Science for Solutions 

NOAA COASTAL OCEAN PROGRAM 

Decision Analysis Series No. 4 

MARINE EUTROPHICATION REVIEW 

PART I: QUANTIFYING THE EFFECTS OF NITROGEN ENRICHMENT 

ON PHYTOPLANKTON IN COASTAL ECOSYSTEMS 

PART 2: BIBLIOGRAPHY WITH ABSTRACTS 

Kenneth R. Hinga 

Heeseon Jeon 

Noelle F. Lewis 

January 1995 

U.S. DEPARTMENT OF COMMERCE 

National Oceanic and Atmospheric Administration 

Coastal Ocean Office 
Hinga, Kenneth R., Jeeseon Heon and Noelle F. Lewis. 1995. Marine Eutrophication Review--Part 1 : 

Quantifying the Effects of Nitrogen Enrichment on Phytoplankton in Coastal Ecosystems; Part 2: 

Bibliography with Abstracts. NOAA Coastal Ocean Program Decision Analysis Series No. 4. NOAA 

Coastal Ocean Office, Silver Spring, MD. Part 1, 36 pp.; Part 2, 120 pp. 

This publication does not constitute an endorsement of any commercial product or 

intend to be an opinion beyond scientific or other results obtained by the National 

Oceanic and Atmospheric Administration (NOAA). No reference shall be made to 

NOAA, or this publication firmshed by NOAA, in any advertising or sales promotion 

which would indicate or imply that NOAA recommends or endorses any proprietary 

product mentioned herein, or whch has as its purpose an interest to cause directly or 

indu-ectly the advertised product to be used or purchased because of thls publication. 
Note to Readers 

The NOAA Coastal Ocean Program (COP) provides a focal point through which the agency, 

together with other organizations with responsibilities for the coastal environment and its 

resources, can make significant strides toward finding solutions to critical problems. By working 

together toward these solutions, we can ensure the sustainability of these coastal resources and 

allow for compatible economic development that will enhance the well-being of the Nation now 

and in fbture generations. The goals of the program parallel those of the NOAA Strategic Plan 

for 1995-2005. 

A specific objective of COP is to provide the highest quality scientific information to coastal 

managers in time for critical decision making and in a format useful for these decisions. To help 

achieve this, COP inaugurated a program of developing documents that would synthesize 

information on issues that were of high priority to coastal managers. To develop such documents, 

a three-step process was used: 1) to compile a list of critical topics in the coastal ocean through 

a survey of coastal resource managers and to prioritize and select those suitable for the document 

series through the use of a panel of multidisciplinary technical experts; 2) to solicit proposals to 

do research on these topics and select principal investigators through a rigorous peer-review 

process; and 3) to develop peer-reviewed documents based on the winning proposals. 

Marine Eutrophication Review is the third title published of the seven topics and associated 

principal investigators which were selected in the initial round. A bibliography of synthesis 

documents was added to the series on the recommendation of a COP management committee and 

is in print (see inside back cover). Other volumes will be published over the next two years on the 

following topics: seagrass restoration technology, coastal watershed restoration, restoring 

streams and anadromous fish habitat affected by logging, and management of cumulative coastal 

environmental impacts. 

As with all of its products, COP is very interested in ascertaining the utility of the Decision 

Analysis Series particularly in regard to its application to the management decision process. 

Therefore, we encourage you to write, fax, call, or Internet us with your comments. Please be 

assured that we will appreciate these comments, either positive or negative, and that they will help 

us direct our fbture efforts. Our address and telephone and fax numbers are on the inside front 

cover. My Internet address is DSCAVIA@HQ.NOAA. GOV. 

' Donald Scavia 

Director 

NOAA Coastal Ocean Program 
MARINE EUTROPHICATION REVIEW 

PART I : QUANTIFYING THE EFFECTS OF NITROGEN ENRICHMENT 

ON PHYTOPLANKTON IN COASTAL ECOSYSTEMS 
TABLE OF CONTENTS 

ABSTRACT 

INTRODUCTION 

DEFINING TERMS 

APPROACHES TO FINDING A RELATIONSHIP BETWEEN 

EUTROPHICATION AND NUTRIENT INPUTS 

NITROGEN AND PRIMARY PRODUCTION IN 

MARINE ECOSYSTEMS 

PHYTOPLANKTON ABUNDANCE AND OTHER MEASURES 

THAT MAY BE RELATED TO EUTROPHICATION 

OTHER COMPLICATING ISSUES 

NITROGEN LOADINGS OR NITROGEN CONCENTRATIONS? 

RELATIONSHIPS BETWEEN NITROGREN AND EUTROPHICATION 

THE ROLE OF ANOTHER FACTOR 

CURVE FITS TO CROSS-SYSTEM AND MERL DATA AND 

COMPARISON TO THE EFFECT FACTORS FROM CHANGES 

OVER TIME 

CONCLUSIONS 

TABLE 

FIGURES 

LITERATURE CITED 
The results of three different approaches are used to investigate relationships between nitrogen 

availability and phytoplankton primary production and abundance in coastal ecosystems. The 

three approaches are controlled experiments in marine enclosures, history of changes in coastal 

ecosystems, and cross-system comparisons. Most systems can be expected to have increased 

primary production with increased nitrogen loadings or long-term average concentration. 

Nitrogen availability alone is not a very precise predictor of phytoplankton production or 

abundance. The magnitude of the change in phytoplankton production or abundance found for 

most systems is typically in the range of 1.4 to 3 times for a doubling in nitrogen loading or 

concentration. However, there are coastal ecosystems which have not followed the general 

relationships found. Until the characteristics which sets those systems apart from other 

systems can be identified, there is no guarantee that any individual system will have the typical 

response. 

Perhaps the paramount question placed before the modern ecologist is "What will happen 

if....?" As this review considers anthropogenic nutrient inputs to coastal environments, the 

question becomes: "What will happen if nutrient loads to a body of water increase?" And, 

conversely: "What will happen if (usually at great expense) nutrient inputs to a coastal body of 

water are reduced?" The answers to these questions are far from trivial to obtain. Ecology as a 

predictive science is still in its infancy. There have been remarkably few planned and 

controlled experiments at the ecosystem level to learn from, and to test the ecologist's ability to 

accurately predict changes. 

Nevertheless, marine ecologists can certainly observe present conditions in coastal ecosystems, 

and they have observed what are presumed to be nutrient-related changes in estuaries. 

Certainly some insights have been gained as to the behavior of coastal ecosystems under 

increased nutrient loadings. 

This review will consider relationships between nutrient loading and phytoplankton production 

in coastal ecosystems. Specifically, the review will examine if there is a simple relationship 

between nutrient loading and phytoplankton production, or surrogates for production, in 

coastal ecosystems. In other words, how well can we predict how much change will occur in a 

coastal ecosystem for any specific change in nutrient loading? 

Before starting on any dialogue it is useful to make sure that participants are using the same 

meanings for their vocabulary. The word "eutrophication" has had a variety of uses when 

applied to coastal ecosystems, and probably many more uses when applied to other 

ecosystems. For this review the definition of eutrophication as proposed by Nixon (1994) will 

be used. His proposed definition is: 

Eutrophication (noun) -- an increase in the rate of supply of organic matter to an ecosystem. 

The rate of supply of organic carbon may be'by either primary production by autotrophs within 

the system (autochthonous carbon) or by an input of organic matter from outside the system 

(allochthonous carbon). The central feature of the definition is the rate of supply of organic 

carbon.. At first consideration, the definition seems distant from the common informal use of 

"eutrophication" to denote an increase in anthropogenic nutrient loading (i.e., cultural 

eutrophication). However, use of Nixon's definition has considerable advantages. 
First, the definition is simple. The definition is based upon a single parameter. There is a lot 

to be said for simplicity. 

Second, the definition makes it easy to separate eutrophication from its causes and its 

consequences. The causes of eutrophication may include increased input of inorganic 

nutrients, decrease in water turbidity, change in hydraulic residence time of the water, a decline 

in grazing pressure or increase in direct inputs of organic matter. The consequences of an 

increased rate of supply of organic carbon may include changes in ecosystem community 

structure, increased rates of oxygen depletion, fish kills, etc. 

Finally, focusing on the rate of organic carbon supply as the central issue makes it a 

straightforward matter to classify the trophic status of coastal ocean ecosystems. The trophic 

status of a coastal ecosystem is simply a measure of the rate of organic carbon supply. 

Definitions of various trophic states were also proposed by Nixon (1994): 

Organic Carbon Supply 

g c M-2 year-1 

oligotrophic 1100 

mesotrophic 101-300 

eutrophic 30 1-500 

hypertrophic >500 

The trophc classification states are value neutral. The definition does not imply that one 

trophic state is better, more desirable, or more valuable than another. 

APPROACHES TO FINDING A RELATIONSHIP BETWEEN EUTROPHICATION AND 

NUTRIENT INPUTS. 

In order to quantify "what will happen in a coastal ecosystem if nutrient inputs go up or 

down," it is necessary to have or to establish a relationship between nutrient loading and rate of 

carbon supply. Carbon supply can be from two sources, primary production within the 

ecosystem and external loadings to the ecosystem. External loadings are likely to be the 

dominant source of carbon in the vicinity of sewage outfalls with primary or no treatment. In 

such areas one would not expect to find a cause-effect relationship between nutrient loadings 

and carbon supply (unless one is considering the processes and procedures used in sewage 

treatment plants themselves). 

For most open-water areas of coastal ecosystems, the carbon supply is likely dominated by 

autotrophic primary production within the system. A very useful relationship to identify would 

then be a relationship between nutrient loadings and primary production. Some care must be 

taken to make sure that coastal systems, or parts of coastal systems, with significant particulate 

loadings are avoided in the development of a nutrient loading vs. production relationship from 

empirical data. 

A number of approaches might be applied to the development of a relationship between nutrient 

loading and primary production. These include: 

1. Controlled experiments with marine ecosystems (mesocosm and enclosure experiments). 

2. Historical records of the response of coastal ecosystems to changes in nutrient loads. 
3. Cross-system comparisons of different coastal ecosystems to see if the primary production 

in those systems is systematically related to nutrient inputs (comparative ecology). 

4. Calculation (modeling) from first principles of physics, chemistry, and ecology. 

Conducting controlled experiments with representative coastal ecosystems would be the most 

direct way to determine the effects of nutrient loadings on coastal ecosystems. Such a direct 

approach was instrumental in determining the response of lakes to nutrient loadings (i.e., 

Shindler et al., 1971; Shindler 1973). Given that coastal ecosystems are considered public 

domain and we are unlikely to want to deliberately damage such an ecosystem, it seems 

unlikely that natural coastal ecosystems will be made available for direct experimentation. 

Direct experiments are possible with coastal ecosystems in miniature. This is the realm of 

enclosures and mesocosms. A variety of marine enclosures have been established to attempt to 

create ecosystems in large containers that have chemical and biological properties of coastal 

ecosystems. Descriptions of, and arguments for the use of, enclosures have been made by 

Davies and Gamble (1979), Steele (1979), Parsons (1981), Lundgren (1985), and Santschi 

(1988). It has been within the marine ecologist's ability for nearly two decades to establish 

enclosures containing ecosystems that, over long periods of time (many months), have biota 

and ecosystem function that are hard to tell apart from natural coastal ecosystems. 

It seems remarkable that given this ability there has not been a widespread use of enclosures. 

Direct experimentation is a powerful, if not the most powerful approach, in scientific 

investigation. Although there have been a number of short term (i.e., up to about a month's 

duration) nutrient experiments in marine enclosures, there has been only one long-term 

eutrophication experiment in marine enclosures. There was a 28-month nutrient-addition 

experiment at the Marine Ecosystems Research Laboratory of the University of Rhode Island 

(Nixon et al., 1984; Kelly et al., 1985; Oviatt et al., 1986a, 1986b, 1989; Keller 1988, 1989; 

Keller & Rice 1989; Hinga 1990). Results of that experiment will appear in the analyses later 

in this paper. 

One might expect it would be possible to learn a great deal from observations on the changes 

that have taken place in coastal ecosystems over time. However, the number of coastal 

ecosystems where adequate measurements exist under two different loading conditions is 

remarkably small. While current conditions and loadings are known in a good number of 

coastal ecosystems, records of conditions under earlier loadings and records of earlier loadings 

are relatively rare. Coastal ecosystems for which adequate data have appeared in the literature 

will be considered below. 

A comparative ecology approach was used by Volenweider (1976) to develop a quantitative 

relationship between nutrient loadings in lakes and the abundance of lake phytoplankton. The 

results of his comparative study demonstrated a clear relationship between phosphorus 

loadings to lakes and the phytoplankton standing stock in lakes. Comparative approaches have 

been applied to marine ecosystems by Boynton et al. (1982), Nixon (1983), Nixon and Pilson 

(1983) and Monbet (1992). 

A modeling approach, while a very desirable goal, is likely out of reach for the near future. 

