Department of 

Agriculture 

Forest Service 

Northeastern Forest 

Experiment Station 

NE-RP-591 

Growth and utrient Status of 

Black Spruce Seedlings As 

Affected by Water Table Depth 

Miroslaw M. Czapowskyj Robert V. Rourke 

Walter J. Grant 
The Authors 

MIROSLAW M. CZAPOWSKYJ, a native of Ukraine and a United States citizen 

since 1957, received a Deiplomforstwirt degree from Ludwig-Maximillians University, Munich, Germany, in 1949. He obtained an M.S. degree in forestry from the 

University of Maine in 1958 and a Ph.D. degree in soils from Rutgers University in 

1962. He joined the Northeastern Forest Experiment Station in 1961 as a research 

forester. Since 1973 he has been with the Station's project in culture of northeastern conifers with emphasis on spruce and fir, at Orono, Maine. 

ROBERT V. ROURKE received B.S. and M.S. degrees in Agronomy from the 

University of Maine, Orono, in 1959 and 1964, respectively. Since 1964 he has 

been a soil scientist with the Maine Life Sciences and Agriculture Experiment Sta- g tion, University of Maine. He is currently conducting soil resource evaluations. 

WALTER J. GRANT, soil scientist, received B.S. and M.S. degrees in Agronomy 

from the University of Maine, Orono, in 1955 and 1957, respectively. He joined the 

USDA Agricultural Research Service in 1957 and has participated in research in 

the New England Region on the evaluation of soil erodibility, on use of cover 

crops, and on crop water relationships. 

Manuscript received for publication 21 January 1986 

1 Abstract 

: A greenhouse experiment was conducted to study the effects of soil water level on 

growth, biomass accretion, and inorganic element uptake by black spruce. Onec year-old containerized seedlings were grown for 3 years at three water table 

depths. All trees survived for the duration of the study confirming that black spruce I has a certain degree of survival tolerance to high water tables. However, tree 

height, diameter growth, and biomass production significantly increased as the 

depth to water table increased. 

The foliar levels of N, P, K, Mg, Fe, Zn, and B increased and those of Cu and Mn 

decreased with the increasing depth to the water table. For ash and Ca, differences were significant but did not follow a consistent trend. In shoots, the level of 

N, Ca, and Mg increased and those of ash, K, Fe, Cu, B, Al, and Mn decreased 

with the increasing depth to the water table. The level of P was not affected by the 

water table. In roots, the level of N and Ca increased and the level of ash, Mn, 

Fe, Al, and Cu decreased with increasing depth to the water table. The level of P, 

Mg, and Zn was significantly different but did not follow any trend. 

Foliar concentration of ash, Ca, Na, Mn, Fe, Zn, Cu, Al, and B increased and concentration of N, P, K, and Mg decreased with the increasing foliage age. In shoots, 

ash, Ca, Al, Fe, and Zn increased and N, P, K, Mg, and B decreased with the 

increasing tree and shoot age. In roots, Fe, Mn, Na, and Al increased and N, P, 

and Cu decreased with the increasing tree age. 

Northeastern Forest Experiment Station 

370 Reed Road, Broomall, PA 19008 

August 1986 
I 

I 

Introduction 

1 Black spruce (Picea mariana (Mill) B.S.P.) is an important 

component of the spruce-fir forest of northern Maine. It occupies about 6 to 7 percent of spruce-fir forest area. The 

species occurs naturally on a variety of soil and site condi- 

~ tions, but most frequently it is associated with swamps, 

extensive flatlands, and fringes of lakes where seomewhat 

poorly drained to very poorly drained soils occur. Many sites 

are occasionally flooded for prolonged periods, but the species seems to be morphologically adapted to survive periodic inundation. 

Since black spruce is the least susceptible of all spruces to 

spruce budworm (Choristoneura fumiferana Clem.) infestations, it has become a favored species for reforestation 

programs. Extensive plantations are being established on 

sites that vary in soil drainage conditions, and containerized 

seedlings are most commonly used as planting stock. Since 

soil drainage is of utmost importance to forest growth, the 

establishment of physiologically healthy plantations on poorly 

drained soil may be a problem. The soil conditions and the 

stresses likely to affect plant growth from excessive soil 

moisture have been well discussed in many publications. 

They were reviewed by Cannell and Jackson (1981). 

Examples of adverse effects on height and diameter growth 

of southern pine seedlings resulting from high water tables 

were presented by Burton (1971), McKee and Shoulders 

(1 974), McKee and others (1 984), and Mueller-Dombois 

(1 964). Little has been written concerning the effects of excessive moisture or of soil drainage on black spruce. 

