol, height should be measured for some subsample of trees to have data

on crown length. Statistically reliable tree height data are also necessary for

determination of stand productivity (m3/ha or m3/ha/year).

Three semimature Scots pine stands were chosen for the investigation and 25

trees were systematically sampled in each of them. The following primary crown

and stem indicators were measured for each sampled tree:

● Quantitative indicators: Stem diameter (DBH; average of maximal

and perpendicular crown diameters); tree height (THG); height of

crown bottom (HCB) and crown diameter (CRD; average of

maximal and perpendicular crown diameters).

● Qualitative indicators (in percent): Crown density (CDN; included

into FHM program); foliage transparency (FTR; included into FHM

program); crown dieback (CDB; included into FHM program) and

crown defoliation (DEF) according to the European forest

monitoring system.

Additionally, increment cores were taken from each tree at breast height and

two increment indicators were estimated — radial tree increment (RI) and basal

area increment (BI) — from 1 to 10 years. Kramer (1986) showed that for assessment of potential tree growth both crown quantity and crown quality should be

considered. The following composite tree crown indicators were used for further

analysis:

● Crown surface area (CSA) as an integrated crown quantity

indicator.

● Total crown losses (TCL) as an integrated crown quality indicator.

Crown surface area and crown volume usually are calculated on the basis of

crown diameter and crown length data, and assume that the crown is cone or parabolic shaped. But more detailed investigations on the basis of extensive study of

crown for felled trees (Kravchenka 1972) showed that such a strictly geometrical

approach can give only a rough approximation to crown surface area or crown

volume. Shape of Scots pine crown depends essentially on the age of tree and crown

class. Crown shape coefficient (K) (ratio of crown surface area or crown volume to

the surface area or volume of cylinder with the same diameter and length) varied

from 0.530 for young and suppressed trees to 0.674 for mature dominant trees

(Kravchenka 1972).

Crown surface area (CSA) was calculated as follows:

CSA = CRD * CRL * π * K (1)

in which CRD — crown diameter, CRL — crown length.

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis
316 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

For the investigated forest stands (70 years), the following crown shape coefficients

(K) were used for different crown classes: 1 (open growth) — 0.660; 2 (dominant) —

0.630; 3 (codominant) — 0.600; 4 (intermediate) — 0.570; 5 (overtopped) — 0.550.

Reduced crown density and increased foliage transparency and dieback were

considered crown losses (losses of branches, shoots, and foliage) because of the

influence of different internal (competition) and external (environmental pollution,

insects, diseases, etc.) factors. Total crown losses (TCL) in percents were calculated

by using the equation:

TCL = 100 - ((100 - DNL) * (100 - DBL) / 100 * (100 - FTL) / 100) (2)

in which DNL — crown losses because of decreased tree crown density (losses of

branches, shoots, and foliage); DBL — crown losses because of increased dieback

(losses of shoots and foliage); FTL — crown losses because of increased foliage

transparency (losses of foliage).

For determination of crown losses, these questions were examined: what can

be considered normal density, normal dieback, and normal foliage transparency?

On the basis of 2 years of FHM data, normal (maximal) crown density for Scots

pine was assessed as 70 percent, normal (minimal) crown dieback as zero, and

normal (minimal) foliage transparency as 5 percent.

Tree crown density losses (DNL) in percent were calculated:

DNL = (NDN - CRD) / NCD * 100 (3)

in which: NCD — normal crown density; CRD — crown density of tree.

Tree crown losses because of increased foliage transparency (FTL) in percent were

calculated:

FTL = ((FTR - NFT) / 100 - NFT) * 100 (4)

in which: NFT — normal foliage transparency; FTR — foliage transparency of tree.

Crown losses because of increased crown dieback should be assessed according to

the analogous formula (4), given that normal dieback for Scots pine equals zero,

DBL = CDB.

The composite crown indicator — relative crown amount (RCA) — is considered a

mostly integrated crown indicator, which includes practically all quantitative and

qualitative crown indicators:

RCA = CSA * NCD / 100 * (100 - TCL) / 100 (5)

in which: CSA — crown surface area; NCD — normal crown density; TCL — total

crown losses.

Results

In order to define the most suitable tree growth indicator as a response function to crown indicators, we correlated radial tree increment (RI) and basal

area increment (BI) for different durations (from 1 to 10 years) with the FHM

crown indicators (table 2). We found that correlation of crown indicators with

radial tree increment is essentially weaker than with basal area increment because basal area increment reflects biomass production much better than ringwidth increments. The same ring width increment can result in very different

biomass increment for trees with different diameters. In addition, foliage transparency is mostly related to radial tree increment between qualitative Scots

pine indicators. Correlation of crown dieback with radial tree increment is weak

and usually statistically insignificant (α= 0.05) for Scots pine.

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 317

Coefficients of correlation were averaged between different durations of

tree increment indicators and foliage transparency for all investigated tree

stands (fig. 1). For 5 to 6 years the correlation of tree increment with crown

indicators essentially increased, but further increase of duration usually caused

stabilization or even decrease of correlation coefficient. Five-year basal area

increment can be considered the most suitable tree increment indicator relating to FHM crown indicators.

FHM qualitative crown indicators and crown defoliation were correlated according to the European forest monitoring system (table 3). Closest correlation with

crown defoliation (according to European system) was detected for foliage transparency. Correlation of crown defoliation with crown dieback is weak and statistically insignificant (α= 0.05).

1 Statistically significant (α=0.05).

Figure 1 — Correlation of tree

increment with foliage transparency for different periods.