Modeling the physics and chemistry of a coastal ecosystem on first principles may be currently 

possible. Unfortunately, the first principles of ecology are not yet sufficiently understood to 

permit useful predictio~s. At the very least, an ecosystem model requires the ability to predict 

when and why different species appear at various times. This ability is surely not yet achieved, 

even in an unperturbed (by human activities) system. The ability to predict how communities 

will change under novel conditions is even further off. 
There are those who might argue that a functional model of the ecosystem is all that is needed 

for prediction of at least some of the consequences of changes in nutrient loadings. In this 

approach, specific species are not modeled. Related groups of species are assumed to have the 

same function (nutrient dynamics, etc.). Functional modeling will certainly provide insights 

into ecosystem functioning and perhaps eventually into effects of external changes on 

ecosystems. Functional modeling is a valuable tool of the ecologist, but at present is more 

likely to be useful as a diagnostic than a predictive tool. In order to use functional models to 

provide predictions of response to change, it is necessary to know the coefficients, rates, and 

equations that describe the functioning of the communities in the modeled system before and 

after change in the parameters of interest. As noted above, it is not yet possible to confidently 

predict which populations will comprise communities after a change in conditions. Unless one 

can demonstrate that all coastal marine communities that could exist in a particular area are 

functionally similar, or one can demonstrate how the functionality will change, it is 

inappropriate to expect functional models to be reliable predictors of change. 

Which of the possible approaches should be used to gain an understanding of the relationship 

between nutrient loadings and the eutrophication of coastal ecosystems? The best answer is 

probably "all of the above." When different independent approaches to the problem provide 

the same results, then marine ecologists can start to have confidence that they have gained a 

defensible ability to predict the response of coastal ecosystems to changes in nutrient loadings. 

The history of lake eutrophication investigations provides a good example of the value of 

having support from independent approaches. In the 1960's there was a great deal of 

discussion and debate concerning the cause(s) of conspicuous thick algal blooms and algal-mat 

growths appearing in many lakes. Were the mats and the blooms a result primarily of 

phosphorus additions or of other agents? When the results of a direct experimental approach 

(Shindler 197 1, 1973) and a comparative approach (Vollenweider 1976) both clearly implicated 

phosphorus as the primary causative agent, a firm basis was provided for the regulation of 

phosphorus in detergents and other sources. 

NITROGEN AND PRIMARY PRODUCTION IN MARINE ECOSYSTEMS 

The nutrient central in discussions of coastal eutrophication is nitrogen. Early in the relatively 

brief history of concern about marine eutrophication, Ryther and Dunstan (197 1) demonstrated 

that additions of nitrogen stimulated growth of marine phytoplankton. Phosphorus additions 

had little if any effect. Nutrient limitation studies have been conducted many times since (e.g. 

Goldman, 1976; Laws and Redalje, 1979; GranCli and Sundback, 1985; D'Elia et al., 1986; 

GranCli et al., 1986; and GranCli et al., 1988, GranCli et al., 1990; Montgomery et al., 1991; 

Oviatt et al., 1994). In general, additions of nitrogen alone, as either nitrate or ammonium, 

promote enhanced growth of phytoplankton (also see the review by Howarth, 1988). 

However there are times when phosphorus limitation is found in environments usually limited 

by nitrogen (e.g. GranCli et al., 1990) and marine environments where phosphorus limitation is 

more common than nitrogen limitation (Myers and herson, 198 1). Silica limitation, resulting 

from nitrogen and phosphorus enrichment, may also have an important influence on the 

composition of phytoplankton communities. 

With the general result that nitrogen is usually the limiting nutrient in coastal ecosystems, and 

provided that long-term responses (i.e., yearly averages) are investigated, it seems reasonable 

to attempt to find relationships between nitrogen and eutrophication, without consideration of 

phosphorus. This assumption has been made in many previous studies, and will be adopted 

here. Should it not be possible to find a satisfactory relationship(s) between nitrogen and 

eutrophication, it may be necessary to retreat and consider phosphorus, and other factors. 
PHYTOPLANKTON ABUNDANCE AND OTHERMEASURES WHICH MAY BE 

RELATED TO EUTROPHICATION 

Primary production becomes the basis of the definition, as given above, of eutrophication in 

coastal systems where external carbon loadings are not significant. Unfortunately, primary 

production is not measured very regularly in many coastal ecosystems. It would be useful if 

surrogate measures related to primary production based upon a more regularly measured 

parameter could be found. This would expand the number of systems available for 

investigation of nitrogen-eutrophication relationships. 

The primary production of phytoplankton is dependent, in part, upon the abundance of the 

phytoplankton. If phytoplankton is of low abundance, there are few "machines" to do the 

work of production and the total production cannot be high. Hence phytoplankton abundance 

and production may be expected to be somewhat related. Measurement of the standing stock of 

phytoplankton is conducted much more often than measurement of production. The standing 

stock of phytoplankton carbon is usually estimated by measuring chlorophyll-a and assuming a 

fixed carbon to chlorophyll-a ratio. 

The instantaneous (i.e., short-term) depth-integrated primary production (i.e., total production 

under 1 m2 of surface) in marine systems can be estimated very well by knowing the 

phytoplankton abundance, the depth of the photic zone (as calculated from the light extinction 

coefficient), and the amount of incident light (Cole and Cloern, 1987; Keller, 1988, 1989). 

Hence, if ambient light and turbidity were the same for all coastal ecosystems, chlorophyll-a 

alone could be used to predict primary production. 

Ambient light availability varies by latitude and local climate. Nixon (1983) assembled light 

data for a variety of coastal embayments and found a modest range in annual average ambient 

light ranging from 329 Ly day-I (Narragansett Bay, and New York Bay) to 461 Ly day-1 (San 

Francisco Bay). The effect of local climate was larger than the latitude effect (30 ON to 41 ON in 

North America). 

Light extinction also varies in coastal systems. Data also assembled by Nixon (1983) showed 

attenuation coefficients ranging from about 1 m-l to 8 m-1 with most of the high values at low 

salinity. (This translates to 1 % light levels at about 1 to about 10 meters depth.) Of course, a 

major determinant of light level in the water column is the amount of phytoplankton material 

itself. 

The abundance of phytoplankton is not always measured at multiple depths. It is more 

common to find measurements of chlorophyll a, and of primary production made only on 

near-surface samples. It is far less useful, from a whole ecosystem point of view, to consider 

possible relationships between phytoplankton abundance and primary production for just nearsurface samples. It is the total carbon input to a coastal ecosystem that is relevant to 

eutrophication as defined here. How well the total water column production can be estimated 

from only near-surface measurements will not be considered in detail here. 

If cross-system comparisons and consideration of changes in coastal ecosystems over time 

were restricted to only those systems for which there is a good record of areal production, there 

would not be much data to consider. In order to attempt to find relationships between nutrients 

and eutrophication, measures of near-surface phytoplankton abundance, and near-surface 

primary production will be considered. It will need to be kept in mind that mixing the various 

types of measures of phytoplankton may obscure relationships for any single parameter. 
OTHER COMPLICATING ISSUES 

Phytoplankton abundance is often estimated from measurement of chlorophyll-a. It should be 

noted that the carbon-chlorophyll ratio is not a constant (e.g., Falkowski, 1980). More 

importantly, it appears that carbon to chlorophyll-a ratios vary systematically with 

eutrophication (Nixon, 1992). The carbon to chlorophyll a ratio in the MERL eutrophication 

experiment, referred to above, decreased by a factor of three from controls to the higher 

enriched tanks. It seems probable that this is a result of shade adaptation and not a result of 

changes in species composition (Oviatt et al., 1989). Such variability may interfere with 

efforts to find a relationship between phytoplankton abundance and eutrophication. 

Phytoplankton abundance may also be derived from direct counts of phytoplankton cells. A 

change in cell count will not be directly proportional to change in biomass if there has been a 

change in average cell size. In one of the ecosystems that will be considered later (Helgoland 

Reede), cell counts were coupled with the volume per cell to provide a better estimate of 

biomass change over time than raw cell counts. 

Some comment must also be made concerning primary production measurements themselves. 

There are a variety of basic techniques used to measure primary production and variability in 

the practice of techniques based upon the same principle. Even the use of a single basic 

technique, such as 14C-based production measurements, the depths of water samples, bottle 

size and material, bottle agitation, and how bottles are incubated, are far from uniform in 

practice. Such variability in technique undoubtedly affects results and may make comparisons 

between different systems misleading. 

Should a relationship be expected between eutrophication and nitrogen loading, or between 

eutrophication and the achieved in-system nitrogen concentration (resulting from changes in 

nitrogen loadings and the other processes affecting nitrogen concentrations)? A priori, there is 

no way to determine which will be a more fruitful approach. Ideally, a description of the 

availability of nitrogen is desired. It is not safe to assume that either inorganic nitrogen 

concentrations or loadings of nitrogen adequately represent nitrogen availability. 

Where measured nitrogen concentrations are used for a basis for a relationship, a long-term 

average concentration of nitrogen must be used. In short time scales, the relationship between 

nitrogen concentrations and phytoplankton abundance, may be inversely related, reflecting 

short-term uptake by blooms. Further, consideration of just the dissolved species of nitrogen, 

or dissolved inorganic nitrogen (DIN; nitrate plus nitrite, plus ammonia), may not represent the 

real availability of nitrogen to the phytoplankton. Some portion of the particulate and dissolved 

organic nitrogen may be subject to rapid remineralization and hence be biologically available. 

If the fraction of the biologically available nitrogen in organic matter is variable in different 

coastal ecosystems, use of DIN only may not provide clear relationships between nitrogen and 

eutrophication. 

If loadings are used as a basis for a relationship, do changes in loadings necessarily reflect a 

corresponding change in availability of nitrogen to phytoplankton? Should loadings calculated 

on a volume basis or loadings on an areal basis be used to represent availability? How does 

flushing rate (or the inverse, the residence time of water) affect the achieved availability of 

nitrogen in the coastal ecosystem? The later question is addressed briefly as follows. 

For a conservative (non-reactive) material introduced into an ideal well-mixed system, 

loadings and achieved concentrations would be directly proportional. In a steady state 
(constant loadings and concentration not increasing) a mass balance can be set up for a 

conservative material where material entering the system must equal the material leaving the 

system (or else the concentration of the material would either be rising or falling). Expressed 

in mathematical shorthand: 

Min is the loading, at a constant rate, of the material M. (This assumes the offshore 

concentration is negligible.) Mout depends upon how much water is exchanged per unit of 

time between the system and more offshore waters. 

Mout = volume of water lost per day * Concentration of M in water 

[Following the units: g/day=liters/day * g/liter] 

As the flushing of a coastal system does not depend upon a material like nitrogen dissolved in 

the water, the volume of water lost per day is constant (long-term average). Hence, in a single 

ideal system, the achieved concentration of a conservative material is directly proportional to 

the loadings. A doubling of a loading would lead to a doubling of achieved concentration, 

once a new steady-state is achieved. (This requires a period of time equal to about three times 

the residence time of the water in the system.) A similar exercise can be done for nonnegligible offshore concentrations, where the net result is that the net difference between the insystem concentration and the offshore concentration is proportional to loading. 

Nitrogen is not a conservative material. The forms of nitrogen undergo many reactions. As 

nitrogen enters a coastal ecosystem from the landward side and passes through a coastal system 

to the open ocean, it may be taken up from dissolved forms into organic matter and 

remineralized (both in the water column and at the benthos) a number of times. In addition 

there can be essentially permanent losses of nitrogen, through long-term burial in sediments 

and denitrification (the conversion of fixed nitrogen to nitrogen gas). While there are very few 

very well constrained nitrogen budgets for coastal ecosystems, it appears that burial is 

generally a small term. Denitrification on the other hand may be quite significant and quite 

variable. In a fast-flushing estuary, there may be little time for denitrification. In a system 

where water has a long residence time, denitrification may account for a major loss. For 

example, it is estimated than 90% of the nitrogen introduced into the Baltic Sea is denitrified 

(Larsson et al., 1985). 

The net result is that in coastal ecosystems with fairly short residence times, the achieved 

concentrations of nitrogen in the system will be at least approximately proportional to changes 

in nitrogen loadings. Even in systems where there is significant denitrification, if the fraction 

of nitrogen lost to denitrification is approximately the same at different loadings, then the 

proportionality between changes in concentration and changes in loading will hold. 

For systems where there is a long residence time of water, it is probable that the relationship 

between nitrogen loadings and concentrations will not be proportional. This is especially true 

where the relative concentrations of the nitrogen species may have also changed over time. A 

higher fraction of nitrate in the inputs (as may happen when sewage treatment plants convert 

more ammonia to nitrate to reduce the oxygen demand of their effluent) may be expected to 

alter the fraction of nitrogen lost to denitrification. (Nitrate is the nitrogen species converted to 

nitrogen gas during denitrificaion.) 
RELATIONSHIPS BETWEEN NITROGEN AND EUTROPHICATION 

There have been a few studies attempting to determine a general relationship between nitrogen 

and eutrophication, or properties related to eutrophication. As might be expected from the 

discussions above, a variety of different measures of nitrogen availability have been used. 

Similarly, different measures of phytoplankton abundance and production have been 

considered. The reader is advised to note the exact parameters plotted on each graph. Many of 

the graphs appear similar but are of different parameters. In addition, some of the plots (those 

generated new for this report) are provided in two versions showing the same data, a log-logaxes version and a linear-axes version. The perspective obtained from inspection of plots may 

be influenced by the log or linear construction of the plot. The dual versions are provided to 

give multiple perspectives. 

This section will look at some previous efforts to find relationships between nitrogen and 

eutrophication from comparative studies and mesocosm experimentation. A new analysis of 

the changes in coastal ecosystems over time will also be presented. In order to compare the 

results obtained from the different measures of nitrogen availability and of phytoplankton 

abundance and production, a factor-of-change of effect (abundance or production) per factor of 

change in nitrogen (loading or DIN concentration) will be calculated. For example, it may be 

found that in a system where the nitrogen loading doubled over time, the phytoplankton 

abundance went up by a factor of 1.5. It may eventually be shown that it is inappropriate to 

'compare the results of abundance changes with production changes. For now, the approach 

allows a quantitative look at eutrophication from a variety of types of information. 