Czapowskyj (1 982) reported that mechanical drainage of 

very poorly drained soils in northern Maine resulted in a 

positive response by young natural black spruce. Thus, 

there is a need to evaluate the influence of the soil water 

table level on black spruce growth and nutrient uptake during initial stages of plantation establishment given the growing interest in artificial regeneration using this species. 

I The objective of this study was to determine the effect of 

different soil water table levels on growth, biomass production, and nutrient accumulation in black spruce seedlings 

growing under greenhouse conditions over three growing 

1 seasons after transplanting. 

I Study 

Black spruce seedlings were planted in plastic pots in the 

spring of 1981 and were grown in the greenhouse with three 

water table regimens for three growing seasons. 

Seedlings 

One-year-old container grown seedlings were obtained in 

May 1981 from the Great Northern Paper Company overwintering beds in Millinocket, Maine. The seedlings were raised 

from seeds obtained from J. D. Irving, Ltd., Juniper, New 

Brunswick-Lot No. SB 58 Restigouche Regiow-and were 

grown in a peat moss-vermiculite mix in Japanese paper 

pots (FH-408) starting in May 1980 for 12 weeks in a greenhouse. During. this period, the seedlings were fertilized 6 

times at 6- to 10-day intervals using commercial fertilizer formulations, 4 times with 20-20-20 followed by two applications of 9-45-15, at a rate of about 12 grams per 1,000 

seedlings plus one application of micronutrients (0.5 gm/ 

1,000 seedlings). In August 1980, the seedlings were transferred from the greenhouse and placed in outside beds to 

harden off and overwinter. During April and May of 1981, the 

seedlings received additional fertilizer treatments; five applications of 9-45-1 5 plus five applications of micronutrients 

at the same rates as the previous year. Mensurational data 

and nutrient concentrations of the seedlings at the beginning 

of this study are given in Table 1. 

Growth Medium 

The seedlings were planted in soil collected from the spodic 

horizon from the Telos soil series (coarse-loamy, mixed, 

frigid, Aquic Haplorthods). This soil is somewhat poorly 

drained and occurs in extensive, contiguous blocks in the 

spruce-fir region of northern Maine. It is associated with the 

moderately well drained Chesuncook and the poorly drained 

Monarda soils. The Telos soil occurs in gently sloping areas 

and has slow or very slow permeability in the substratum 

because of its dense, low-porosity nature. The upper soil 

layers above the dense basal till are moderately permeable 

and of lower density. The soils are extremely to very 

strongly acid (pH 4.2 to 5.0). The combination of little relief 

and slow permeability result in a high water table in the 

spring and fall and in periods of high rainfall during the 

growing season. Soil was transported to the greenhouse, air 

dried, and sieved through a sieve with 1.3-cm openings. 

Study Layout 

Sixty plastic pots, 30 cm in height with a surface area of 6.5 

dm2, were filled with approximately 20 kg of soil. Each pot 

was planted with three seedlings, for a total of 180 seedlings. A 3-cm layer of peat moss was placed on the soil 

surface in each pot to simulate the organic layer usually 

found on forest sites. The pots were then placed in a 1.25-m 

x 2.50-m water tank with a fixed overflow elevation to simulate three water table levels. The polyethylene-lined plywood 

tank was constructed in such a way that each water table 

regime contained 20 pots. The water table levels were 4, 10, 

and 20 cm below the mineral soil surface--WT 4, WT 10, 

and WT 20, respectively. \ 

Water was automatically pumped from an overflow reservoir 

into the tank at timed intervals in order to replace losses due 

to evaporation. At the initiation of the study, the pH of the 
Table 1.-Physical characteristics and concentration of nutrient elements in 1- 

year-old containerized seedlings prior to planting' 

Seedling 

characteristic and Foliage Shoot Root Total 

nutrient element seedling 

Height, cm - 13.4 - - RCD,~ mm - 2.0 - - 

Oven-dry weight, g 0.29 .ll 0.19 0.59 

Nitrogen, percent 1.43 .85 .54 1 .04 

Phosphorus, percent .27 .I2 .29 .25 

Potassium, percent 1.08 .86 .74 .78 

Calcium, percent .87 .35 .79 .76 

Magnesium, percent .13 .I2 .39 .22 

Manganese, mglkg 200 100 100 183 

Iron, mglkg 181 174 2056 745 

Zinc, mglkg 124 125 366 205 

Copper, mglkg 16 60 42 32 

Boron, mglkg 31 20 76 44 

Aluminum, mglkg 100 100 200 133 

Ash, percent 4.44 2.76 7.60 5.21 

'Based on a sample of 25 seedlings. 