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis

Table 2 Table 2 — Correlation of radial and basal area increments with primary FHM crown indicators for different periods. Table 2 

Crown Length of period (years) Length of period (years)

1 2 3 4 5 6 7 8 9 10

 indicator Correlation coefficient (significance level) Correlation coefficient (significance level)

Radial increment Radial increment

Density 0.296 0.260 0.302 0.363 1

0.417 10.439 10.434 10.426 10.417 0.403

0.159 0.220 0.151 0.085 0.043 0.032 0.034 0.038 0.042 0.056

Transparency -0.368 -0.300 0.370 1

-0.473 1-0.578 1-0.623 1-0.628 1-0.624 1-0.618 1-0.541

0.076 0.154 0.074 0.019 0.003 0.001 0.001 0.001 0.001 0.007

Dieback -0.036 -0.174 -0.128 -0.137 -0.148 -0.173 -0.223 -0.236 -0.247 -0.237

0.867 0.417 0.548 0.519 0.489 0.416 0.295 0.266 0.244 0.261

Basal area increment area increment

Density 1

0.451 10.446 10.489 10.541 10.555 10.554 10.532 10.517 10.507 10.491

0.027 0.029 0.015 0.006 0.005 0.005 0.007 0.010 0.011 0.012

Transparency 1

-0.533 1-0.495 1-0.569 1-0.665 1-0.710 1-0.726 1-0.706 1-0.616 1-0.688 1-0.623

0.007 0.013 0.004 0.000 0.000 0.000 0.000 0.000 0.000 0.001

Dieback -0.057 -0.240 -0.202 -0.242 -0.251 -0.269 -0.323 -0.374 -0.349 -0.320

0.791 0.245 0.315 0.243 0.273 0.210 0.123 0.072 0.095 0.126

0.60

0.50

0.40

0.30

Correlation coefficient

–

–

–

––––––––

1 2 3 4 5 6 7 8 9 10

Length of period, years

Radial Increment

Basal Area Increment
318 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Five-year basal area increment and different crown indicators were correlated

(fig. 2). Crown length is more closely related to tree increment than crown diameter. Foliage transparency and crown density area are most strongly related to tree

increment from primary crown qualitative indicators. Foliage transparency is also

closely correlated with crown defoliation according to the European forest monitoring system (table 3).

Relative crown amount, which integrates quantitative and qualitative crown

indicators, is most strongly correlated with tree increment from composite crown

indicators. Correlation of total crown losses, as a composite qualitative crown indicator with tree increment is closer to that for crown surface area, which integrates

quantitative crown indicators. Correlations of primary qualitative crown indicators (with the exception of crown dieback) with tree increment in general is larger

than correlations of quantitative primary crown indicators (fig.␣ 2).

Finally, the influence of crown quality on the growth of trees with a different

crown quantity (crown surface area) was investigated. Results of multiple regression analysis showed (fig. 3) that for suppressed trees with small crowns the influence of crown quality (total crown losses) is much less than for trees with well

developed crowns (big crown surface area). Multiple regression coefficient equal

about 0.7; consequently, almost one-half of basal area increment variation can be

explained on the basis of composite qualitative and quantitative crown indicators.

Figure 3 — Influence of qualitative and quantitative composite

crown indicators on tree increment.

Figure 2 — Correlation between

5-year basal area increment and

different crown indicators. Key:

CDM - crown diameter,

CLG - crown length,

LCR - live crown ratio,

CDN - crown density,

FTR - foliage transparency,

CDB - crown dieback,

CSA - crown surface area,

TCL - total crown losses,

RCA - relative crown amount.

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis

160

140

120

100

80

60

40

20

Basal area increment, cm

2

BI5 = 42.667 + 0.701CSA - 0.148CLS - 0.00727CSA * CLS

(R = 0.678)

Crown Losses, pctCrown surface area, m2

160

100

40

0 10 20 30 40 50 60 70 80 90 100

–

–

–

–

–

–

–

–

–

–

–

–

–

–

–

–

–

Correlation coefficient
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 319

Discussion

For meaningful interpretation of forest monitoring data, the possibility to assess

general features of forests, such as productivity, sustainability, etc., on the basis of

easily detected external indicators is very important. Analysis of FHM indicators

showed that the list of primary indicators measured and evaluated during field

work is not sufficient for biological interpretation of collected data and estimation

of potential tree growth and its anthropogenical changes. For assessment of tree

growth on the basis of crown indicators, both crown quality and crown quantity

should be evaluated and measured. The FHM field work program includes only

one quantitative tree crown indicator — crown diameter — and one semiquantative

— live crown ratio. On the basis of these two indicators it is impossible to assess

crown quantity (crown surface area, crown volume). Tree height should be measured for some subsample of trees in order to have data on crown length. Statistically reliable tree height data are also necessary for determination of stand

productivity.

Five-year basal area increment can be considered the most suitable tree increment indicator relating to FHM crown indicators. Correlation of radial (linear) increment with crown indicators is weaker because the same linear increment can

result in different biomass increases. The composite tree crown indicator of total

crown losses integrates all tree FHM crown qualitative indicators (crown density,

foliage transparency, and crown dieback) and reflects general crown quality. Total

crown losses correlates more closely with tree growth than the composite crown

quantity indicator — crown surface area. An elaborated multiregresional model

showed that on the basis of qualitative and quantitative crown indicators, a reliable assessment of tree growth is possible.

Besides separate tree productivity (increment), assessment of productivity of

entire monitored stands, such as biomass, volume, and basal area per unit of area,

is absolutely necessary. According to the FHM field work program, basal area measurement is included in the list of tree site measurements, but really is not statistically based. Basal area of monitored stands calculated on the basis of measured tree

diameters on subplots would be more appropriate. Tree height measurements should

be adopted to supplement tree diameter measurements on subplots as special

subsamples. Simplified methodology for minimally statistically reliable tree height

measurements was developed by the Lithuanian FHM project and is presented in

the report for 1995 field season.