Nixon (1992) prepared plots of primary production and chlorophyll-a against DIN input (both 

on an areal and volume basis) for a variety of marine systems ranging from the open ocean to 

heavily nutrient-loaded estuaries (Figures 1 to 4). Both the production and abundance of 

phytoplankton correlate with nitrogen loadings. 

In viewing Figures 1 to 4 it should be kept in mind that the plots are log-log plots. Hence the 

scatter of points is a significant magnitude. If a more limited range is examined, the 

significance of the scatter becomes evident. Nixon and Pilson (1983) and Nixon (1983) 

prepared cross-system plots for estuaries only. Their results are replotted in Figure 5. In these 

graphs, the correlation between nitrogen and production is not so evident. Two estuaries with 

the same DIN input rate may have up to a factor of five difference in production. 

The results of two other cross-estuary approaches appear in Figures 6 and 7. Figure 6 shows 

an effort similar to that of Nixon and Pilson (1983) except that the estuaries were broken into 

segments (typically 4 segments), so as not to average high and low nutrient portions of 

estuaries, and the segments were plotted individually. Nixon and Pilson (1983) used wholeestuary averages. Figure 7 shows an annual average DIN vs. annual average chlorophyll plot 

for individual stations in a variety of estuaries (from Monbet, 1992). 

While there may be a general trend of increasing phytoplankton abundance with increasing 

nitrogen availability, nitrogen alone is an imprecise predictor for a public official faced with a 

decision as to whether the benefit will be worth the expense to reduce, for example, the 

nitrogen inputs to an estuary by 30%. Estuaries, portions of estuaries, and individual stations 

in estuaries with similar nitrogen loadings or DIN concentrations may have quite different (at 

least from a practical management perspective) phytoplankton abundance and production. 

From a historical perspective, examination of a portion, limited to high nutrient levels, of the 

Vollenweider plot of corrected phosphate concentrations vs. chlorophyll (which was an 
influential milestone in lake management), on a linear plot (Figure 8) shows scatter much like 

that in Figures 5 and 6. While the general principle of phosphorus control on lake trophic 

status was strongly supported by the original Vollenweider relationship, the relationship 

established also did not have the precision to accurately (i.e., within a factor of 2) predict the 

chlorophyll abundance in any individual lake. 

The scatter in the cross estuary studies begs the question of how have individual coastal 

ecosystems responded to changes in nitrogen. Will each individual coastal ecosystem respond 

to nitrogen additions with an increase in phytoplankton abundance or production? If so, would 

all ecosystems have the same magnitude of response for the same change in nitrogen loading or 

concentration? Or, do coastal ecosystems of vary in sensitivity to changes in nitrogen? 

What has been learned from controlled experiments? A 28-month eutrophication experiment 

was conducted in the MERL marine enclosures using 9 enclosures. Three of the enclosures 

were kept as controls and 6 enclosures received daily additions of nitrogen, phosphorus, and 

silicate to create a gradient of loading. Figures 9, 10, and 11, show that measures of 

phytoplankton abundance and production increase reasonably steadily with increasing nitrogen 

loading or average concentration. The variability about the trend appears to result primarily 

from influence of the benthlc communities (which developed into quite different communities 

in the different enclosures) on the phytoplankton in the water column. Nevertheless, at least 

for this one environment, increases in nutrient loading resulted in eutrophication as defined 

earlier. 

What has been the record of changes in coastal ecosystems over time? Table 1 provides details 

on a number of systems where a history of the conditions in the system is reasonably well 

constrained. For most systems, the available parameters are phytoplankton abundance (either 

as directly-measured chlorophyll or as calculated chlorophyll from cell counts) and nitrogen 

concentrations. The measured average abundance at two nitrogen concentration levels (usually 

near the begining and end of the reported period of changes in nitrogen concentration) are 

plotted in Figure 12. While many of the trends for individual systems combine to create an 

overall trend consistant with those in Figures 1-4, is clear that the slopes of the trends are not 

all parallel. For two of the systems, the change in abundance decreases with higher nitrogen 

concentrations, running counter to the expected overall trend. 

Additional systems and production data can be considered by calculating a normalized 

parameter for each system. The parameter calculated here is the magnitude of change of 

abundance or production that is calculated to result from a doubling in nitrogen concentration or 

loading. A calculated "change factor" of 2, for example, indicates that there was a doubling in 

abundance or production for a doubling in nitrogen loading or concentration. A change factor 

of 1 would indicate that there was no change in abundance or production with a change in 

nitrogen concentration or loading. A change factor of less than one indicates that there was a 

decrease in abundance or production while there was an increase in nitrogen concentration or 

loading. (See the legend for Figure 13 for details on the method of calculation.) The values 

for the magnitude of the effect are plotted against the average DIN concentration for the system 

(i.e., midpoint of the system's DIN concentrations). 

Changes observed in different systems for a doubling in nitrogen ranged from a value of 0.60 

to 6.99. The values 0.60 for a region of the Upper Chesapeake indicated that there was a 

decrease in abundance of phytoplankton over a period in which there was increasing DIN 

concentrations. Similarly, the value of 0.64 for the Upper Pamlico, indicated that there was an 

increase in abundance of phytoplankton over a time period in which there was decreasing DIN 

concentrations. Most of the calculated change factors fall between about 1.1 and 3. It is 

interesting that for the three stations in the system for which there are measures of both 

abundance and production, Kanaehoe Bay, the increase in production was, at two of the three 
stations, far greater than the increase in abundance. The Kanaeohe Bay results are from a 

relatively short-duration study after a sewage diversion and may suffer from averaging a 

relatively short period (Table 1). There was no clear trend in the magnitude of response in 

relation to the overall DIN concentrations in the various systems. However, the data do not 

clearly rule out a trend. 

It is not possible to determine how much of the range in factors found in Figure 13 and slopes 

in Figure 12 may result from inadequacies in the data and the use of different measures of 

phytoplankton abundance and production. It is discouraging to note that two studies of a 

similar portion of Chesapeake Bay (Upper Chesapeake Bay and Chesapeake Region VI in 

Table 1) which encompass different but overlapping time periods, provide very pictures of the 

response of phytoplankton abundance to increases in nitrogen concentrations. Yet, it seems 

improbable that the variability can be accounted for simply by scatter. It is unlikely a single 

magnitude of response to nitrogen increases applies to all the systems. The two systems with 

trends running counter to the general trend are particularly telling. There are also systems 

where, without any change in DIN, phytoplankton concentrations have increased. (Such 

examples cannot be plotted on Figure 13 as the effect value would be infinity.) For example, 

the Bokniseck station in the Keil Bight had increases in phytoplankton abundance and 

production over a period when there is no apparent increase in DIN (see Gerlach, 1990). 

THE ROLE OF ANOTHER FACTOR 

Perhaps it is simply unrealistic to expect that nitrogen availability alone will be a precise 

predictor of (long-term) phytoplankton abundance or production in coastal ecosystems. At the 

very least, phytoplankton abundance is also controlled by grazing. Phytoplankton production 

is also controlled by light availability that may in turn depend upon inorganic particles, 

unrelated to nitrogen, suspended in the water. Coastal ecosystems are also subject to 

anthropogenic inputs of other chemicals that may alter phytoplankton response to nitrogen. 

Revisiting the data assembled by Monbet (1992), shown earlier in Figure 7, demonstrates that 

an additional, somewhat subtle, factor helps control the abundance of phytoplankton in coastal 

ecosystems. Monbet (1992) divided the estuaries in hls data set into macrotidal (tidal range >2 

m) and microtidal (tidal range < 2 m). The macrotidal and microtidal points fall into 2 nearly 

distinct groups as shown in Figure 14. For the same concentration of nitrogen, macrotidal 

estuaries generally support lower phytoplankton abundances than microtidal estuaries. 

Presumably, the higher tidal range estuaries have greater mixing energies which in turn exhibits 

some type of control on phytoplankton abundance. Two mechanisms can be readily 

hypothesized. First, more rapid vertical mixing may keep individual phytoplankton cells out of 

the photic zone a sufficient fraction of the time to slow overall growth. Second, more vertical 

mixing may bring phytoplankton near to the benthos where they are subject to more rapid direct 

grazing by benthic organisms. Cloern (199 1 and personal communication) had previously 

noted that phytoplankton abundance in Northern San Francisco Bay was correlated to the neapspring cycle of the tides. 

Only one of the estuary segments shown in Figure 6 would be classified macrotidal (with a 

tidal range of only 2.1 m) by Monbet's definition. Nevertheless, if the data in Figure 6 are 

sorted into those segments with tidal ranges 21.2 m and 4.2 m, the set with the higher tidal 

ranges falls low relative to the set with the lower tidal range (Figure 15). Although these data 

do not provide a strong test of the effect of tidal height (lacking data from systems with high 

nitrogen concentrations and low tidal range), they are consistent with the trend found by 

Monbet (1992). 
CURVE FITS TO CROSS-SYSTEM AND MERL DATA AND COMPARISON TO THE 

EFFECT FACTORS FROM CHANGES OVER TIME 

Once the cross-estuary data is sorted into groups by tidal height, it is possible to calculate the 

magnitude of the change in abundance for a change in nitrogen for each group. This allows a 

comparison of the cross-estuary data to changes over time data. Figure 14 shows the Monbet 

(1992) data with power functions fit to the two groups of data. Of the types of equations for 

whch curve fits were tried (power, linear, log, and exponential) the power curve provided the 

best combination of correlation coefficients and normal distribution of residuals for both the 

Monbet (1992) data and the MERL data. Figure 15 shows the MERL abundance data 

(previously shown in Figure 9) fit with a power function. In Figure 16, a power function is fit 

to the data from segments of estuaries with tidal ranges less than 1.2 m. Changes in abundance 

expected from doublings in nitrogen concentration may be calculated directly from the fitted 

curves. Both groups of Monbet (1992) data and the MERL experiment indicate that a doubling 

of nitrogen concentrations will lead to approximately a 1.4 times increase in phytoplankton 

abundance. The data from the estuary segments with tidal heights 4.2 m indicates an increase 

of only 1.1 times for a doubling of nitrogen concentrations (although the correlation coefficient 

is rather low). Doubling values from the cross-estuary studies and the MERL experiment are 

within the range of those found in changes over time in individual estuaries (Figure 13). 

Cross-system comparisons, history of changes in coastal ecosystems, and experimentation all 

support the general idea that if an individual coastal ecosystem is enriched with nitrogen it will, 

probably have higher average abundances of phytoplankton and higher average phytoplankton 

primary production. 

Unfortunately, cross-system comparisons, based only upon nitrogen loading or 

concentrations, do not provide a very precise predictor of the phytoplankton abundance or 

production that will occur in individual systems. Other factors in addition to nitrogen need to 

be taken into account. Tidal height in coastal systems has been shown to have a large influence 

on the abundance of phytoplankton that will be supported by a given average level of DIN in 

the system. It appears that factors that may influence phytoplankton in addition to nitrogen and 

tidal height will need to be considered in order to be able to provide a more precise predictive 

relationship for phytoplankton abundance and production. 

The magnitude of the response for a doubling in nitrogen loading or average concentration is 

most likely to result in an increase in abundance or production by a factor between 1.4 and 3. 

The responses of most coastal ecosystems will probably fall within this range. 

However, there are systems that have not responded in the "normal" manner to nitrogen 

enrichment. Some coastal ecosystems have had changes in phytoplankton abundance without, 

or in the opposite direction of, changes in nitrogen concentration. Some coastal ecosystems do 

not follow along the trend of nitrogen vs. abundance or production curves, even when the 

systems are sorted by tidal height. 

There is, at present, no guarantee that in any individual system, reductions in nitrogen loadings 

will result in proportionate reductions (i.e., between 1.4 and 3 for a doubling in nitrogen) in 

phytoplankton abundance or production. There is no characteristic of estuaries identified here 

which will identify coastal ecosystems that will have unusual responses to nitrogen loadings. 

Table 1, continued. 

Phytoplankton Annual 

System Years Salinity % Average [DIN] Parameters Averages Sources 

pmolesniter and Measured [Nl or 

factor change in Loading 

[DIN] or loading 

Eastern Oosterschelde 1980-1989 27-30 52 hlorophyll and IDIN] Wetsteyn and Krom

(The Netherlands) 2.1 x Primary Nienhuis and Small, 

Production per 

area 

Helgoland Reed (North 1962- 1984 30-32 22.6 Cell volumes [DIN] Gerlach, 1990 

Sea) I .6x 

Halsskov Rev (Belt 1975- 1982 -20 7.2 Priamary [EN] (6) Nielsen and Ertebje

Sea, Baltic) 3.7~ Production per 

area 

Gulf of Finland (Baltic) 1972-1985 6 22.6 Chlorophyll [CN] (6) Kononen and Niemi

1.3~ Konenen, 1988 

Tolo Harbor 1978- 1985 25 4.4 Cell Counts [DIN] Hodgkiss and Chan, 

5.0~ 

Tolo Channel 32 1.9 Cell Counts [Dm] Hodgkiss and Chan, 

3.2~ 

Upper Pamlico 1972- 1992 0.5-5 13.6 Chlorophyll [Dm] Stanley, Personal Co

1.9~ 

1. Not detined, from discussion appers to be total DIN. 

2. Change was calculated as the average of 1960-1969 compared to the average of 1980- 1989 

3. Patuxent River to South RiverIAnnapolis. 

4. South RiverIAnnapolis to Bay Bridgehlagorhy River. 

5. Bay BridgeIMagothy River to susquehanna Flats. 

6. Not defined. 
Figure 1. From Nixon (1992). Labeled systems are: (1) Sargasso Sea; (2) North Pacific 

gyre; (3) Choptank River estuary; (4) the Patuxent estuary; (5) Pamlico River estuary; (6) 

Potomac River estuary; (7) main stem of the Chesapeake Bay; (8) Patapsco River estuary; 

(9) Continental Shelf off New York; (10) Georges Bank; (1 1) Peru Upwelling; (13) 

Delaware Bay; (14) Kaneohe Bay; (15) Baltic Sea. Open triangles are from the MERL 

eutrophication experiment. For data sources and further information see Nixon (1992). 