2Root collar diameter. 

water in the tank was adjusted to match the typical pH of 

forest soil solution. 

At the end of October in each growing season, the pots 

were drained and placed in an outside unheated trailer for 

overwintering. At mid-April of the second and third growing 

seasons they were transferred to the greenhouse and arranged in the same order and placed under the same water 

table regime as they were during the first growing season. 

Growth Analyses 

Survival counts, height, and diameter (at ground line) were 

measured at 2-week intervals from the beginning of May 

to mid-October during each growing season. Each year at 

the end of October one seedling was harvested from each 

pot for growth and biomass measurements as well as to 

determine nutrient concentrations of foliage, shoots, and 

roots. Roots were separated and washed in distilled water, 

foliage was separated from the shoots, and the components 

were dried at 65°C to constant weight. Because the trees 

were small, three composite samples were prepared from 20 

trees in each treatment. Samples were ground to pass a 

20-mesh screen and stored for chemical analyses. 

Chemical Analyses 

The samples were dry-ashed at 480°C, and ash content was 

determined gravimetrically. The ash was taken up in dilute 

HCL, and phosphorus (P), calcium (Ca), magnesium (Mg), 

potassium (K), sodium (Na), manganese (Mn), zinc (Zn), 

iron (Fe), aluminum (Al), and boron (B) were determined by 

inductively coupled plasma emission spectroscopy (ICP) 

at the Universeity of Maine. Nitrogen (N) was determined 

from Kjeldahl digestions by an autoanalyzer. All analyses 

were performed in duplicate on the oven-dry (65°C) samples. 

Data Analysis 

The data were tabulated by seedling age, component age, 

and by the depth to water level. Tree growth attributes 

(height, diameter, and oven-dry weight), percentage of ash, 

and nutrient concentrations were analyzed statistically by 

analysis of variance. Duncan's new multiple range test was 

used for mean separations. 

Results and Discussions 

Survival 

Seedling survival was not affected by the depth to the water 

table. All trees remained alive for the duration of the study, 

indicating the ability of black spruce to survive under high 

water table levels. 

Buds started to break 1 week after the pots were returned 

from the overwintering environment to the greenhouse. Buds 

broke first on trees with treatment WT-20. The WT-4 and 
WT-10 treatments had bud break delayed for 6 to 8 days. 

Delayed bud break on trees subjected to waterlogging is 

a common phenomenon that has been observed in the past 

(Coutts 1981). In addition, the foliage of trees with WT-4 and 

WT-10 regimens was consistently chlorotic, as is typical of 

saturated soil conditions. 

Height and Diameter Growth 

Biweekly measurements taken during the growing season 

revealed that shoots grew for about 8 weeks regardless 

of the water table regime. However, the diameter growth of 

seedlings in the WT-4 regime ceased after 12 weeks, but 

continued in the WT-10 and WT-20 regimes for a total of 20 

weeks. 

Average annual height and diameter growth (Table 2) show 

tree response to differences in water table levels. These 

responses were significant (p = 0.05) and both the mean 

height and mean diameter were ordered as follows: WT-4 R 

< WT-10 R < WT-20 R. This pattern held for the entire study 

period. 

Table 2.-Mean height and diameter at root collar of 

black spruce as affected by depth to water 

table and growing season 

Tree 

characteristic WT-4 WT-10 WT-20 

Tree Age, 1 + 1-40 trees sampled 

Height, cm 1 8.2b2 19.9a 20.2a 

RCD,' mm 2.9~ 3.2b 3.5a 

Tree age, 1 + 240 trees sampled 

Height, cm 26.4~ 30.2b 35.7a 

RCD, mm 4.0~ 5.0b 6.2a 

Tree age, 1 + 3--20 trees sampled 

Height, cm 29.6~ 34.0b 41.9a 

RCD, mm 5.lc 6.5b 8.5a 

'Root collar diameter. 

'Lower case letters designate differences between water table 

levels of the same year at the 0.05 level, using Duncan's New 

Multiple Range Test. 

Biomass Allocation 

Data presented in Table 3 show the means and differences 

in biomass allocation by tree component for the three water 

table regimens. For each of the 3 years, biomass responded 

to water table regimes similarly to tree height and diameter. 