Table 3 — Results of correlation of qualitative FHM crown indicators with crown defoliation (significance level). Table 3

Stand Crown density Foliage transparency Crown dieback Total crown losses

(pct) (pct) (pct) (pct)

1 1-0.813 10.878 0.215 10.801

(0.000) (0.000) 0.314) (0.000)

2 1

-0.830 10.931 10.404 10.818

(0.000) (0.000) (0.049) (0.000)

3 1

-0.781 1-0.823 0.206 1-0.789

 (0.000) (0.000) (0.334) (0.000)

1statistically significant (α = 0.05)

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis
320 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

In Lithuania, as well as in other Baltic countries, three different forest monitoring systems were introduced during the last decade:

● The European International Cooperative Program on Assessment and

Monitoring of Air Pollution Effects of Forests (ICP-Forests) began in 1987,

and information on the condition of Lithuanian forests is presented every

year. This very simple and operative method was supplemented by the

forest soil monitoring subsystem from 1993 and more sophisticated second-level forest monitoring methods from 1995.

● From 1993 Lithuania and other Baltic countries (Latvia and Estonia)

joined the International Integrated Monitoring Program on Small

Catchments. The pilot phase of this program (Integrated Monitoring in

Terrestrial Reference Areas of Europe and North America) was started in

1988 within the UN/ECE Convention on Long-range Transboundary Air

Pollution. On the basis of financial support from Nordic countries, two to

three very well- equipped Integrated Monitoring Stations were established

in afforested background areas in each Baltic country during 1993-1995.

● The Forest Health Monitoring Program, with sound ecologically based

methodology and emphasis on biodiversity of forest ecosystems, may be a

very good extension of the Forest Monitoring System. It may help in filling

a gap between the very simple European Forest Monitoring System and

the very sophisticated and expensive Integrated Monitoring System.

Better coordination and coadaptation between those three systems are necessary. Baltic countries could be a very good case study for such international and

interprogram activities.

References

Braker, O.; Gaggen S. 1987. Tree ring analysis in the Swiss forest decline study. In: Forest decline and reproduction. Tree ring analysis in the Swiss forest decline study

Laxenburg; 124-129.

Forest Health Monitoring. 1994. Field methods guide. Washington: U.S. Environmental Protection Agency. Field methods guide

Juknys, R. 1991. Growth and productivity of Scots pine under polluted environment. Krasnojarsk. Ph.D. dissertation; 342 p. Growth and productivity of Scots pine under polluted environment

(in Russian).

Juknys, R. 1995. Trends of Lithuanian environment during transitional period. Environmental Research, Engineering, and Trends of Lithuanian environment during transitional period

Management 1: 15-24.

Kontic, R.; Winkler-Seifert, A. 1987. Comparative studies on the annual ring pattern and crown conditions of conifers. In: Comparative studies on the annual ring pattern and crown conditions of conifers

Forest decline and reproduction. Laxenburg; 143-152.

Kramer, H. 1986. Relation between crown parameters and volume increment of Relation between crown parameters and volume increment of Picea abies abries stands damaged by envi- stands damaged by environmental pollution. Scandinavian Journal of Forest Research 1: 251-263. ronmental pollution

Kravchenko, G. 1972. Regularities of pine growth. Moscow; 1688 p. [in Russian]. Regularities of pine growth

Ozolincius, R. 1996. Forest condition assessment in the regional network of forest monitoring. Annual Report. Kaunas; 36 p. Forest condition assessment in the regional network of forest monitoring

Schweingruber, F. 1985. Abrupt changes in growth reflected in tree ring sequences as an expression of biotic and abiotic

influences. In: Inventorying and monitoring endangered forests. Zurich; 291-295. influences

Session IV Forest Health Monitoring Indicators and Their Interpretability: A Lithuanian Case Study Juknys, Augustaitis
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 321

SESSION V

Photosynthetically Active Radiation (PAR) Measurements

and Their Use in Forest Health Monitoring

Craig Palmer, Chair
322 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 323

Abstract

Photosynthetically active radiation (PAR) has been measured in pure pine and mixed pine forests on 15 sites

in two transects in Poland: the “climatic” (from the western to the eastern border), and the “Silesian” (from

the most to the less polluted part of the country). PAR was measured by using the standard procedure developed by the USDA Forest Service’s Forest Health Monitoring Program. At each forest site PAR was measured

at the “central point” and at six points 7.3 m from the central point. PAR was also measured at three additional points (at the location of litterfall collection). Eight out of 15 forests were classified as pure pine, and

7 as mixed pine. Two series of measurements were completed in September/October 1994 and in July/August

1995. The first measurements showed that the mixed pine forests were more shaded than the pure pine forests. Average leaf area index (LAI) for mixed pine forests was 2.03 and 1.78 for pure pine. Mixed forests were

characterized by greater seasonal changeability of LAI because of a higher amount of broadleaved trees and

shrubs. LAI and transmitted PAR (TPAR) standard deviation on a particular forest site may be treated as

an index of internal forest differentiation. The indexes differ between particular forests, but average values 

for pure pine and mixed pine forest are almost identical. The first attempt to correlate PAR indexes and other

ecological indexes suggest that there is no correlation between PAR and litterfall and the rate of litter decomposition.