DIN INPUT, mmol m-3 y-' 
Figure 2. Primary Production by phytoplankton (14~ uptake) as a function of the annual 

input of dissolved inorganic nitrogen per unit area of a wide range of marine ecosystems. 

Systems are identified in Figure 1. The line represents a Redfield carbon to nitrogen 

molar ratio of 6.625 and gives a rough indication of how much carbon would be fixed at a 

given rate of nitrogen input without any recycling of nitrogen. 

DIN INPUT, moles m-2 y-' 
Figure 3. Mean annual concentrations of phytoplankton chlorophyll as a function of the 

annual input of dissolved inorganic nitrogen per unit volume of a wide range of marine 

ecosystems. Systems are identified in Figure 1. 

0.1 1 ' ' ' '1"1 I ,,,,I , , , I, ,,,I 

0. I I 10 lo2 10) 10' 

DIN INPUT, mmol m-3 y-I 
Figure 4. Mean annual concentrations of phytoplankton chlorophyll as a function of the 

annual input of dissolved inorganic nitrogen per unit area of a wide range of marine 

ecosystems. Systems are identified in Figure 1. 

0.1 1 1 11 1111 I I1 111111 I I l 11 Ill1 

0.01 0.1 1 10 100 

DIN INPUT, moles y-l 
Figure 5. Redrawn from Nixon (1983) and Nixon and Pilson (1983). 

Coastal ecosystems represented are: Long Island Sound, Kaneohe Bay, 

Chesapeake Bay, Narragansett Bay, Patuxent River, Apalachicola Bay, 

South San Francisco Bay, North San Francisco Bay, and 

Lower New York Bay. 

DIN input rate, mmol m-3 yfl 

DIN input rate. mmol yfl 
Figure 6. Unpublished data (compiled by K. Hinga, V. Berounsky, B. Kopp, A. Keller, 

A. Desbonnet, S. Pavignano, D. Stanley, M. Pilson, S. Nixon, and V. Lee). Each point 

represents a segment of an estuary. Estuaries were divided into typically four segments 

and data averaged for the segment. Averages may include up to 10 years of data. 

Estuaries represented are: Narragansett Bay, Delaware Inland Bays, Galveston Bay, 

Tampa Bay, San Francisco Bay, Buzzards Bay, Long Island Sound, Delaware Bay, 

Albemarle/Pamlico Bays, Barataria Bay, and Hudson River Estuary. 

1 10 100 1000 

Average DIN, Witer 

l,,,l,,,l,,,l, ., 

a 

- 

-a 

- 

a 

3. a 

- 

a a 

:a a 

a a 

- 

a a 

0. a :- a a - 

l"'I"'1"'l'" 

0 40 80 120 160 200 

Average DIN, pglliter 
Figure 7. Data from Monbet (1992). 

Annual Average DIN Concentration, prnoles 1-I 

Annual Average DIN Concentration, pmoles 1-I 
Figure 8. Vollenweider (1976) data (estimated) for 22 lakes with high phosphorus 

concentrations. . 

40 80 120 160 

Average Phosphorus Concentration, pgfliter 
22 

Figure 9. Data from MERL eutrophication Experiment (Frithsen et al., 1985). 

MERL Average DIN. pnoles~liter 

10 100 

MERL Average DIN, pnolefliter 
Figure 10. Data From MERL Eutrophication Experiment (Frithsen et al., 1985). 

Nitrogen addltlon rate 

mmol/m3/yr 
Figure 11. Data From MEW Eutrophication Experiment (Oviatt et al., 1986 and Keller, 

1988). 14~ production is from 14c uptake of water samples from the MERL enclosures. 

Net production is the result of whole-enclosure oxygen changes over a 24-hour period. 

14C PRODUCTKIN 

---f-- NET PRODUCTIVITY 

Net production, 28 month average 

14C production, 1 year of data 

100 I . Nitrogen addition rate 

mmollm3/yr 
Figure 12a. Chlorophyll concentrations at two different nitrogen 

concentrations for each system or station. See Table 1 for details 

on. each system. 

[DIN] or Total [N] 

----C)-- U. Chesapeake r Gulf of Finland 

Chesapeake IV . Helgoland 

---(>-- Chesapeake V Oosterschelde 

Chesapeake VI * Kaneohe, OF 

U. Parnlico Kaneohe,SE 

Tolo Harbor Kaneohe. CE 

---tE+-- Tolo channel 
Figure 12b. Legend same as Figure 12a. 

[DIN] or Total [N] 

---* U. Chesapeake -+ Gulf of Finland 

--* Chesapeake IV Helgoland 

----O-- Chesapeake V Oosterschelde 

* Chesapeake VI m Kaneohe. OF 

U. Parnlico Kaneohe, SE 

Tolo Harbor Kaneohe, CE 

Tolo channel 
Figure 13. For each system (See Table l), the data for the "effect" (chlorophyll, 

production, cell counts, or cell volume,) was regressed against the nitrogen concentration 

over the period of time when nitrogen and effects were changing. From the resultant 

linear relationship between effect and nitrogen, the predicted effect level was calculated 

for nitrogen values of 213 and 413 of the mean concentration for the system while the 

change was taking place. Thefactor for change in effect is the ratio of the calculated 

effects at 413 a d 213 of the mean nitrogen concentration. For all but four of the systems 

there was an increase in nitrogen over time. Two of the systems, Tampa Bay and 

Kanaehoe Bay, the change in nitrogen concentrations (and effect) were decreases 

resulting from sewage diversion. In the Oosterschlede nitrogen concentrations decreased 

as a result of a building of a storm-surge barrier. In the Parnlico river, the nitrogen 

concentrations were also going down over time. 

Average [Dm 

Upper Chesapeake + Kanacohe Bay, Chl 

Tampa Bay, Loadings Gulf of Finland 

Helogland Reede A Tolo Harbor 

v Tolo Channel X Kanaeoe Bay. Production 

+ Halskov Rev + Upper Pamlico 

0 Ches IV 0 Ches V 

Ches VI b Oosterschelde. Chl 

0 Oosterschelde. Rod 
Figure 14. Data from Monbet (1992) fitted with a power function. Data is divided into 

stations with tidal ranges >2 m (crosses) and 4 m (open circles) and each set fit 

separately. 

+ - >2m Chlorophyll Y = 0.626~O.~~' ? = 0.501 

0 - Rm Chlorophylly = 3.785~O.~~~ ? = 0.421 

Annual Average DIN Concentration, prnoles 1-I 

Annual Average DIN Concentration, prnoles 1-I 
Figure 15. Data shown in Figure 6 divided into two groups by tidal height in the 

segment. 

@ CHL Ave, pgtliter, Tide Range ~1.2 m 

o CHL Ave, paliter, Tide Range H.2 m 

o 40 80 120 160 200 

AVG DIN (uM) 

10 100 

AVG DIN (uM) 
Figure 16. Data from MERL eutrophication experiment fitted with a power function. 

10 100 

MERL Average DIN, pnoleslliter 
Figure 17. Data from Figures 6 and 14 with a power function fit to the data with tidal 

height 4.2 m. 

-0- CHL Ave, pgliter, Tide Range 4.2 rn 

o CHL Ave, pgtliter, Tide Range 21.2 m 

10 100 

AVG DIN (uM) 
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MARINE EUTROPHICATION REVIEW 

PART 2: BIBLIOGRAPHY WITH ABSTRACTS 
TABLE OF CONTENTS 

INTRODUCTION 1 

ASSEMBLY OF THE BIBLIOGRAPHY 1 

BIBLIOGRAPHY FORMAT 

Subject Lists 

Main Reference List 

ELECTRONIC VERSIONS OF THE BIBLIOGRAPHY 2 

RESOURCES USED IN THE BIBLIOGRAPHY (TABLES 1-4) 3 

SUBJECT LISTS 

Phytoplankton Abundance 

Phytoplankton Composition 

Red Tides And Nuisance Blooms 

Macrophytes 

Oxygen Concentration 

MAIN REFERENCE LIST 11 

APPENDIX: Keyword List 117 
INTRODUCTION 

Modem societies have increased the rate of supply of nutrients to estuaries and coastal marine environments 

from many sources over the rates in the pre-industrial era and times of lower population density. The addition 

of nutrients, or eutrophication, is considered a major problem in coastal areas. Typical symptoms attributed 

to eutrophication are: increases in algal biomass and productivity, nuisance algal blooms, algal mats and 

scums, increased bacterial activity, odor problems, turbidity, and depletion of oxygen in bottom waters. Long 

term and secondary effects may include changes in the composition of pelagic and benthic communities, 

reduction of species diversity, decreased abundance of submerged aquatic plants due to epiphetic or 

planktonic shading, disruption of food chains, and the possible occurrence of toxic phytoplankton blooms 

(such as "red tides"). On a positve note, a reasonable level of nutrient enrichment may increase primary 

productivity without much negative effect and may be converted into increased abundance and yield of 

desireable and harvestable species. 

It is likely that all the consequences of estuarine eutrophication are not known. Even for those known effects, 

a clear understanding of the magnitude of many of the effects resulting from increases in nutrient loadings is 

lacking. Equally important, there are likely some effects attributed to nutrient additions to coastal 

environments which are not actually caused by the increases in nutrients. This may occur because other 

materials may be added to the marine environment, or there are other alterations of the environment which are 

occurring simultaneously with nutrient additions. 

In order to make a step toward a better understanding of the effects of eutrophication, the NOAA Coastal 

Ocean Program commissioned the assembly of this bibliography on marine eutrophication. Literature which 

may be useful in understanding marine eutrophication is in diverse sources. This bibliography is intended to 

facilitate the finding of literature either for the scientist or manager who may want to develop a clearer picture 

of marine eutrophication. The bibliography is not a compilation of data sources which might be used for new 

analysis of eutrophication. 

ASSEMBLY OF THE BIBLIOGRAPHY 

The 438 references in this bibliography were assembled by searching a number of data bases, utilization of 

symposia volumes concerning eutrophication, special journal volumes on .eutrophication, and books on 

nutrients in the marine environment. The data searched, and the years covered by the sources are listed in 

Table 1. The keywords used for those searches are listed in Table 2. 

The symposia volumes and books which were used for sources are listed in Table 3. Special journal volumes 

are listed in Table 4. The judgement for including papers from these sources was based upon the content of 

the individual papers. Our intent was to include papers which had research results or other insights 

concerning marine eutrophication and not to include papers which simply mentioned the topic. Nor did we 

attempt to compile a listing of data sources. There is not always a clear distinction between these two 

categories and other preparers might have sorted papers somewhat differently. 

Finally, papers which were listed in the references of the papers located by the above two techniques were 

included where they met the criteria for inclusion. 
BIBLIOGRAPHY FORMAT 

Subject Indexes 

There are five lists identifying papers contaning information in five major subject divisions. These are: 

phytoplankton abundance 

macrophytes 

red tides and other nuisance blooms 

phytoplankton community assemblage 

oxygen concentrations 

Each reference in these lists gives author and year. The complete reference is found in the main reference list 

described below. 

Main Reference List 

The references are listed alphabetically by author. The year of publication appears next, followed by the title 

of the work in bold text. The complete citation to the work appears next. Keywords highlighting the content 

of the referenced work appear in brackets after the citation. Keywords supplied with the original work were 

used when available. Keywords were provided for other references. Appendix A is a compiled list of the 

keywords found in the bibliography. The complete abstract of the paper follows the citation unless there was 

no abstract. 

ELECTRONIC VERSIONS OF THE BIBLIOGRAPHY. 

The primary version of the bibliography was assembled in Endnote PlusTM software for the MacintoshTM. 