The differences were significant for all years and the means 

were ordered as follows: WT-4 < WT-10 < WT-20. By expressing the data in Table 3 in relative terms (percent), the 

experiment wise (years x water level) allocation of extra 

biomass attributable to better drainage was ordered as follows: foliage > shoots > roots. The differences in root:shoot 

ratios were negligible: 37;38;41 percent for WT-4, WT-10, 

and WT-20, respectively, at the conclusion of the study. 

All three growth measurements (height, diameter, and biomass) provide additional evidence that the high water table 

had a negative effect on the performance of black spruce. 

Trees that were grown under the high water tables (WT4 and WT-10) showed signs of stress from poor aeration resulting in reduced oxygen diffusion. The stress symptoms-- 

reduced growth, chlorosis, abscission of the foliage, and 

deformation of root systems-were consistent with those reported by Norby and Kozlowski (1 983) for other species. 

Further examination of the data given in Table 2 and 3 and 

Figure 1 shows that lowering the water level by 6 cm (from 4 

to 10 cm) resulted in height increases of 15 percent, diameter increases of 27 percent, and total biomass increases of 

68 percent. Height increased by 42 percent, diameter increased by 67 percent, and biomass nearly doubled when 

the water table was lowered to 20 cm. 

Concentrations and Distribution of Ash and Mineral 

Elements 

The effects of both water level and seedling age on concentrations of ash, and selected elements in the foliage, shoots, 

and roots are readily apparent in Table 4. Because of a 

lack of trends and the small magnitudes in differences, the 

values for ash and Al concentrations were presented as 

averages of all three water table levels (Table 4). A summary of analysis of variance showing significant differences 

is presented in Table 5. Changes in nutrient composition 

varied considerably among individual components, water table levels, and component age classes. 

Needles. Mean concentrations of N, P, K, Mg, Fe, B, and 

Zn increased and those of Mn and Cu decreased with increasing depth to water table. For ash and Ca, the differences in concentration were significant but did not follow a 

consistent trend. Concentrations of A1 did not differ between 

water levels. 

Concentrations of ash and all mineral elements varied 

greatly with foliage age. The overall mean concentrations of 

ash, Ca, Na, Al, Zn, Fe, Cu, and B increased and those of 

N, P, K, and Mg decreased with an increasing foliage age 

from current-year to 2- and 3-year-old needles. Inconsistencies were observed in Mg concentrations, but the differences were small (Table 4). 

Shoots. Mean concentrations of N, Ca, and Mg increased 

and those of ash, K, Fe, Cu, B, Mn, and Al decreased along 

the gradient of increasing depth to water table level. No 
Table 3.-Mean component weight of black spruce seedlings raised under three depths of water table for a period 

of 3 years (average of 20 trees sampled) 

Biomass WT-4 WT-10 WT-20 

g percent g percent g percent 

Tree Age, 1 + 1 

Foliage 0.8c1 42 l.lb 44 1.4a 45 

Shoots .6c 32 .8b 32 1 .Oa 32 

Roots .5b 26 .6b 24 .7a 23 

~ Total 1.9~ 100 2.5b 100 3.la 100 

1 Root: Shoot Ratio .29 

='I 0002 .68 

I 

Foliage 

Shoots 

Roots 

Tree Age, 1 + 2 

2.8~ 43 3.8b 43 6.9a 40 

2.0b 30 2.6b 29 5.7a 32 

1.8b 27 2.5b 28 4.8a 28 

Total 

Root: Shoot Ratio .38 

1 "1 000 69 

Foliage 

Shoots 

~ Roots I 

Total 

Tree Age, t + 3 

4.8~ 41 8.2b 42 13.la 38 

3.5~ 30 6.0b 31 11.8a 35 

3.4~ 29 5.4b 27 9.2a 27 

1 1.7~ 1 00 19.6b 100 34.la 100 

Root: Shoot Ratio .37 

"1 000 .69 

'Lower case letters designate differences between water table levels of the same year at the 0.05 level, using Duncan's New Multiple 

Rtnge Test. 

1,000 needles weight. 