Introduction

Photosynthetically active radiation (PAR) was measured in pine forests of Poland

under the Polish-American cooperative program entitled “The Effect of Pollution

and Climate Change on Forest Ecosystems.” This program includes climatic, ecological, geobotanical, and soil research along two transects. The first transect (eight

sites) runs east-west along the 52nd parallel and is associated with the increasing

continentality of climate. The second transect (also of eight sites) runs from the

most polluted areas of Upper Silesia to the relatively unpolluted part of Poland

around the Bialowieza Forest. The most north-easterly point (K011) is common to

both transects; and out of a total of 15 sites, 8 were categorized as pure pine forest,

and the other 7 as mixed pine forest (fig.1). All the stands contain trees between 70

to 100 years old (Breymeyer and others 1995).

This study measured PAR along the climatic and Silesian transects in Poland

by using an analysis of the different amounts of light reaching the forest floor and

procedures of the USDA Forest Service’s Forest Health Monitoring Program (FHM

1994), in order to explain the rate of some ecological processes.

Methods

PAR research on the transect sites began in 1994. The first series of measurements

at the study sites were recorded in September and October 1994, and the subsequent series in July and August 1995. Unfavorable weather conditions forced the

abandonment of measurements at three sites in October 1995. Thus, the series of

measurements at three forest sites were incomplete in October 1995.

Organic matter decomposition was sampled at each site at a central point

(Breymeyer and others 1995), and the PAR measurements were made at the central

point and at six other points that were 7.3 m from the central point (fig. 2). In addition,

Photosynthetically Active Radiation Measurements in Pure

Pine and Mixed Pine Forests in Poland1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Geographer, Institute of Geography

and Spatial Organization, Polish Academy of Sciences, 00-927 Warsaw,

Krakowskie Przedmiescie 30, Poland.

Jaroslaw Smialkowski2
324 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Session V Photosynthetically Active Radiation Measurements in Pure Pine and Mixed Pine Forests in Poland Smialkowski

3 Mention of trade names or products is

for information only and does not imply endorsement by the U.S. Department of Agriculture.

Figure 1 — Forest sites studied

on the climatic and Silesian

transects in Poland.

Figure 2 — Schematic diagram

of PAR measurements on the

forest sites.

PAR was measured at three locations where litterfall was collected. Thus, PAR was

measured at a total of 10 points. Analysis of results from the first series of

measurements (autumn 1994) showed that data collected in conditions of variable

cloud cover (especially with convection cloudiness) were difficult to interpret.

Transmitted PAR (TPAR) and leaf area index (LAI) for the forest site were calculated

by using incoming light below the canopy and at an ambient station in an open

area. The under canopy measurement was a single ceptometer measurement lasting

10 to 15 seconds. The simultaneously collected data from the ambient station (located

in an open area) were stored as 1-minute averages (FHM 1994; Decagon Devices,

Inc. 1989; LI-COR, Inc. 1990)3. A false result is obtained (LAI may be below 0 and

TPAR above 1) if the sun shines during the 15-second under canopy measurement,

and is then covered by cloud for the rest of minute (Campbell 1991). Similar

distortions of the results may be linked to the distance between a measurement

point under the canopy and the ambient station (which is sometimes 300 - 500 m).

Uneven convection cloudiness may easily allow the sun to shine over a forest stand,

while the ambient station remains in shade. Because of the problems associated

with variable cloudiness, the measurements in 1995 were only made in sunny, cloudfree weather, or at times with minimal cloud cover.

K092 K098

K061

K023 K055 K117 KS011

K126

S023

S002

S060 S005

S126

S028

S011

N

30o

90o

150o 210o

270o

330o

- Measurement points

- Central point
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 325

Results

The results of the three series of measurements show that mixed pine forests are

characterized by lower levels of PAR reaching the forest floor than pure pine forests.

The mean for PAR in mixed pine forests was 0.25, while that for the pure pine forests

was 0.21 (fig. 3). Still greater differences were noted in the case of LAI, which varied

between 1.3 and 3 at the study sites. The mean LAI for pure pine forests was 1.78, and

that for mixed pine 2.03 (fig. 4). These differences are associated with the greater proportion of broadleaved trees and shrubs in the mixed pine forests.

Comparison of summer and autumn measurements yielded these results: all

sites for which both summer and autumn data are available showed LAI values to be higher in summer because of the presence of leaves. An exception

was site S023, in which the autumn results may be erroneous because of cloudiness. When data for this point are deleted, the mean difference in summer and

autumn LAI in mixed pine forest is 1.61 (i.e., a figure greater than the 1.36

recorded from pure pine forests) (fig. 5).

Standard deviations of the measured values of TPAR and LAI may be interpreted as a measure of heterogeneity of the forest sites under study. Standard deviation values for LAI were not related to the type of forest, but rather to its internal structure. In the case of LAI these ranged from 0.4 to 1.3. The greatest value was

noted for site S005, which had a considerable number of young spruce trees growing in the forest. The mean LAI standard deviation in pure pine forests was 0.75,

while that for mixed pine forests was 0.77 (fig. 6).

Preliminary attempts were made to correlate the PAR measurements and the

other ecological indices (e.g., litterfall and litter decomposition rates). However,

despite the fact that LAI is sometimes calculated based on litterfall, statistically

significant linkage has not yet been found, except for the inverse correlation between LAI and TPAR.

Figure 3 — Average TPAR on the forest sites. Figure 4 — Average LAI on the forest sites.