The file is about 985 kilobites and is supplied upon request with the hard copy. ASCII versions of the main 

reference list for both MacintoshTM and PCTM compatible computers can be supplied upon request (about 500 

kilobites). 
RESOURCES USED IN THE BIBLIOGRAPHY 

Table 1. Data Bases Utilized For Preparation Of Bibliography 

Electronic Data Bases Years Covered 

Aquatic Sciences and Fisheries Abstracts 1978-1992 

science Citation Index 1989-1991 

NOAA Libraries Catalog 

Government Documents Catalog Service 1976-1991 

Paper Copy Abstracts 

Aquatic Sciences and Fisheries Abstracts 1970- 1977 

Deep Sea Research 1970-1991 

Dissertation Abstracts 1970-1990 

Pollution Abstracts 1970-1991 

Table 2. Words Used In Searches Of Electronic Databases 

Abnormality 

Abundance 

Algae 

Anoxia 

Assemblage 

Biomass 

Bivalves 

Bloom 

Brown Tides 

Chlorophyll 

Clams 

Coast 

Community 

Coral 

Cycles 

Deficiency 

Denitrification 

Depletion 

Diatom 

Dinoflagellates 

Discharge 

Diversity 

Dynamics 

Eelgrass 

Effluents 

Energy 

Enriched 

Enrichment 

Epifauna 

Epiphytes 

Estuaries 

Eutrophication 

Eutrophied 

Fauna 

Fish 

Function 

Growth 

Hypoxia 

Infauna 

Light 

Macroalgae 

Macrofauna 

Macrophyte 

Meiofauna 

Metabolism 

Microalgae 

Nitrate 

Nitrification 

Nitrite 

Nitrogen 

Number 

Nutrients 

Organic 

Organic Matter 

Pathology 

pH Phosphate 

Phosphorus 

Photosynthesis 

Pollution 

Population 

Primary Production 

Primary Productivity 

Red Tides 

Redox 

Response 

Seagrass 

Sewage 

Species Composition 

Standing Crop 

Structure 

Sublethal 

Submerged Vascular Plants 

Sulfate 

Sulfur 

Turbidity 

Waste 

Water Quality 

Zooplankton 
Table 3. Symposia Volumes And Edited Books With Papers On Eutrophication 

Which Were Used In The Assembly Of The Bibliography 

Anderson, D. M., A. W. White, and D. G. Baden 

(1985) Toxic Dinoflagellates. (Proceedings 

of The Third International Conference on 

Toxic Dinoflagellates, St. Andrews, New 

Brunswick, Canada 8-12 June, 1985). New 

York, Elsevier. 

Carpenter, E. J. and D. G. Capone (1983) 

Nitrogen in the Marine Environment. New 

York, Academic Press. 

Cosper, E. M., V. M. Bricelj, and E. J. Carpenter 

(1989) Novel Phytoplankton Blooms: 

Causes and Impacts of Recurrent Brown 

Tides and other Unusual Blooms. (Papers 

from a conference at Stony Brook, NY, 27-28 

October, 1988). Berlin, Springer-Verlag. 

Dennis, L. and H. H. Seliger (1979) Toxic 

Dinoflagellate Blooms. (Proceedings of the 

Second International Conference on Toxic 

Dinoflagellate Blooms, Key Biscayne, 

Florida, 31 October - 5 November, 1978). 

North Holland, Elsevier. 

GranCli, E., B. Sundstrom, L. Edler, and D. M. 

Anderson (1990) Toxic Marine 

Phytoplankton. (Proceedings of the Fourth 

International Conference on Toxic Marine 

Phytoplankton, Lund, Sweden, 26-30 June, 

1989). New York, Elsevier. 

Hallegraeff, M. and J. L. Maclean (1989) Biology 

Epidemology and Management of 

Pyrodinium Red Tides. (Proceedings of the 

management and training workshop, Bandar 

Seri Begawan, Brunei Darussalam, 

Philippines, 23-30 May, 1989). Manila, 

Fisheries Department, Ministry of 

Development. 

Kennedy, V. S. (1984) The Estuary as a Filter. 

Orlando, Academic Press. 

Likens, G. E. (1972) Nutrients and 

Eutrophication: The Limiting Nutrient 

Controversy. (Proceedings of the Symposium 

on Nutrients and Eutrophication: The 

Limiting Nutrient Controversy, W.K. Kellog 

Biological Station, Michigan State 

University, 11-12 February, 1971). 

Lawrence, Kansas, American Society of 

Limnology and Oceanography and the Allen 

Press. 

LoCicero, V. R. (1975) Toxic Dinoflagellate 

Blooms. (Proceedings of the First 

International Conference on Toxic 

Dinoflagellate Blooms, Boston, 4-6 

November, 1974) (MIT Sea Grant Report 

MITSG 75-8). Wakefield, Massachusetts, 

Massachusetts Science & Technology 

Foundation. 

Neilson, B:J. and L. E. Cronin (1981) Estuaries 

and Nutrients. Clifton, New Jersey, Humana 

Press. 

Okaichi, T., D. M. Anderson, and T. Nemoto 

(1988) Red Tides: Biology, Environmental 

Science, and Toxicology. (Proceedings of the 

International Symposium on Red Tides, 

Takamastu Perfecture, Japan, 10- 14 

November, 1987). New York, Elsevier. 

Smayda, T. J. and Y. Shimizu (1993) Toxic 

Phytoplankton Blooms in the Sea. 

(Proceedings of the Fifth International 

Conference on Toxic Marine Phytoplankton, 

Newport, Rhode Island, USA., 28 October 

- 1 November 199 1). Amsterdam, Elsevier. 

Tyson, R. V. and T. H. Pearson (1991) Modern 

and Ancient ContinentaIShelfAnoxia. 

Special Publication No. 58 of The Geological 

Society. London, The Geolocical Society. 

Vollenweider, R.A., R. Marchetti, and R. Viviani 

(1992) Marine Coastal Eutrophication. 

(Proceedings of an International conference, 

bologna, Italy, 21-24 march 1990. Reprinted 

from the journal Science of the Total 

Environment, Supplement, 1992) Amsterdam, 

Elsevier. 
Table 4. Special journal volumes concerning eutrophication used in the assembly 

of the bibliography 

Ambio Volume 19, Issue 3 (1990) Special Issue: 

Marine Eutrophication. 

Kieler Meeresforschungen, Sonderheft 6 (1988) 

The Baltic Sea Environment, HistoryEutrophication-Recruitment-Ecotoxicology. 

(Proceedings of the 10th Symposium of the Baltic 

Marine Biologists, 29 September - 3 October 

1987). 

Marine Pollution Bulletin Volume 20, Issue 7 

(1989) Pollution in the Far East. 

Marine Pollution Bulletin Volume 23 (199 1) 

Environmental Management and Appropriate 

Use of Enclosed Coastal Seas. (Proceedings of 

the International Conference on the 

Environmental Management of ~ndosed Coastal 

Seas '90, held in Kobe, Hyogo Prefecture, Japan, 

3-6 August, 1990). 

Netherlands Journal of Sea Research Volume 19, 

Issue 3&4 (1984). Nutrients in Estuaries: 

Eutrophication and Other Aspects. 

(proceedings of the ESBA/Hydrobiological 

Society Symposium, Texel, The Netherlands, 11- 

13 September, 1984). 
SUBJECT INDEXES 

SUBJECT INDEX: Phytoplankton Abundance 

Words Searched for: abundance; bloom; chlorophyll; chl a; algal growth 

(142 references) 

Andersin, A.-B., J. Lassig, L. Parkkonen, and H. 

Sandler (1978) 

Anderson, D. M. (1987) 

Anderson, D. M. (1992) 

Arzul, G. and P. Gentien (1990) 

Austen, M. C., R. M. Warwick, and M. C. Rosado 

(1989) 

Beatty, L. L. (1991) 

Beukema, J. J. (1990) 

Beukema, J. J. (1991) 

Beukema, J. J. and G. C. CadBe (1986) 

Beukema, J. J. and G. C. Cadte (1991) 

Birch, P. B. and J. 0. Gabrielson (1984) 

Bodeanu, N. (1992) 

Boni, L., E. Carpent, D. Wynne, and M. Reti 

(1989) 

Brand, L. E., M. D. Gottfried, C. C. Baylon, and 

N. S. Romer (1991) 

Breuer, G. and W. Schramm (1988) 

Brockmann, U., G. Billen, and W. W. C. Gieskes 

(1988) 

Brockmann, U. and E. Dahl(1990) 

Brockmann, U. H., E. Dahl, and K. Eberlein 

(1985) 

Brown, J. R., R. J. Gowen, and D. S. McLusky 

(1987) 

CadBe, G. C. (1986) 

CadBe, G. C. (1990) 

CadBe, G. C. (1992) 

CadBe, G. C. and J. Hegeman (1991) 

Cederwall, H. and R. Elmgren (1980) 

Cosper, E. M., W. Dennison, A. Milligan, E. J. 

Carpenter, C. Lee, J. Holzapfel, and L. 

Milanese (1989) 

Cosper, E. M., C. Lee, and E. J. Carpenter (1990) 

Costa, J. E. (1988) 

Dauer, D. M. and W. G. Conner (1980) 

Dauvin, J.-C. and F. Gentil(1989) 

Dzurica, S., C. Lee, E. M. Cosper, and E. J. 

Carpenter (1989) 

Emara, H.I., M.A. Shriadah, Th.H. Moustafa and 

M.S. El-Deek (1992) 

Fanuko, N. (1984) 

Friligos, N. (1985) 

Gerlach, S. A. (1990) 

Giovanaridi, F. and E. Tromellini (1992) 

Gowen, R. J., P. Tett, and K. J. Jones (1992) 

GranBli, E., P. Carlsson, P. Olsson, B. Sundstrom, 

W. GranBli, and 0. Lindahl(1989) 

GranBli, E., W. Graneli, and L. Rydberg (1986) 

GranCli, E., S. Schulz, U. Schiewer, D. 

Gedziorwska, W. Kaiser, and M. Plinski 

(1988) 

Grassle, J. P. and J. F. Grassle (1984) 

Gray, J. S. (1979) 

Haas, L. W., S. J. Hastings, and K. L. Webb 

(1981) 

Hagmeier, E. (1978) 

Hardy, J. T. and Z. Zubayli (1976) 

Heinanen, A. P. (1991) 

Hinga, K. R. (1992) 

Hobbie, J. E. and J. J. Cole (1984) 

Hodgkiss, I. J. and B. S. S. Chan (1987) 

Howells, G. P. (1972) 

Hull, S. C. (1987) 

Hungspreugs, M., W. Utoomprkporn, S. 

Dharmvanij, and P. Sompongchaiyakul 

(1989) 

Imai, I., S. Itakura, and K. Itoh (1991) 

Jochem, F. and B. Babenerd (1989) 

Johansson, J.O.R. and R.R. Lewis, I11 (1992) 

Josefson, A. B. (1987) 

Josefson, A. B. and R. Rosenberg (1988) 

Kaiser, W., A. Irmisch, D. Nehring, F. Georgi, 

and G. Bruel (1990) 

Kat, M. (1987) 

Kautsky, N., H. Kautsky, U. Kautsky, and M. 

Waern (1986) 

Keller, A. A. (1989) 

Keller, A. A. and R. L. Rice (1989) 

Keller, A. A. and R. L. Rice (1990) 

Kononen, K. and A. Niemi (1984) 

Kroncke, I. (1990) 

Laws, E. A. and D. G. Redalje (1979) 

Leppakoski, E. (1980) 

Maclean, J. L. (1984) 

Maclean, J. L. (1989) 

Maestrini, S. Y. and E. GranBli (1991) 

Mattson, J. and 0. Linden (1983) 

McLusky, D. S., M. Teare, and P. Phizacklea 

(1980) 

Millner, R. S. (1980) 

Mingazzini, M., A. Rinaldi, and g. Montanari 

(1992) 

Morton, B. (1989) 

Moshiri, G. A., N. G. Aumen, and W. G. 

Crumpton (198 1) 

Murakarni, A. (1973) 

Nielsen, A. and G. Ertebjerg (1984) 

Nishijima, T. and Y. Hata (1991) 
Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P. 

Donaghay, B. Sullivan, S. Seitzinger, D. 

Rudnick, and J. Frithsen (1984) 

Okaichi, T. (1987) 

Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst 

(1984) 

Olsson, G. E., B. Sundstrom, and L. Edler (1987) 

Oviatt, C. A., P. Lane, F. F. 111, and P. Donaghay 

(1988) 

Paerl, H. W. (1988) 

Pastorok, R. A. and G. R. Bilyard (1985) 

Pearson, T. H. (1980) 

Pearson, T. H. (1987) 

Pearson, T. H., A. B. Josefson, and R. Rosenberg 

(1985) 

Pearson, T. H. and R. Rosenberg (1976) 

Pearson, T. H. and R. Rosenberg (1978) 

Pearson, T. H. and S. 0. Stanley (1979) 

Pekkari, S. (1973) 

Perciasepe, R. (1992) 

Persson, L.-E. (1987) 

Phillips, D. J. H. and S. Tanabe (1989) 

Pommb, F. (1992) 

Price, K. S., D. A. Flemer, J. L. Taft, G. B. 

Mackiernan, W. Nehlsen, R. B. Biggs, N. 

H. Burger, and D. A. Blaylock (1985) 

Raman, A. V. and K. P. Prakash (1989) 

Renk, H., J. Nakonieczny, and S. Ochocki (1988) 

Revelante, N. and M. Gilmartin (1976) 

Ritz, D. A., M. E. Lewis, and M. Shen (1989) 

Rosenberg, R. (1985) 

Rosenberg, R., J. S. Gray, A. B. Josefson, and T. 

H. Pearson (1987) 

Schiewer, U., R. Borner, and N. Wasmund (1988) 

Seliger, H. H. (1989) 

Sellner, K. G. and M. M. Olson (1985) 

7 

Silva, E. S. (1985) 

Skjoldal, H. R. and I. Dundas (1991) 

Smayda, T. J. (1989) 

Smayda, T. J. and A. W. White (1990) 

Soulsby, P. G., D. Lowthion, and M. Houston 

(1982) 

Spencer, C. P. (1985) 

Stockner, J. G., D. D. Cliff, and K. R. S. 