Table 4.-Mean concentrations of ash and nutrient elements in black spruce as related to water table level and seedling age 

Tree Tree Component W-4 + 10 + 20 WT-4 WT-10 W-20 WT-4 W-10 W-20 

component age age class (average) 

Year Year Ash, percent N, percent P, percent 

Needles 1-1 O+ 1 5.77 (.67)' 1.37cZ 1.62b 1.97a .14a .14a .17a 

1-2 0 3.92 (.70) .62b .67b 1.12a .08a .08a .10a 

1 +2 5.49 (.76) .64b .73b 1.00a .07b .06b .08a 

1-3 0 4.32 (.22) .76b .65b 1.07a .09a .O9a .lla 

1 5.35 (.26) .74b .68b 1.05a .07a .08a .08a 

2+3 6.49 (.42) .68b .65b .98a .07a .07a .07a -- - - 

Shoots 1-1 O+ 1 1.68 (.25) .80b .86b 1.00a .I 1 a .09a .10a 

1-2 0 1.44 (.17) .35b .38b .53a .05b .05b .06a 

1 +2 1.54 (.16) .27b .27b .33a .03a .03a .04a 

1-3 0 1.65 (.08) .33a .38a .46a .07a .07a .08a 

1 1.63 (.lo) .28a .31 a .38a .05a .04a .04a 

2+3 1.74 (.11) .26a .22a .34a .03a .03a .03a 

Roots 1-1 - 3.85 (.59) .66a .68a .85a .10a .08a .09a 

1-2 - 3.63 (.52) .50a .45a .52a .05a .05a .05a 

1-3 - 4.1 1 (1.29) .57a .39b .48ab .06a .05ab .05a 
1 Table 4.--Continued 

t 

Tree Tree Component WT-4 WT-10 WT-20 WT-4 WT-10 WT-20 WT-4 WT-10 WT-20 

component aae aae class 

Year 

Needles 1-1 

1-2 

Shoots 1-1 

1-2 

Year 

O+ 1 1.24a 

0 .71 b 

1 +2 1.29a 

0 .71a 

1 1.15a 

2+3 1.54a 

0+1 .37a 

0 .27b 

1 +2 .38a 

0 .27b 

1 .31a 

2+3 .37a 

Ca, percent 

.93b 

.70b 

1.18a 

.67a 

1.05a 

1.28a 

.30b 

.29b 

.36a 

.30ab 

.34a 

.37a 

K, percent 

.83ab 

.53b 

.37b 

.72a 

.54a 

.35a 

.21 ab 

.14a 

.13a 

.17a 

.19a 

.10a 

Mg, percent 

.14b 

Roots 1-1 - .25a .25a .27a .38a .36ab .28b .14a .llb .12ab 

1-2 - .2& .31 b .34a .29a .31 a .27a .12a .12a .13a 

1-3 - .26b .29ab .33a .31 a .29a .34a .I 5a .lla .14a 

Table 4.--Continued 

Tree Tree Component WT-4 WT-10 WT-20 WT-4 WT-10 WT-20 WT-4 WT-10 WT-20 

component age age class 

Year Year Mn, percent Zn, mglkg Cu, mglkg 

Needles 1-1 O+ 1 .21 a .15b .llb 222a 272a 244a 16a 12b llb 

1-2 0 .27a .18b .12c 116b 163a 189a 3a 2a 3a 

1 +2 .30a .18b .12c 2296 31 2a 297a 4ab 3b 5a 

1-3 0 .41a .25b .12c 128a 125a 141a 10a 9a 8a 

1 .54a .30b .15c 205a 21 7a 238a 10a 9a 8a 

2+3 .48a .28b .13c 269a 282a 292a 10a 8b 8b 

I Shoots 1-1 O+ 1 .07a .05b .03c 120a 139a 127a 17a 15b 12c 

1-2 0 .10a .07b .03c 72b 10la 1 05a 8a 8a 7a 

1 +2 .09a .06b .03c 93b 118a 95b 8a 8a 6b 

1-3 0 .14a .lob .04c 99a 95a 92a 10a 7b 6b 

1 .14a .08b .03c 99a 107a 94a 7a 5a 7a 

2+3 .12a .07b .03c 101a lOla 90a 7a 7a 10a 

Roots 1-1 - .06a .04b .03c 114b 155a ll8b 19a 18ab 16b 

1-2 - .08a .05b .02c 116a 145a 134a 17a 14b 9c 

1-3 - .lla .08b .03c 126a 130a 125a 17a 13b 13b 

I 

Table 4.-Continued 

Tree Tree Component WT-4 WT-10 m-20 WT-4 WT-10 WT-20 WT-4 + WT-10 + WT-20 

comoonent aae aae class laveraae) 

Year Year Fe, mglkg B, mglkg Al, mglkg 

Needles 1-1 0+1 181ab 21 5a 120b 56a 61a 66a 270 (1 32) 

1-2 0 95b 90b 183a 54b 56b 66a 238 (1 17) 