Session V Photosynthetically Active Radiation Measurements in Pure Pine and Mixed Pine Forests in Poland Smialkowski

K092 K061 K023 K011 S005 S028 S002 S011 K098 K055 K117 K126 S126 S060 S023

SITE

Pure pine Mixed pine

Avg = 0.25 Avg = 0.21

0.5

0.4

0.3

0.2

0.1

0

AVG TPAR

–

–

–

–

–

––––––––––––––

K092 K061 K023 K011 S005 S028 S002 S011 K098 K055 K117 K126 S126 S060 S023

SITE

Pure pine Mixed pine

Avg = 1.78 Avg = 2.03

3.5

3

2.5

2

1.5

1

0.5

0

AVG LAI

–

–

–

–

–

–

–

––––––––––––––
326 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Acknowledgments

This work was part of a project entitled “U.S.-Poland Cooperative Project on the

Status and Long-Term Trends in Forest Ecosystems: Climate, Pollution and Forest

Health.” The program was financed by the U.S. Environmental Protection Agency,

USDA Forest Service, USDA Foreign Agriculture Service, Polish Academy of Sciences,

Polish Bureau of Global Environmental Facilities (GEF), Forest Research Institute,

Michigan Technological University, Bowling Green State University, and the Institute

of Geography and Spatial Organization of the Polish Academy of Sciences.

References

Breymeyer, A.; Degorski, M.; Roo-Zielinska, E.; Solon, J.; Smialkowski, J. 1995. Effects of atmospheric deposition and climate

change in forest ecosystems in central and eastern Europe and the United States, 1994-1995 report. change in forest ecosystems in central and eastern Europe and the United States, 1994-1995 report. Warsaw: Institute of change in forest ecosystems in central and eastern Europe and the United States, 1994-1995 report.

Geography and Spatial Organization, Polish Academy of Sciences.

Campbell, G.S. 1991. Application note: canopy leaf area index from sunfleck ceptometer PAR measurements. Application note: canopy leaf area index from sunfleck ceptometer PAR measurements. Pullman, WA: Application note: canopy leaf area index from sunfleck ceptometer PAR measurements.

Decagon Devices, Inc.

Decagon Devices, Inc. 1989. Sunfleck ceptometer user’s manual. Sunfleck ceptometer user’s manual. Pullman, WA: Decagon Devices, Inc . Sunfleck ceptometer user’s manual.

FHM 1994. PAR. In: Tallent-Halsell, N.G., ed. PAR. Forest health monitoring field methods guide. Vol 1. Las Vegas, NV: U.S. EPA, Forest health monitoring field methods guide

Environmental Monitoring Systems Laboratory.

LI-COR, Inc. 1990. LI-1000 datalogger instruction manual, Lincoln, NE: LI-COR, Inc. LI-1000 datalogger instruction manual, Lincoln, NE

Figure 6 — Internal differentiation of the forest sites measured as

standard deviation of LAI.

Figure 5 — Differences in LAI between autumn and summer on the

forest sites. Average value for mixed pine in parenthesis is the value

when data for site SO23 is deleted due to cloudiness at the site (see

text).

Session V Photosynthetically Active Radiation Measurements in Pure Pine and Mixed Pine Forests in Poland Smialkowski

K092 K061 K023 K011 S005 S028 S002 S011 K098 K055 K117 K126 S126 S060 S023

Pure pine Mixed pine

Avg = 1.36 Avg = 1.28 (1.61)

2.5

2

1.5

1

0.5

0

-0.5

AVG LAI

–

–

–

–

–

–

––––––––––––––

K092 K061 K023 K011 S005 S028 S002 S011 K098 K055 K117 K126 S126 S060 S023

Avg = 0.75 Avg = 0.77

1.4

1.2

1

0.8

0.6

0.4

0.2

0

STD LAI

–

–

–

–

–

–

–

––––––––––––––

STD (LAI)
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 327

Abstract

Effective management of forest resources requires access to current and consistent geospatial information

that can be shared by resource managers and the public. Geospatial information describing our land and

natural resources comes from many sources and is most effective when stored in a geospatial database and

used in a geographic information system (GIS). The information on the location and condition of current

vegetation is one of the key elements in resource management. Remote sensing data, such as aerial photographs, satellite imagery, aerial video, and data collected by other remote devices, are primary sources for

mapping vegetation. Comparison of images acquired several days or several years apart can assist in determining changes that occurred in that time period. The Forest Service, U.S. Department of Agriculture, has

been using various remote sensing and associated technologies to map vegetation and monitor changes. Current needs include an increased training and awareness in use of this technology, as well as development of

new applications, standards, and guidelines.

Introduction

Geospatial information about land and natural resources is essential to effective

management of natural resources. Data from various remote sensing instruments

make a substantial contribution to the geospatial information needed in the USDA

Forest Service (FS). Remote sensing is a source of current and repeatable information on the location, quantity, and quality of land cover and other resource information, particularly vegetation. It allows change detection and monitoring of land

cover and vegetation over time. Data from various remote sensing tools form an

important source for various geographic information system (GIS) databases and

subsequent analysis. Many types of data are already in digital form, and others can

be digitized. Satellite imagery covers large areas and has the locational precision

and spatial resolution to satisfy many natural resources mapping requirements.

For many applications satellite imagery needs to be used in conjunction with a

closer view provided by aerial photographs and airborne video images. The global

positioning system (GPS) used in conjunction with satellite imagery helps managers determine changes in land cover over time.

This paper examines the ability to integrate remote sensing imagery in a GIS

combined with ancillary data, including cartographic feature files (CFFs), digital

elevation models (DEMs), digital orthophotoquads (DOQs), and other resource data

layers to provide a solid geospatial data foundation for support of Forest Service

management of natural resources.

Data Collection Through Remote Sensing

The USDA Forest Service uses data from a variety of remote sensing instruments,

such as the advanced very high resolution radiometer (AVHRR ), which uses data

obtained through the U.S. Geological Survey (USGS) EROS Data Center (EDC),

from the U.S. Department of Commerce, and from the National Oceanic and

Atmospheric Administration (NOAA) weather satellites. It has relatively course

spatial resolution of 1.1 by 1.1 km pixel size with five spectral bands. Broad area

Remote Sensing Applied to Resource Management1

1 An abbreviated version of this paper

was presented at the International

Symposium on Air Pollution and

Climate Change Effects on Forest

Ecosystems, February 5-9, 1996,

Riverside, California.