Shortreed (1979) 

Stockner, J. G. and K. S. Shortreed (1988) 

Sukhanova, I. N., M. V. Flint, G. Hilbaum, V. 

Karamfilov, A. I. Kopylov, E. Matveea, 

T. N. Rat'kova, and A. F. Sazhin (1988) 

Takahashi, M. and N. Fukazawa (1982) 

Takano, H. (1987) 

Tamminen, T. (1990) 

Taslakian, M. J. and J. T. Hardy (1976) 

Thompson, G. B. and J. Ho (1981) 

Tsutsumi, H., T. Kikuchi, M. Tanaka, K. Imasaka, 

and M. Miyazaki (1991) 

Vidakovi'c, J. (1983) 

Vukadin, H. (1991) 

Walker, D. I. and R. F. G. Ormond (1982) 

Wallstrom, K. (1988) 

Wang, Z. (1987) 

Weigelt, M. and H. Rumohr (1986) 

Weston, D. P. (1990) 

Widbom, B. and R. Elmgren (1988) 

Wittberg, M. and W. Hunte (1992) 

Wong, P. S. (1987) 

Wu, R. S. S. (1982) 

Wyatt, T. and J. Horwood (1973) 

Yang, D. B. (1987) 

Yi, S. K., J.4. Hong, and J. H. Lee (1982) 

Zingone, A., M. Monitresor, and D. Marino 

(1990) 
SUBJECT INDEX: Phytoplankton Composition 

Words searched for: assemblage; community structure; species composition; species richness 

(25 references) 

Abdul-Hussein, M. M. and C. F. Mason (1988) 

Anderson, N. J., B. Rippey, and A. C. Stevenson 

(1990) 

Beauchamp, S. T. and J. Kerekes (1989) 

B odeanu, N. (1 992) 

Crema, R., A. Castelli, and D. Prevedelli (1991) 

DeGroodt, E. G., H. F. J. Los, T. A. Nauta, A. A. 

Markus, and I. deVries (1992) 

Dortch, Q., D. Milstead, N.N. Rabalais, S.e. 

Lorenz, D.G. Redalje, M.J. Dagg, M.E. 

Turner, and T.E. Whitledge (1992) 

Gerlach, S. A. (1990) 

Grantli, E., H. Persson, and L. Edler (1986) 

Josefson, A. B. (1987) 

Kononen, K. and A. Niemi (1984) 

Kroncke, I. (1990) 

Manen, A., I. Haahtela, H. Ilvessalo, and J. 

Lehto (1984) 

Moore, C. G. and T. H. Pearson (1984) 

Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P. 

Donaghay, B. Sullivan, S. Seitzinger, D. 

Rudnick, and J. Frithsen (1984) 

Oviatt, C. A., P. Lane, F. F. 111, and P. Donaghay 

(1988) 

Pastorok, R. A. and G. R. Bilyard (1985) 

Schultz, S., G. Ertebjerg, G. Behrends, G. Breuel, 

P. Ciszewski, U. Horstmann, K. 

Kononen, E. Kostrichkina, J. Leppanen, 

F. Mghlenberg, 0. Sandstrom, M. 

Viitasalo, and T. Willtn (1992) 

Skjoldal, H. (1992) 

Sukhanova, I. N., M. V. Flint, G. Hilbaum, V. 

Karamfilov, A. I. Kopylov, E. Matveea, 

T. N. Rat'kova, and A. F. Sazhin (1988) 

Swartz, R. C., F. A. Cole, D. W. Schults, and W. 

A. DeBen (1986) 

Thompson, G. B. and J. Ho (1981) 

Tomascik, T. and F. Sander (1987) 

Valente, R. M., D. C. Rhoads, J. D. Germano, and 

V. J. Cabelli (1992) 

Wittberg, M. and W. Hunte (1992) 
SUBJECT INDEX : Red Tides and Nuisance Blooms 

Words searched for: red tide; brown tide; nuisance blooms 

(38 references) 

Anderson, D. M. (1992) 

Arzul, G. and P. Gentien (1990) 

Aubert, M. (1992) 

Cho, C.-H. (1991) 

Cosper, E. M., W. Dennison, A. Milligan, E. J. 

Carpenter, C. Lee, J. Holzapfel, and L. 

Milanese (1989) 

Cosper, E. M., C. Lee, and E. J. Carpenter (1990) 

Doucette, G. J. (1989) 

Dzurica, S., C. Lee, E. M. Cosper, and E. J. 

Carpenter (1989) 

Fiedler, P. C. (1982) 

Hodgkiss, I. J. and B. S. S. Chan (1987) 

Hungspreugs, M., W. Utoomprkporn, S. 

Dharmvanij, and P. Sompongchaiyakul 

(1989) 

Imai, I., S. Itakura, and K. Itoh (1991) 

Jingzhong, Z., D. Liping, and Q. Baoping (1985) 

Keller, A. A. and R. L. Rice (1989) 

Keller, A. A. and R. L. Rice (1990) 

Lam, C. W. Y. and K. C. Ho (1987) 

Liaci, L. S. (1980) 

Maclean, J. L. (1984) 

Maclean, J. L. (1989) 

Morton, B. (1989) 

Murakami, A. (1973) 

Nishijima, T. and Y. Hata (1991) 

Nixon, S. W. (1989) 

Ochi, T. (1987) 

Okaichi, T. (1987) 

Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst 

(1984) 

Phillips, D. J. H. and S. Tanabe (1989) 

Reguera, B. and Y. Oshima (1990) 

Seliger, H. H. (1989) 

Sellner, K. G. and M. M. Olson (1985) 

Silva, E. S. (1985) 

Smayda, T. J. (1992) 

Sukhanova, I. N., M. V. Flint, G. Hilbaum, V. 

Karamfilov, A. I. Kopylov, E. Matveea, 

T. N. Rat'kova, and A. F. Sazhin (1988) 

Takahashi, M. and N. Fukazawa (1982) 

Wang, Z. (1987) 

Wong, P. S. (1987) 

Wyatt, T. and J. Horwood (1973) 

Yang, D. B. (1987) 

SUBJECT INDEX: Macrophytes 

Words searched for: seaweeds; macroalgae. 

(20 references) 

Baden, S. P., L.-0. Loo, L. Pihl, and R. 

Rosenberg (1990) 

Birch, P. B. and J. 0. Gabrielson (1984) 

Cheney, D. P. (1992) 

Conolly, N. J. and E. A. Drew (1985) 

Fletcher, R. L., V. Cuomo, and I. Palomba (1990) 

Harlin, M. M. and B. Throne-Miller (1981) 

Hull, S. C. (1987) 

Kautsky, N., H. Kautsky, U. Kautsky, and M. 

Waern (1986) 

Lee, V. and S. Olsen (1985) 

Lenzi, M. (1992) 

Mdcinen, A., I. Haahtela, H. Ilvessalo, and J. 

Lehto (1984) 

Menesguen, A. (1992) 

Pekkari, S. (1973) 

Quinlan, A. V. (1982) 

Raffaelli, D., J. Limia, S. Hull, and S. Pont (1991) 

Schramm, W., D. Abele, and G. Breuer (1988) 

Smith, D. W. (1984) 

Soulsby, P. G., D. Lowthion, and H. A. C. 

Montgomery (1985) 

Tubbs, C. R. and J. M. Tubbs (1983) 

Vogt, H. and W. Schrarnm (1991) 
SUBJECT INDEX: Oxygen Concentration 

Words searched for: anoxia; hypoxia 

(65 references) 

Andersin, A.-B., J. Lassig, L. Parkkonen, and H. 

Sandler (1978) 

Andersson, L. and L. Rydberg (1988) 

Aneer, G. (1987) 

Baden, S. P., L.-0. Loo, L. Pihl, and R. 

Rosenberg (1990) 

Boesch, D. F. (1983) 

Brockmann, U., G. Billen, and W. W. C. Gieskes 

(1988) 

Degobbis, D. (1989) 

Dethlefsen, V. and H. v. Westernhagen (1983) 

Dortch, Q., D. Milstead, N.N. Rabalais, S.e. 

Lorenz, D.G. Redalje, M.J. Dagg, M.E. 

Turner, and T.E. Whitledge (1992) 

Emara, H.I., M.A. Shridah, Th.H. Moustafa, and 

M.S. El-Deek (1992) 

Friligos, N. (1981) 

Gerlach, S. A. (1990) 

Griggs, G. B. and T. S. Hopkins (1976) 

Herdendorf, C. E. (1986) 

Hognestad, P. T. (1987) 

Hosie, D. A., R. F. Uglow, L. Hagennan, T. 

Sondergaard, and W. Weile (1991) 

Justi'c, D. (1987) 

Justi'c, D. (1991) 

Kohler, A. and F. Holzel(1980) 

Lam, C. W. Y. and K. C. Ho (1987) 

Leppakoski, E. (1980) 

LISS, L. I. S. S. (1990) 

Malone, T. C. (1991) 

Millner, R. S. (1980) 

Mills, C. A., S. I. Heaney, C. Butterwick, J. E. 

Corry, and J. M. Elliott (1990) 

Morton, B. (1989) 

Nehring, D., S. Schulz, and W. Kaiser (1984) 

Nehring, D. (1992) 

Nilsson, P., B. Jonsson, I. L. Swanberg, and K. 

Sundback (1991) 

Nixon, S. W. and V. M. Berounsky (1984) 

Nixon, S. W., C. A. Oviatt, J. Garber, and V. Lee 

(1976) 

Ochi, T. (1987) 

Okaichi, T. (1987) 

Oviatt, C. A., A. A. Keller, P. A. Sampou, and L. 

L. Beatty (1986) 

Oviatt, C. A., J. G. Quinn, J. T. Maughan, J. T. 

Ellis, B. K. Sullivan, J. N. Gearing, P. J. 

Gearing, C. D. Hunt, P. A. Sampou, and 

J. S. Latimer (1987) 

Parker, C. A. (1983) 

Parker, C. A. and J. E. O'Reilly (1991) 

Pastorok, R. A. and G. R. Bilyard (1985) 

Pearson, T. H. (1980) 

Perciasepe, R. (1 992) 

Phillips, D. J. H. and S. Tanabe (1989) 

Phoel, W. C., K. L. Webb, and C. F. ~'~lia'(1981) 

Polak, J. and G. D. Haffner (1978) 

Portnoy, J. W. (1991) 

Price, K. S., D. A. Flemer, J. L. Taft, G. B. 

Mackiernan, W. Nehlsen, R. B. Biggs, N. 

H. Burger, and D. A. Blaylock (1985) 

Rabalais, N. N. and D. F. Boesch (1986-1987) 

Rabalais, N. N., R. E. Turner, J. W.J. Wiseman, 

and D. F. Boesch (1991) 

Reys, E. and M. Merino (1991) 

Riisgtd, H. U. and E. Poulsen (1981) 

Rosenberg, R. (1985) 

Rosenberg, R., B. Hellman, and B. Johansson 

(1991) 

Rosenberg, R. and L.-0. Loo (1988) 

Rydberg, L., L. Edler, S. Floderus, and W. 

GranCli (1990) 

Sales, D., A. Gbmez, and D. Cantero (1983) 

Sampou, P. and C. A. Oviatt (1991) 

Sampou, P. A. (1989) 

Stachowitsch, M. (1991) 

Tolmazin, D. (1985) 

Topping, G. (1976) 

Van Es, F. B., V. A. M.A., L. A. Bouwman, and 

H. G. J. Schroder (1980) 

Vollenweider, R.A., A. Rinaldi, and G. Montanari 

(1992) 

Weigelt, M. and H. Rumohr (1986) 

Welsh, B. L. and F. C. Eller (1991) 

Whiteledge, T. E. (1985) 

Wu, R. S. S. (1982) 
11 

MAIN REFERENCE LIST 

Abdul-Hussein, M. M. and C. F. Mason (1988) 

The phytoplankton community of a 

eutrophic reservoir. Hydrobiologia 169:265- 

277. 

{phytoplankton; reservior; community; pH; 

composition; assemblage) 

The dynamics of the phytoplankton community 

of a eutrophic reservoir are described for a two year 

period. Fifty-eight species were recorded, 25 of 

them common. Bacillariophyta dominated during 

the winter and early spring and Chlorophyta during 

late spring, to be replaced by a bloom of 

Cyanophyta. The mean and peak biomass of 

phytoplankton was 8.6 mg 1-I and 40.8 mg 1-I in 

1981, and 8.3 mg 1-I and 37.6 mg 1-I in 1982. 

Temperature accounted for 67.3% and pH for 8% of 

the variation in total phytoplankton biomass over 

the two year period, using regression technique. 

Both horizontal and vertical patchiness, measured 

as an index of mean crowding, were recorded in the 

reservoir. Horizontal aggregations were associated 

with spring blooms of Chlorophyta and summer 

blooms of Cyanophyta, while vertical aggregations 

were most marked during the summer bloom of 

Cyanophyta. Concentrations of phytoplankton were 

influenced by wind, the prevailing southwesterly 

wind accumulating algae in the northeasterly arm of 

the reservoir during much of the year. 

Andersin, A.-B., J. Lassig, L. Parkkonen, and H. 

Sandler (1978) The decline of macrofauna in 

the deeper parts of Baltic proper and the 

Gulf of Finland. Kieler Meeresforschungen 

Sonderheft 4:23-30. 