1 +2 270b 155b 465a 65a 69a 75a' 481 (1 77) 

1-3 0 145a 135a 164a 67a 60a 64a 268 (41) 

1 160b 252a 222ab 76a 72a 69a 389 (59) 

2+3 41 5b 437a 446a 84a 74a 84a 667 (66) 

Shoots 1-1 0+1 145a 92ab 55b 15a 14a 14a 108 (58) 

1-2 0 119a 48b 54b 14a 13ab 12b 96 (1 6) 

1 +2 99a 86a 80a 12b 15a llc 144 (21 

1-3 0 69a 70a 77a 13a 13a lla 116 (1 5) 

1 74a 81 a 68a 13a 12a 10a 123 (1 6) 

2+3 120a 165a 86a 8a 8a 9a 204 (43) 

Roots 1-1 - 1839a 1 096a 121 4a 24a 21ab 16b 1481 

- (225) 

1-2 4660a 1 059b 1 029b 32a 23b 17b 2257 

- (348) 

1-3 5967a 1387b 9961, 32a 19b 16b 2916 (594) 

'Standard deviations are in parentheses. 

'Letters designate difference due to soil drainage conditions during the same year at the 0.05 level using Duncan's New Multiple Range 

Test. 
Table 5.-Summary of analysis of variance showing significant effect for nutrient concentration in black spruce growing in soil with three 

different water table levels - - 

Source of 

variation Ash N P Ca K Ma Mn Fe Al Zn B Cu Na 

Water level (W) 

Age (4 

lnteraction (WxA) 

Water level (W) 

Age (A) 

lnteraction (WxA) 

Water level (W) 

Age (A) Interaction (WxA) 

"F'' values 

NEEDLES 

30** 249** 5** - 20** 44'* 22** 19** 

- 12** 4- 3- 

SHOOTS 

12" 392" 13** 13*' 

19*' 36** 4** 16** 

2* 8** 3** 2* 

ROOTS 

5'* 196** 96** 8** 

- 46** 1P 55" - 14** 19** - 

'Significant at 5-percent level "Significant at 1 -percent level 

NEEDLES 

SHOOTS T 

TOTAL 

TREE 

ABOVE 

GROUND 

T 

Figure 1 .-Mean diameter, height, and component biomass of black spruce as related to 

water table level. 
significant differences in concentrations of P were found due 

to water table level. For Zn, the differences were significant 

but did not follow any trend. 

Concentrations of ash and all mineral elements except Fe 

and Cu varied with the age of seedling shoots. The overall 

mean concentrations of ash, Ca, Al, Fe, and Zn increased 

and those of N, P, K, Mg, and B decreased with increasing 

shoot age. The increases of Cu and Mn concentrations were 

somewhat inconsistent in respect to component age class. 

The values were also affected by wood-bark ratio, which 

was not quantified in this study, for respective tree ages. 

Roots. Along the gradient of water table depth from 4 to 20 

cm, the mean concentration of N and Ca in roots significantly increased and that of ash, Mn, B Fe, Al, and Cu significantly decreased. The values for P, Mg, and Zn were 

significantly different but did not follow any trend. Concentrations of K did not significantly differ between three water 

levels. 

Mean concentration of ash, Fe, Mn, Na, and A1 significantly 

increased and N, P, and Cu significantly decreased with 

increasing tree age. The overall concentrations of K, Mg, Zn, 

and B did not significantly differ between 1 -, 2-, and 3-yearold trees. 

Changes in element concentrations in trees growing under 

different soil water table levels were consistent with results 

reported in the literature. This is due to an aerobic to anaerobic gradient in soil conditions with progressively poorer 

soil drainage. It is not surprising that in the absence of oxygen nutrient absorption by roots in the majority of the trees 

is significantly curtailed, because roots grow very poorly 

in waterlogged soils. It appears that black spruce has some 

degree of tolerance to high water tables, but the mechanism 

is unknown to the authors. 

The differences in inorganic element concentrations that 

occurred with plant age were. consistent with the literature. 

Concentrations of Ca rapidly increased between current and 

older tree components. On the other hand, concentrations 

of N, P, K, and some micronutrients decreased sharply between the first and second growing seasons. This decrease 

was partly due to dilution by tree growth as well as due to 

fertilization of seedlings prior to the first growing season 

in the greenhouse, with nutrient reserves by the end of the 

first year likely being exhausted. 