2 Program Leader, Integration of Remote Sensing, Engineering Staff,

USDA Forest Service, 2222 West 2300

South, Salt Lake City, UT 84119.

Henry M. Lachowski2
328 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

coverage is obtained with a swath width of 2,700 kilometers. Temporal resolution

is excellent with repeat coverage over a given geographic area every 12 hours. Major

applications for this data include: vegetation discrimination, vegetation biomass,

snow/ice discrimination, vegetation/crop stress, and geothermal mapping. This

imagery is currently used by the Forest Service in association with the USGS on

development of a global vegetation data set and with the United Nations on

verification for a global vegetation characterization data base.

Another remote sensing instrument is the Landsat Multispectral Scanner (MSS).

MSS imagery has a spatial resolution of 80 m, spectral resolution in 4 bands, revisits the same geographic location every 16 days, and covers an area of 185 by 170 km

per scene. The Landsat program gathered digital MSS data from 1972 through 1992.

The result is a 20-year time span of data that can support evaluations of change in

landscapes or land cover over a longer time period than any available earth observation system. MSS data is available through the EDC working in cooperation with

the National Aeronautics and Space Administration (NASA) Landsat Pathfinder

Program. Pathfinder efforts are focused on evaluation of global change by using

available remote sensing technologies. The North American Landscape Characterization (NALC) Project, a component of the Landsat Pathfinder Program, is developing an archive of MSS data and production of three-date georeferenced data sets,

called triplicates, acquired in 1970’s, 1980’s and early 1990’s. The NALC images are

geometrically rectified, georeferenced, and placed into a UTM map projection. Pixels are resampled into a 60 by 60 m size format for compatibility with the 30 by 30

m Landsat Thematic Mapper (TM) data resolution.

TM imagery has 30 m spatial resolution with seven spectral bands. The satellite covers the same geographic location every 16 days, each scene covering 185 by

170 km. The Landsat Program is the longest running program in the collection of

multispectral, digital data of the earth’s land surface from space. The temporal extent of the collection, the characteristics and quality of Landsat data, and the ability

to collect new data directly comparable to that in the archive, make Landsat data a

unique resource to address a broad range of issues in ecosystem management. An

interagency consortium called the Multi-Resolution Land Characteristics Monitoring System (MRLC) has been established to purchase TM imagery to cover the

United States (not including Alaska). This group has selected the appropriate scenes

and developed image processing, georeferencing, terrain correction, clustering and

classification strategies. This Federal consortium includes U.S. Environmental Protection Agency, EMAP; U.S. Fish and Wildlife Service, GAP Program; USGS EDC

and National Water Quality Assessment Program; and NOAA Coast Watch Change

Analysis Program. High resolution airborne video and digital cameras provide a

powerful tool for ecosystem management. A camera can be mounted in a small

airplane and imagery can be viewed as soon as the flight is complete (Bobbe 1995).

Video imagery linked with GPS provided location information for each frame, allowing users to quickly locate specific areas on maps or images. Video images can

be digitized and used with other GIS layers.

Use and Access to Remote Sensing Data

Applications of the various remote sensing sources vary widely among resource

management agencies, depending on their specific requirements. Users of digital

imagery range from novices, who simply use imagery as a backdrop for overlaying

GIS information, to expert users, who do classification of landscapes for input into

large area assessments or forest planning efforts. Thus, a wide range of user needs

must be met in the form of technology transfer and training to fullfil the Forest

Service’s mission. Access to remote sensing data should rapidly increase for several

reasons, among them, new cooperative agreements among agencies to share data

Session V Remote Sensing Applied to Resource Management Lachowski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 329

acquisition costs, and improved information access on the information super

highway. Currently, many potential users of remote sensing are either not aware of

how to access the data or believe that the cost of acquiring the data is prohibitively

high. Improved access to the data, both physically and financially, will likely increase

the number of users.

New data procurements, such as the one recently made by the Forest Service,

will provide AVHRR, Landsat Multispectral Scanner triplicates, and Landsat TM

imagery, for all lands in the continental U.S. This low cost purchase will provide

imagery except for Alaska at very low cost to the entire Forest Service at three

different scales. Purchases such as these provide inexpensive and easy access to

data and will surely spur interest in using digital imagery for applications such as

Resources Planning Act (RPA) assessments and forest planning. Access to the

internet now gives users the capability to preview satellite imagery acquired from

several platforms including Landsat. This will allow users to assess the condition

of the imagery (e.g., cloud cover) and will ensure that the user is confident in the

quality of the imagery before purchasing. Because internet access will also provide

an easy method for ordering and transferring digital imagery, the process of data

acquisition that was recently time-consuming and inefficient at best will be streamlined. Several new satellite sensors are being developed and are scheduled to be

launched within the next decade. Once these plans come to fruition, users of remote sensing data will have several new options. These sensors will allow for different perspectives of the landscape because of the differences in spectral, spatial,

and temporal resolutions of the imagery they will provide.

Applications of Remote Sensing

With new GIS hardware and software available to the Forest Service, and existing

GIS and image processing capabilities, the use of digital imagery for effective resource management will continue to increase. Virtually everyone in the Forest Service will have the capability to view and manipulate imagery in some way. The

Forest Service’s use of remote sensing has been shaped not only by technological

advances, but by the changing needs for information (Lachowski 1995). Increasingly, the synoptic view that is offered by remote sensing has become more important as issues such as endangered species and cumulative effects have shaped management activities on Forest Service lands. The need for information on the type

and condition of the vegetation is almost always critical to these issues and remote

sensing plays an important role in collecting this information.