{abundance; benthic; macrofauna; oxygen; hypoxia; 

Baltic; hydrogen sulfide) 

An attempt is made to describe the large-scale 

changes in the benthic soft bottom macrofauna in 

the deep parts of the Bornholm Basin, the Gulf of 

Gda'nsk, the Central Basin and the Gulf of Finland, 

from the beginning of Baltic zoobenthos research to 

the present day. The authors also try to correlate 

these changes with fluctuations in the oxygen 

content and salinity in near-bottom water layer. The 

paper surveys the literature and presents recent and 

earlier unpublished results. 

During the later part of last century and the first 

decades of the twentieth century no area of the 

Baltic Sea seemed to have been totally devoid of 

macrofauna. Unfortunately there are considerable 

gaps in our knowledge of the time before the 

middle of this century. The most striking decline 

has taken place, generally speaking, after the 

exceptionally great inflow in 1951-1952, and the 

subsequent prolonged stagnation. 

The first records of "dead" bottoms in the 

Bornholm basin are from 1948, when no 

macrofauna was recorded below 80 m. Records from 

1954 show that the deepest parts of the Eastern 

Gotland Basin and the deep area between 0land and 

Gotland were devoid of macrofauna at that time, but 

that the deep areas of the northernmost Baltic 

proper and the Gulf of Finland were still populated. 

The change continued and during the 1960s the 

communities dominated by lamellibrachs in the 

Bornholm and Gda'nsk Deeps disappeared, and were 

subsequently replaced by polychaete communities. 

These have been wiped out during periods of bad 

oxygen conditions, but quickly re-established when 

conditions improved. The lammelibrach 

community has not been restored. In the Northern 

Central Basin and the Gulf of Finland the 

depopulation of the deep bottoms probably began 

later, in the late 50's. In the 70s practically no 

macrofauna has been recorded below the permanent 

halocline in the Central Basin (except the 

southernmost parts of it) and the Gulf of Finland. 

During the 60s and 70s the area with periodically 

unfavorable oxygen conditions has covered about 

100,000 km2, which is 25% of the total area of the 

Baltic Sea. 

Anderson, D. M. (1987) Toxic algal blooms 

and red tides: A global perspective. In: 

T. Okaichi, D. M. Anderson and T. Nemoto (ed.) 

Red Tides: Biology, Environmental Science, and 

Toxicology. Elsevier, New York. 

{toxic; bloom) 

The literature on toxic algal blooms and red tides 

documents a global increase in the frequency, 

magnitude, and geographic extent of these events 

over the last two decades. Some of this increase is 

undoubtedly a result of the increased awareness and 

analytical capabilities of the scientific community, 

but a strong correlation between the number of red 

tides and the degree of coastal pollution or 

utilization of coastal waters for aquaculture argue 

that there are other contributing factors. It also 

appears likely that toxic algal species have spread 

within regions over spatial scales of hundreds of 

kilometers, moving with major water currents and 

storms. Long distance transport of species across 

oceans~may have occurred as well, but the evidence 

is not conclusive and the hypothesis controversial. 

Anderson, D. M. (1992) Toxic dinoflagellate 

blooms and red tides in New England and 

abroad: A biogeographic and physiologic 

perspective. 31st Annual Symposium of the 

Northeast Algal Society, Woods Hole. 

{dinoflagellate; red tide; bloom; Alexandrium; Gulf 

of Maine) 
Toxic red tides of the dinoflagellate Alexandrium 

fundyense have become persistent phenomena over 

the last two decades in the southern Gulf of Maine, 

having been common occurrences for hundreds of 

years further to the north. This paper will explore 

the bloom dynamics of this organism in the Gulf, 

as well as the physiological characteristics of the 

cells that influence their toxin production. 

Additional studies will be presented that relate to 

the population biology of Alexandrium, both 

within the Gulf of Maine, and throughout the 

world. Through a combination of traditional field 

phytoplankton ecology, physical oceanographic 

measurements, remote sensing, toxin analysis, 

morphological analysis, and molecular studies, the 

complex and fascinating characteristics of this 

important organism are gradually becoming known. 

Topics to be emphasized include the long-distance 

transport and confinement of A lexandrium 

populations within a coastal current that originates 

in the rivers of southern Maine; the potential 

impact of environmental conditions on the 

production of toxins in Alexandrium; the role of 

dormant cysts in the bloom dynamics and 

population distribution of Alexandrium; the 

possible global dispersal of A lexandri um 

populations via ship traffic; and forensic RFLP 

analysis of globally distributed Alexandrium 

populations. 

Anderson, D. M., A. W. White, and D. G. Baden 

(ed.) (1985) Toxic Dinoflagellates 

(Proceedings of The Third International 

Conference on Toxic Dinoflagellates, St. 

Andrews, New Brunswick, Canada 8-12 

June, 1985). Elsevier, New York. 

No abstract 

Anderson, N. J., B. Rippey, and A. C. Stevenson 

(1990) Change to a diatom assemblage in 

a eutrophic lake following point source 

nutrient re-direction: a 

palaeolimmnological approach. Freshwater 

Biology 23:205-2 17. 

{diatom; assemblage; lake; nutrient) 

No abstract. 

Andersson, L. and L. Rydberg (1988) Trends in 

nutrient and oxygen conditions within 

Kattegat: Effects of local nutrient 

supply. Estuarine, Coastal and Shelf Science 

26:559-579. 

{nitrogen; oxygen; phosphorus; nutrient; Baltic) 

The Kattegat forms the outer part of the Baltic 

estuary. It is characterized by a stable two-layer 

stratification maintained by approximately equal 

supplies of low saline water from the Baltic and 

high saline oceanic water from the Skagerrak. The 

nutrient supply to these waters increased rapidly 

during the past decades and oxygen deficits have 

been reported from different parts of the estuary. In 

this paper, we have calculated trends in nutrient and 

oxygen concentrations within the surface and deep 

waters of the Kattegat and adjacent waters. This has 

been done with available data for the past decades, 

with reference to nutrient supply and phytoplankton 

~roduction. 

Oxygen concentrations within the deep water 

decreased from 4.58 to 4.08 ml 1-I between 1971 

and 1982, indicating a 50% increase in oxygen 

consum~tion. Concentrations of Tot-N, Tot-P and 

inorganic nitrogen increased simultan~ously, both 

in the surface water during the winter and in the 

deep-water during the summer. Changes in Tot-N 

and Tot-P were dominated by the Baltic water, 

while local supply to the Kattegat dominated the 

changes in inorganic nitrogen. Increases in Tot-N 

and Tot-P suggest a successively increasing ~ ~ 

biomass. 

The importance of local nutrient supply to the 

Kattegat was studied comparing expected nutrient 

concentration within the surface water (due to 

exchange with adjacent water) with actually 

observed concentrations. 

Aneer, G. (1985) Some speculations about 

the Baltic herring (Clupea harengus 

m e m b r a s ) in connection with the 

eutrophication of the Baltic Sea. Canadian 

Journal of Fisheries and Aquatic Science 42 

(suppl.l):83-90. 

{Baltic; herring; filamentous; algae; egg; 

mortality) 

In this paper the hypothesis is put forward that 

Baltic herring's (Clupea harengus membras) 

spawning time, spring or autumn, is determined by 

feeding conditions during the adult phase and thus 

not generally fixed. The present "absence" or 

autumn spawners is thought to be the result of 

improved feeding condition during the latest decades 

as a result of the eutrophication of the Baltic Sea. 

During two spawning ground studies carried out in 

1978 and 1982 unusually high mortality rates were 

noted for eggs in situ. In 1982, during 4 week close 

to peak of spawning, the mortlaity increased 

substantially, especially for eggs among 

filamentous algae. A significant difference was 

noted between eggs on coarser algae and those 

among filamentous algae @ < 0.001). During this 

period the average mortalities were 33 and 75% 

respectively. Very low levels of oxygen were 

measured at night among the filamentous algae. An 

increase in the amounts of this type of algae as a 

response to eutrophication might constitute a new 

hazard to the reproductive success of the Baltic 

herring. 

Aneer, G. (1987) High natural mortality of 

Baltic herring (Clupea harengus) eggs 
ca&ed by algal exydates? Marine Biology 

94: 163-169. 

{Baltic; hemng; egg; toxic algal exudates; oxygen) 

Results from field studies in 1978 and 1982 

together with results from a laboratory study carried 

out in 1984 indicate that the unusually high natural 

mortality of Baltic herring (Clupea harengus L.) 

eggs observed in situ is presumably caused by toxic 

exudates released by filamentous brown algae 

during a limited period which happens to coincide 

with the peak of spawning. Oxygen deficiency is 

no longer believed to be the major cause of the 

observed unnaturally high mortality. It cannot, 

however, be entirely ruled out. 

Ansari, Z. A., B. S. Ingole, and A. H. Parulekar 

(1986) Effect of high organic enrichment 

of benthic polychaete pbpulation in an 

estuary. Marine Pollution Bulletin 17:361-365. 

{Benthic; polychaete; productivity; diversity; 

biomass; density; sewage) 

The benthic ~olvchaete fauna of an estuarine L. 

region receiving domestic sewage and wastes from a 

nearby fish landing jetty was compared to that of a 

site having normal organic enrichment. The 

population density, biomass and species diversity 

were greater at the sewage receiving site. Some 

species showed preference for sewage site and their 

numbers were significantly greater at the sewage 

site while others~ecies showed no differences. 

Total counts and the biomass were significantly 

higher at the site of organic enrichment. A 

quantitatively enhanced polychaete population 

indicates that input of-high organic matter 

stimulate benthic productivity. 

Antonius, A. (1985) Coral diseases in the 

Indo-Pacific: A first record. P. S.Z. N. I: 

Marine Ecology 6: 197-2 18. 

{coral; disease; Indo-Pacific; Red Sea; Philippines) 

The so-called "band" diseases of reef corals, the 

White Band Disease (WBD) and the Black Band 

Disease (BBD), were observed in the Red Sea and in 

the Philippines. Since they were previously known 

only from the western Atlantic, this is the first 

record for the Indo-Pacific region. WBD is neither 

infectious nor contagious and the pathogens is 

unknown. BBD is highly infectious and contagious 

and is caused by the cyanophyte Phormidium 

corallyticum. Susceptibility to WBD occurs 

throughout the order Scleractinia, but the effects of 

the disease are most severe among Acroporidae. 

Susceptibility to BBD seems to be restricted to 

Faviidae, with Platygyra and Goniastrea species the 

most heavily afflicted. WBD cannot be influenced 

by any parameter tested so far. BBD can be 

enhanced by light and by water eutrophication, and 

its advance can be stopped by antibiotics. An active 

WBD seems to be a precondition for the origin of 

BBD. 

Anttila, R. (1973) Effect of sewage on the 

fish fauna in the Helsinki area. OIKOS 

Supplementum 15:226-229. 

{sewage; fish; species) 

The results given are based on fishery studies 

begun by the City of Helsinki in 1969 on the 

effects of sewage on the fish fauna and fishing in 

the Helsinki sea area. The main research methods 

are test fishing with a series of nets, and fishery 

inquiries. 

Sewage has brought about considerable changes in 

the quality and quantity of fish fauna in the 

Helsinki sea area. The changes in the fish fauna are 

similar to those generally recognized as deriving 

from sewage. The fish species susceptible to 

pollution have suffered a decline andlor moved to 

purer areas. Such species are brown trout Salmo 

trutta(L)., whitefish Coregonus lavaretus (L.), 

burbot Lota lota (L.) and in Leusiscus idus (L.). Of 

the major edible fish, pikeperch Lucioperca 

lucioperca L. has withstood pollution best. 

The fish biomass in the more eutrophic bays is 

considerable in size compared with pure areas. In 

addition to the deterioration in the fish species 

structure there are also flavour defects in fish over a 

sea area of 3,000-4,000 hectares. Flavour defects in 

fish are caused by the secondary phenomena of 

sewage and oil that have entered the water. The 

mercury and pesticides contents observed in fish 

have been relatively low. 

Arndt, H. (1988) Dynamics and production 

of a natural population of Brachionus 

plicatilis (Rotaria, Monogonota) in a 

eutrophicated inner coastal water of the 

Baltic. Kieler Meeresforschungen, Sonderheft 

6: 147-153. 

{rotifer; Brachionus; growth; zooplankton; Baltic) 

The dynamic and production of a natural 

population of the rotifer Brachionus Plicatilis 

(O.F.MULLER) were studied by regular field 

sampling at routine stations in the shallow Darss 

Zingst estuary, southern Baltic .(3-7 %o S). 

Investigations of the horizontal distribution 

revealed significant population growth during the 

summer months at salinities above 3 %o S and at 

those stations characterized by high eutrophication. 

The first individuals hatch from resting eggs in 

May. Significant reproduction occurs from June to 

September, when temperatures are above 15 "C. 

Mixing rates were highest during the exponential 

growth phase. Instantaneous rates of growth, birth, 

and mortality were estimated. Mean P/B ratios for 

the growing season were high (around 0.7 d-l) and 

were in the range of values obtained from mass 

cultures in the field under subtropical conditions. In 

Barther Bodden annual biomass production for 1982 

was 1.1 g fw m-3. In its natural habitat, B. 
plicatilis serves as a food source for fish juveniles 

and the mysid Neomysis integer. The seasonal 

dynamics of the rotifer population were not 

significantly affected by predators. 

Arts, G. H. P., G. V. d. Velide, J. G. M. Roelofs, 

and C. A. M. V. Swaay (1990) Successional 

changes in the soft-water macrophyte 

vegetation of (sub)Atlantic, sandy, 

lowland regions during this century. 