Comparisons with Published Data 

To examine the relationships among nutrients in foliage, 

studies by lngestad (1 967), Weetman (1 968), and Adams 

and Allen (1 985) utilized nutrient proportions, expressing all 

nutrient concentrations relative to N concentration which is 

taken to be 100 percent. Consequently, we compared our 

data on current year foliage with those suggested by Ingestad (1 967) for optimum nutrient proportions, with the levels 

given by Weetman (1968) for untreated mature black spruce 

stands and by Swan (1 969) for levels sufficient for good to 

very good growth of black spruce seedlings (Table 6). 

Table 6.--Comparisons among current year's foliar nutrient concentrations and proportions of black spruce 

reported by several authors 

Current study lngestad (1 966, Swan (1 969) Weetman (1 968) 

Nutrient Cont. 1967) (Seedlings, (seedlings, for (mature stand 

Seedlings WT-4 WT-10 WT-20 optimum proportion) good growth) untreated) 

N 

Percent 1.43 0.69 0.66 1.10 - 1.50-2.50 0.94 

Proportion 100 100 100 100 100 100 100 100 

P 

Percent 0.27 0.08 0.08 0.10 - 0.18-0.30 0.19 

Proportion 19 12 12 9 8-1 5 12-1 2 20 

K 

Percent 1.08 0.62 0.62 0.70 - 40-80 0.53 

Proportion 76 90 94 64 50-1 00 27-32 56 

Mg Percent 0.13 0.14 0.13 0.17 - 0.1 2-0.25 0.14 

Proportion 9 20 20 15 5-10 8-1 0 15 

Ca 

Percent 0.71 0.71 0.68 0.84 - 0.1 5-0.40 0.36 

Proportion 50 103 103 76 5-1 0 10-1 6 38 
Data in Table 6 show that the foliage of containerized seedlings prior to planting exhibited sufficient levels of N, P, and 

Mg for good to very good growth, and those of K and Ca are 

considered to be far above the optimum levels. 

In planted seedlings, the relationships in the current year's 

foliage during the second and third growing seasons drastically changed. Concentrations of N on plots with 4 and 10 

cm to the water table decreased to the levels of acute deficiency, and those with 20 cm to the water table decreased 

to the levels of moderate deficiency. Concentrations of P 

dropped to the levels of 'acute deficiency on all plots. Concentrations of K and Mg remained at levels sufficient for 

good to very good growth, and those of Ca remained at luxury consumption levels. 

With decreasing N concentration, there was an increase in 

relative concentration of K, Mg, and Ca. Because of the 

decrease in absolute concentration of P, a decrease in relative levels has to be expected. 

Comparisons of nutrient levels reported in this study for 

branches, shoots, and roots were not made because data 

on macro-nutrient levels in black spruce components other 

than the current year's foliage is nonexistent. 

Furthermore, comparisons of nutrient levels reported in this 

study with those required for adequate or optimum growth 

cannot be made because information on macro-nutrient 

levels in black spruce components, other than the current 

year's foliage, is also nonexistent. Therefore, until furtner 

data become available it seems reasonable to accept the 

levels and ranges given by Young and Carpenter (1967) for 

branches, stems, and roots of red spruce, by McKee and 

others (1 984) for 2-year-old loblolly pine (Pinus taeda L.) 

grown under flooded and drained soil conditions, and by 

Czapowskyj (1979) for balsam fir as affected by soil drainage. Elemental levels in the foliage, shoots, and roots reported in the current study are generally comparable, and 

the trends are consistent with those reported by authors 

referred to above for the respective tree components of 

other coniferous species. 

Conclusions 

The following conclusions can be drawn from these results: 

Containerized seedlings received adequate nourishment 

for good to very good growth during the initial growth 

period. 

Survival of black spruce seedlings was not affected by 

water table depths of 4, 10, and 20 cm during a period of 

3 years following transplanting. 

An increase of the water table depth below the mineral soil 

surface significantly enhanced the growth and biomass 

production of the foliage, shoots, and roots. 

As the depth to the water table increased: (a) the levels of 

N increased and those of Na, Mn, Cu, and Al decreased 

in foliage, shoots, and roots; and (b) the levels of P, K, Zn, 

Fe, and B increased only in the foliage and Mg increased 

in both foliage and shoots. 

A positive response in black spruce growth would be 

expected if plantations were established on sites where 

the water table could be maintained to a depth of at least 

20 cm. 

Literature Cited 

Adams, M. B.; and Allen, H. L. Nutrient proportions in 

foliage of semi-mature loblolly pine. Plant and Soil. 

86:24-34; 1985. 