Mapping Vegetation and Land Cover Characteristics

The Forest Service has a wealth of experience with using digital imagery for projects

ranging in scales from RPA assessments to forest and project plans. Perhaps the

most common use of digital imagery in the agency is for mapping vegetation and

land cover characteristics. For example, digital imagery from Landsat Thematic

Mapper has been used to map vegetation and model old-growth forests in California (Warbington 1993), Washington, and Oregon (Congalton 1993, Tepley 1993).

These successful, region-wide projects are examples of how satellite imagery can

be useful at the forest level for mapping vegetation and land cover. In addition,

numerous individual forests in every region have used digital satellite imagery to

map vegetation characteristics (Evans 1992, Gonzales 1992). Applications of this

information serve a variety of purposes ranging from multi-region and multi-agency

planning (Columbia River Basin Assessment) to forest planning efforts (Warbington

1993), and project level applications such as wild and scenic river planning

(Linderman 1995) and range allotment mapping and analysis (Martinez 1994). On

a broad scale, the Forest Service has used digital satellite imagery, such as AVHRR

Session V Remote Sensing Applied to Resource Management Lachowski
330 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

data from NOAA weather satellites, to map forest land for the entire U.S. and Mexico.

These maps are used for RPA assessments and are useful as a baseline analysis

for forests at the national level. Currently, other studies are planned to assess

the feasibility of linking this information with forest inventory and analysis

(FIA) inventories.

Change Detection

In addition to mapping vegetation and land cover, Forest Service managers have

recognized the usefulness of multi-date digital imagery in performing change detection for mapping insect and disease outbreaks as well as for updating management activities and forest plan monitoring (Maus 1992). Because each National Forest

is mandated by law to monitor the effects of their management over time, the ability to perform change detection is potentially the most effective use of digital imagery from remote sensing. Performing change detection may also be the most

underutilized application of digital imagery. This is particularly true as the span of

digital imagery available for an area becomes ever wider and the costs of older

data continue to plummet. Using digital imagery for performing change detection

offers many advantages over traditional methods. The traditional approach for

mapping change was to compare two maps, each interpreted independently. Often, false changes would result because of differences in classification schemes or

mapping techniques. With advances in geocoding of digital imagery, image-to-image comparisons can be made where spectral values between two or more dates of

imagery can be compared. These techniques can help eliminate problems associated with traditional map comparison methods. As awareness of these simple techniques for performing change detection increases and more resource managers have

access to the necessary computer hardware and software, the demand for such

analysis will certainly increase.

Ecological Unit Mapping

Current direction of the Forest Service requires that an ecological approach to resource management be used. To ensure ecosystem management is applied over

various scales of landscapes, the National Hierarchy of Ecological Units was developed (Avers 1993). Within the National Hierarchy, various scales of units are being

mapped and used for planning efforts at large scale (multi-region), medium scale

(multi-forest), and small scale (multi-district). The boundaries of these units will be

based on ecological influences, social/political considerations, and multi-agency

needs. Currently, remote sensing is playing a major role in the delineation of these

boundaries at all levels. Digital imagery from satellites is being heavily used for the

smaller scale units (e.g., ecoregion, sections, and subsections). The larger scale units

(e.g., land type associations, and landtype phases) are being mapped using combinations of aerial photographs and satellite imagery (Fallon 1994). Once ecological

units are mapped, their characteristics can be monitored over time by using remote

sensing techniques. Additional information can be added to ecological units for

project level analysis that can also be derived from remote sensing.

Disaster Assessment, Forest Pest Management, and Other Areas

Recent fires in large ecosystems such as the Greater Yellowstone Area and the Payette

National Forest have been mapped by using digital imagery from satellites. Other

landmark disasters, such as the Mount St. Helens eruption and the Mississippi

River flooding, have also been mapped from satellite imagery. Although the use of

satellite imagery for such events is somewhat rare, it is often the only type of current information available or accessible. Satellite imagery will continue to fulfill a

role in these assessments and will likely increase as access to and interest in digital

imagery increases.

Session V Remote Sensing Applied to Resource Management Lachowski
USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998. 331

Current Needs

Training and Awareness on How to Use the Data

New users of remote sensing imagery will need to know opportunities and

pitfalls of using the new technology. They will need to know when it is appropriate to use a particular type of imagery and when it may not be. With personnel reductions, fewer employees are likely to have graduated from recent

university remote sensing programs; thus, more will need to be trained. Using

remote sensing imagery will likely be a skill they acquire to compliment a wide

range of other job responsibilities they will have. New and experienced users

will need technical instruction in how to best use digital imagery for different

scales of projects ranging from simple applications, such as using imagery as a

backdrop for updating GIS layers to more complex applications like automated

image analysis for vegetation mapping and change detection. Users of digital

imagery will need to share ideas and techniques for using new types of digital

imagery. This transfer of information among users will need to take place between all levels of the Forest Service, including Ranger Districts, National Forests, regions, research stations, and State and Private Forestry.

New Sensor Applications Testing and Guidelines

As new types of imagery become available they will need to be assessed for cost

effectiveness as well as application suitability. Some new sensors will have

hyperspectral bands with very narrow wavelengths. Others will have higher spatial resolution than current satellite sensors. Still others will acquire imagery at

different times of the day such as early morning. However, increased access to remote sensing information may also increase potential for abuse of the technology.

As applications are developed for new types of imagery, guidelines for using the

imagery will need to be developed to inform prospective users of pitfalls and potentials. Pilot projects will need to be carried out to evaluate imagery for meeting

various user requirements.

Data Standards and Methods

The Forest Service will need to provide standards and guidelines in the collection,

design, use, and storage of remote sensing related data. These types of standards

will allow for information sharing between districts, forests, and regions. Information sharing across administrative boundaries will be critical to the success of ecosystem management practices. Standardized naming conventions, file structures,

and methodologies will promote information sharing and development of corporate database structures in the FS. These processes will simplify the use of geospatial

data, including data acquired through remote sensing and make the application of

these data sources to resource management easier and more efficient.

Acknowledgments

I would like to thank Laurie Dunn for technical editing of this manuscript.

References

Avers, Pete 1993. National hierarchical framework of ecological units. Washington DC: USDA Forest Service. National hierarchical framework of ecological units

Bobbe, Tom; McKean, Jim 1995. Evaluation of a digital camera system for natural resource management. Evaluation of a digital camera system for natural resource management. Earth Observation Evaluation of a digital camera system for natural resource management.

Magazine; 45-48.

Congalton, Russ; Green, Kass; Tepley, John 1993. Mapping-old growth forests on National Forest and Park Lands in the

Pacific Northwest from remotely sensed data. Pacific Northwest from remotely sensed data. Photogrammetric Engineering and Remote Sensing 59:(4) 529-535. Pacific Northwest from remotely sensed data.

Session V Remote Sensing Applied to Resource Management Lachowski
332 USDA Forest Service Gen.Tech.Rep. PSW-GTR-166. 1998.

Evans, David; Schoelerman, Lynn; Melvin, Tom. 1992. Landsat thematic mapper data for development and interpretation of Landsat thematic mapper data for development and interpretation of

vegetation information on the Kisatchie National Forest. In: Greer, Jerry, ed. Remote sensing and natural resource man- vegetation information on the Kisatchie National Forest

agement: proceedings of the fourth Forest Service remote sensing applications conference; 1992 April 6; Bethesda, MA:

American Society for Photogrammety and Remote Sensing; 117-124.

Fallon, Don; Svalberg, Terry; Tart, Dave; Maus, Paul; Wirth,Tim; Lachowski, Henry. 1994. Use of satellite imagery and digital Use of satellite imagery and digital

elevation models in the Bridger-Teton integrated resource inventory. elevation models in the Bridger-Teton integrated resource inventory. In: Greer, Jerry, ed. Remote sensing and ecosystem elevation models in the Bridger-Teton integrated resource inventory.

management: proceedings of the fifth Forest Service remote sensing applications conference; 1994 April 11; Bethesda,

MA: American Society for Photogrammety and Remote Sensing; 77-83.

Gonzales, Jesse; Maus, Paul. 1992. Vegetation classification and old growth modeling in the Jemez Mountains. In: Greer, Vegetation classification and old growth modeling in the Jemez Mountains

Jerry, ed. Remote sensing and natural resource management: proceedings of the fourth Forest Service remote sensing

applications conference; 1992 April 6; Bethesda, MA: American Society for Photogrammety and Remote Sensing.

Lachowski, Henry. 1995. Guidelines for the use of digital imagery for vegetation mapping. Washington, DC. USDA Forest Guidelines for the use of digital imagery for vegetation mapping

Service; EM-7140-25.

Linderman, Carl. 1995. East fork of the Jemez wild and scenic river plan, draft version. Santa Fe National Forest. East fork of the Jemez wild and scenic river plan, draft version

Martinez, Roberto; Miller, Larry; Whitney, Jeff; Lachowski, Henry; Maus, Paul; Gonzales, Jesse; Powell, Jay. 1994. An evalua- An evaluation of the utility of remote sensing for range management. In: Greer, Jerry, ed. Remote sensing and ecosystem manage- tion of the utility of remote sensing for range management

ment: proceedings of the fifth Forest Service remote sensing applications conference; 1994 April 11; Bethesda, MA: American Society for Photogrammety and Remote Sensing.

Maus, Paul; Landrum, Vaughan; Johnson, Jan; Lachowski, Henry; Platt, Bruce; Schanta, Mike. 1992. Utilizing satellite data

and GIS to map land cover change. Proceedings; Vancouver, B.C. Canada. and GIS to map land cover change

Tepley, John; Green, Kass. 1991. Old-growth forest: how much remains? Geographic Information Systems 5: 18-21. Old-growth forest: how much remains?

Warbington, Ralph; Askevold, Ray; Simmons, W. 1993. Building a regional GIS planning database for the National Forests of Building a regional GIS planning database for the National Forests of

California. The Compiler 11(4) 43-45. California

Session V Remote Sensing Applied to Resource Management Lachowski
The Forest Service, U.S. Department of Agriculture, is responsible for Federal

Leadership in forestry.

It carries out this role through four main activities:

• Protection and management of resources on 191 million acres of National Forest System

lands

• Cooperation with State and local governments, forest industries, and private landowners

to help protect and manage non-Federal forest and associated range and watershed

lands

• Participation with other agencies in human resource and community assistance programs

to improve living conditions in rural areas

• Research on all aspects of forestry, rangeland management, and forest resources

utilization.

The Pacific Southwest Research Station

• Represents the research branch of the Forest Service in California, Hawaii, American

Samoa, and the western Pacific.

The United States Department of

Agriculture (USDA) prohibits

discrimination in all its programs

and activities on the basis of race,

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USDA’s TARGET Center at

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To file a complaint of discrimination, write

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Office of Civil Rights

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or call

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USDA is an equal opportunity

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General Technical ReportResearch StationPacific SouthwestForest Serviceof AgricultureDepartmentUnited States

 February 5-9, 1996 Riverside, California

on Forest Ecosystemson Air Pollution and Climate Change EffectsProceedings of the International Symposium