Burton, J.D. Prolonged flooding inhibits growth of loblolly pine seedlings. Res. Note SO-124. New Orleans, 

LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station; 1971. 4 p. 

Cannell, R.Q.; Jackson, Michael B. Alleviating aeration 

stresses. In: Arkin, G. F.; Taylor, H. M., eds. Modifying 

the root environment to reduce crop stress. Monog. 

A.S.A.E. 1981 :141-192. 

Coutts, M.P. Effects of waterlogging on water relations of 

actively-growing and dormant sitka spruce seedlings. 

Annals of Botany 47:747-753; 1981. 

Czapowskyj, M. Foliar nutrient concentrations in balsam 

fir as affected by soil drainage and methods of slash 

disposal. Res. Note NE-278. Broomall, PA: US. Department of Agriculture, Forest Service, Northeastern Forest 

Experiment Station; 1979. 4 p. 

Czapowskyl, M. M. Response of a young black spruce 

stand to soil draining control of competition and fertilization. Agronomy Abstracts. 1982:263. 

Ingestad, T. Methods for uniform optimum fertilization of 

forest tree plants. Proc. IUFRO Cong. 3, 265-269; 1967. 

McKee, W.H., Jr.; Shoulders, E. Slash pine biomass response to site preparation and soil properties. Soil 

Science Society of America Proceedings. 38:144-148; 

1974. 
McKee, W.H., Jr.; Hook, D.D.; DeBell, D.S.; Askew, J.L. 

Growth and nutrient status of loblolly pine seedlings 

in relation to flooding and phosphorus. Soil Science 

Society of American Journal. 48:1438-1441; 1984. 

Mueller-Dombois, Dieter. Effect of depth to water table on 

height growth of tree seedlings in a greenhouse. Forest Science. 10:306-316; 1964. 

I Norby, Richard J.; Kozlowski, T.T. Flooding and SO, stress 

interaction in Betula papyrifera and 6. nigra seedlings. 

Forest Science. 29(4):739-750; 1983. 

Swan, H. Stewart D. Relationships between nutrient supply, growth and nutrient concentrations in the foliage 

1 of black spruce and jack pine. Woodlands Rep. No. 

19. Ottawa, ON: Pulp and Paper Research lnstitute of 

Canada; 1969.46 p. 

Weetman, G.F. The nitrogen fertilization of three black 

1 spruce stands. Woodlands Rep. No. 6 Ottawa, ON: Pulp 

and Paper Research Institute of Canada; 1968. 45 p. 

Young, Harold E.; Carpenter, Paul M. Weight, nutrient element and productivity studies of seedlings and sap 

lings of eight tree species in natural ecosystems. 

Orono, ME: University of Maine Life Science and Agriculture Experiment Station; 1967; Tech. Bull. No. 28. 39 p. 

~ Acknowledgment 

We thank the Great Northern Paper Company, Millinocket, 

Maine, for donating the seedlings; we also thank George R. 

Benoit for his help in planning the study. 
Czapowskyj, Miroslaw M.; Rourke, Robert V.; Grant, Walter J. Growth and 

nutrient status of black spruce seedlings as affected by water table depth. 

NE-RP-591. Broomall, PA: U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station; 1986. 9 p. 

The objective of this study was to determine the effect of different soil water 

table levels on growth, biomass production, and nutrient accumulation in 

black spruce seedlings growing under greenhouse conditions over three 

growing seasons after transplanting. 

I ODC 181.32 Picea mariana (Mill) B.S.P. 

Keywords: black spruce biomass, water table, nutrients 
Headquarters of the Northeastern Forest Experiment Station are in Broomall, Pa. 

Field laboratories are maintained at: 

Amherst, Massachusetts, in cooperation with the University of Massachusetts. 

Berea, Kentucky, in cooperation with Berea College. 

Burlington, Vermont, in cooperation with the University of Vermont. 

Delaware, Ohio. 

Durham, New Hampshire, in cooperation with the University of New Hampshire. 

Hamden, Connecticut, in cooperation with Yale University. 

Morgantown, West Virginia, in cooperation with West Virginia University, 

Morgantown. 

Orono, Maine, in cooperation with the University of Maine, Orono. 

Parsons, West Virginia. 

Princeton, West Virginia. 

Syracuse, New York, in cooperation with the State University of New York 

College of Environmental Sciences and Forestry at Syracuse University, 

Syracuse. 

University Park, Pennsylvania, in cooperation with the Pennsylvania State 

University. 

Warren, Pennsylvania. 

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