Freshwater Research 24287-294. 

{vegetation; macrophytes) 

SUMMARY 

1. Considerable changes in macrophyte vegetation 

can be noticed in 146 originally soft waters, when 

data on the recent aquatic vegetation are compared 

with historical information from the period 1900- 

60. Changes in nutrient status (N, P, and C) and 

accumulation of organic material can be regarded as 

the operative factors. 

2. The processes observed in soft waters are 

acidification, eutrophication and water hardening. 

Which process dominates depends on the type of 

soft water. 

3. Acidification as well as eutrophication of water 

bodies may ultimately result in the total 

disappearance of all aquatic macrophytes, with the 

exception of the floating-leafed nymphaeids 

Nymphaea alba (L.) and Nuphar lutea (L.) Sm. 

Observed successional stages are described and 

summarized. 

Arzul, G. and P. Gentien (1990) Chemical 

typology of coastal sediments in relation 

to red tides. In: E. GranBli, B. Sundstrom, L. 

Edler and D. M. Anderson (ed.) Toxic Marine 

Phytoplankton. Elsevier, New York. 

{red tide; phytoplankton; bloom; organic matter; 

sediments; copper; nutrients; nitrogen; phosphorus; 

Alexandrium) 

The study of different chemical parameters 

characterizing the coastal sediments showed 

possible relationships between high levels of 

organic matter and total copper concentrations, and 

"red tide" occurrences. Sediment extracts had toxic 

effects on the growth of Alexandrium tamarensis in 

April, however, it was not observed in sediment 

elutriated in July. 

Aubert, M. (1992) Sanitary consequences of 

eutrophication and related ecological 

disequilibria in the marine environment. 

In: R. A. Vollenweider, Marchetti and R. Viviani 

(ed.) Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{nuisance bloom, dinoflagelate, Dinophysis) 

Considering the rates and dynamics of marine 

primary productivity it appears quite clear that 

besides the essential factors such as nutrient pools 

and inputs of energy, the function of growthpromoting or inhibitory telemediators (exocrines) 

present a very important control mechanism, in 

particular as far as the species compositions and 

periodic successions of phytoplankton are 

concerned. However, this mechanism also 

ultimately influences the overall equilibria. We 

must take into account many facts which show the 

importance of this last factor. 

Such massive, semi-monospecies dinolflagellate 

blooms are, almost without exception, 

environmentally harmful, many of them also 

leading to the toxicity of sea food and relevant 

human pathology. However, in specific cases, 

such as the DSP intoxications induced by a number 

of species of Dinophysis, even the slight increase 

in population densities, e.g. >200 cells dm-3, may 

cause quite dramatic pathology among shellfish 

consumers and, of course, also important economic 

losses for mariculture activities. 

Austen, M. C., R. M. Warwick, and M. C. Rosado 

(1989) Meiobenthic and macrobenthic 

community structure along a putative 

pollution gradient in Southern Portugal. 

Marine Pollution Bulletin 20:398-405. 

{benthos; Portugal; community; diversity; 

abundance; sewage) 

Macro- and meiobenthos were sampled from Ria 

Formosa, Portugal along putative sewage pollution 

gradients in summer 1987 and winter 1988. The 

status of the benthos was assessed using 

multidimensional scaling ordination, ABC plots, 

and a variety of univariate indices of community 

structure. Meiobenthos in both seasons and 

macrobenthos in winter appeared to respond to 

sewage enrichment only in the immediate vicinity 

of the sewage outfalls and channels but ABC plots 

indicated that macrobenthic communities were 

moderately stressed throughout the area in summer. 

In the summer, human digging for shellfish results 

in considerable sediment turnover. Since meiofauna 

may be less affected by physical sediment 

disturbance than macrofauna this may explain why 

the macrofauna communities were apparently 

disturbed relative to the meiofauna even beyond the 

influence of the sewage. Sampling meiofauna with 

the macrofauna significantly increased our 

understanding of mans impact in this area. There 

was little correlation between macro- and 

meiobenthos in diversity, abundances, number of 

taxa and biomass and these idices gave no 

indication of a organic enrichment 

Axelard, D. M., G. C. B. Poore, G. H. Arnott, J. 

Bauld, V. Brown, R. R. C. Edwards, and N. J. 

Hickman (198 1) The effects of treated 

sewage discharge on the biota of Port 

Philip Bay, Vitoria, Australia. In: B. J. 

Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 
{bacteria; nitrification; denitrification; sewage; 

microalgae; macrophytes; macrobenthos; 

zooplankton; phytoplankton; Australia; diversity; 

fish) 

The Werribee sewage-treatment farm contributes 

more than half of the total nitrogen and phosphorus 

input to Port Philip Bay. This study attempted to 

determine the fate of these nutrients and their effect 

on the biota of the Bay. This was addressed by 

comparing community composition, biomass, 

productivity, or process rates in the Werribee area 

of the Bay with that in Bay areas more remote from 

nutrient discharge. 

Rates of bacterial nitrification and denitrification 

were greatest in sediments closest to the sewage 

discharge point. Up to 15 percent of the inorganic 

nitrogen discharge may be lost via sequential 

nitrification-denitrification in a 4 km2 area. 

Epibenthic microalgal biomass and productivity 

were found to be five times greater at Werribee than 

at a control station. The Werribee macrophyte 

community showed reduced species diversity, 

dominance of fast growing opportunistic species, 

loss of large brown algal species, and occasional 

algal blooms, as compared to communities in 

nutrient-poor areas of the Bay. Classification 

analysis revealed offshore and nearshore groups of 

macrobenthos in a 2 km2 area near Werribee; 

species diversity was greatest offshore. The 

Werribee offshore macrofauna was typical of that 

along the whole northwestern coast of the Bay. In 

summer, fish biomass at Werribee was equal to that 

found at stations remote from sewage discharge, 

however, community composition differed. At 

Werribee, the nearshore fish community was 

dominated by juveniles and small species, while 

offshore older and larger fish were more abundant. 

Phytoplankton productivity decreased with 

increasing distance from sewage outfalls only 

during summer. Nitrogen is probably the nutrient 

critical to phytoplankton biomass production in the 

Bay but light and/or temperature may limit 

productivity over much of the non-summer period. 

Baywide phytoplankton productivity was similar to 

that of non-eutrophic coastal marine waters. 

Zooplankton standing crop was low compared to 

that for many estuaries and marine embayments. 

Densities of zooplankton in the Werribee area were 

highly variable, and crustacean species found at 

Wembee were also distributed throughout the Bay. 

These findings suggest that sewage discharge had 

affected benthic more than planktonic communities, 

but that the measurable impact of the discharge is 

limited to a few hundred meters around the outfalls. 

Baden, S. P., L. 0. Loo, L. Pihl, and R. 

Rosenberg (1990) Effects of eutrophication 

on benthic communities including fish: 

Swedish west coast. Ambio 19: 113-122. 

{oxygen; hypoxia; seaweeds; macroalgae; increase; 

decrease; macrofauna; lobster; fish; floating; 

mortality) 

The southern Kattegat is susceptible to 

eutrophication due to shallow mean depth (23m) 

and a strong halocline which reduces intrusion of 

oxygen to bottom waters. The effects of 

eutrophication were first observed in the area in the 

autumn of 1980. Since then investigations on 

primary producers, fish, lobsters, and benthic 

infauna have been conducted to document the effects 

of eutrophication. Above the halocline in the 

Laholm Bay a change in the macrophyte species 

from Fucus spp. to filamentous green algae has 

been observed. Mortality of benthic macrofauna, 

mainly bivalves, has been observed in most years 

and seasons. The recruitment of flatfish has not 

been negatively affected. Around the halocline the 

benthic infauna were seriously affected by oxygen 

deficiency. Below the halocline, fish disappeared 

and lobsters emerged from burrows when oxygen 

saturation declined below 40%. When oxygen 

saturation decreased below 15% Norway lobsters 

were immobilized and their blood-pigment 

concentration decreased. Benthic infaunal species 

emerged from the sediment. Lobsters died when 

saturation dropped to lo%, while many infaunal 

species tolerated levels of 7-5% for some weeks. 

Analysis of stomach content indicated that neither 

fish nor lobsters died from lack of food, but from 

hypoxia. Subsequent to the reoxygenation of the 

bottom water during winter, flatfish and benthic 

infauna recovered whereas cod and lobster 

population did not. 

Beatty, L. L. (1991) The response of benthic 

suspension feeders and their grazing 

impact on phytoplankton in eutrophied 

coastal ecosystems. Dissertation for Ph.D., 

The University of Rhode Island. 

{benthos; MERL; nutrient; composition; 

abundance; biomass) 

The eutrophication of coastal waters from landderived nutrients affects all biological aspects of the 

estuarine ecosytem. Changes to the benthic 

community can be pronounced. Not only do 

benthic species respond to nutrient-induced 

phytoplankton blooms, but these animals may 

have a subsequent impact on the system. 

A sixteen-month study was conducted with daily 

nutrient loading. One of the nutrient levels included 

enhanced silicon to promote diatom growth and 

potentially lead to a healthier food chain. Cores 

were taken monthly to follow changes in benthic 

species composition. Laboratory experiments were 

performed on five occasions from March to October 

to determined the role of benthic grazing in control 

versus nutrient-enriched systems. Cores were taken 

from mesocosms and removal rates of natural 

assemblages of phytoplankton were determined. 
A two to five-fold increase in abundance and 

biomass was observed in enriched over control 

mesocosms. A shift in species composition 

included the rapid response of facultative 

suspension-feeding spinoid polychaetes, Polydora 

ligni and Streblospio benedicti, and amphipod, 

Ampleisca abdita and Corophium sp., to 

phytoplankton compared to the other nutrient 

systems, although differences were not statistically 

significant by the second summer. Zooplankton 

initially responded to phytoplankton blooms but 

adult benthic suspension feeders dominated the 

grazer biomass within three months of nutrient 

additions. 

Benthic grazers could remove up to 75% of water 

column phytoplankton per day. Rates were higher 

in nutrient-enriched systems where animal biomass 

was greater than in control closures. Phytoplankton 

>10 um and water temperatures over 15°C were 

also significant factors in observed feeding rates. 

Benthic species successfully competed with 

zooplankton for food resources; alttough their 

weight-specific feeding rates are lower than those 

for copepods, opportunistic characteristics of 

spinoid polychaetes and amphipods and their ability 

to deposit feed when water column phytoplankton 

is depleted contribute to their success. 

The results of this study are most applicable to 

shallow, well-mixed coastal areas. As water column 

depths decrease and/or food is more accessible to the 

benthos, benthic-pelagic coupling increases in 

importance. 

Beauchamp, S. T. and J. Kerekes (1989) Effects 

of acidity and DOC on phytoplankton 

community structure and production in 

three acid lakes (Nova Scotia). Water, Air, 

and Soil Pollution 46:323-333. 

{acidity; lakes; phytoplankton; assemblage) 

Phytoplankton community structure varied 

between the three lakes and between years within 

lakes. The Beaverskin Lake phytoplankton 

community was dominated by cyanophytes and 

chlorophytes in the summer and chrysophytes in 

the winter. Kejimkujik Lake was dominated by 

bacillariophytes in the summer of 1979 but no 

single group dominant in 1980 or 1981. 

Pebbleoggitch Lake phytoplankton consisted 

mainly of chlorophytes in 1979 but low biomass 

and no dominant groups characterized this lake 

during the growth season of 1980. Daily integral 

planktonic primary production measured 

simultaneously in the three lakes showed that in 

both years annual planktonic primary production 

was highest in the clear water lake, Beaverskin 

Lake, which also had lower dissolved organic 

carbon (DOC) concentration compared to the two 

dystrophic lakes. In the clear water lake annual 

production was similar between years but in the 

two colored lakes annual production was 40% 

higher in the second year. The observed increases in 

annual production between years in the colored 

lakes were largely due to changes in euphotic depth-. 

resulting from variations in hydrology and DOC 

export from the lake catchments. Lower discharges 

in the colored lakes in 1980 were accompanied by 

lower DOC concentration in the clear water lake did 

not produce significant changes in water color, 

light extinction coefficient nor annual production 

between year. Rates of primary production at light 

optimum (P-max) were consistently higher in the 

most colored, acidic lake indicating that relatively 

high rates of autotrophic production will occur 

under acidic conditions if nutrient supply is 

maintained. 

Becker, D. S., N. L. Gerrish, and K. K. Chew 

(1981-1982) Influence of organic 

enrichment on demersal fishes. Coastal 

Ocean Pollution Assessment News 1:50-52. 

{fish; organic enrichment; demersal; Puget Sound) 

No abstract 

Bell, P. F. (1991) Status of eutrophication 

in the Great Barrier Reef lagoon. Marine 

Pollution Bulletin 23:89-93. 

{lagoon; phytoplankton; nutrients) 

Historical data on the levels of nutrients and 

phytoplankton in the GBR lagoon are reviewed. 

The results indicate that background levels of PPO4 and phytoplankton have increased significantly 

over the past 50-60 years and that the levels appear 

be at or above the eutrophication threshold level for 

coral reef waters. Other data indicate that river 

discharge probability has a major impact on the 

nutrient status of the GBR lagoon, but other factors 

such the nitrogen-fixing blue-green alga, 

Trichodesmium, could also be important. 

Trichodesmium has the ability to introduce large 

amounts of new nitrogen and it appears that the 

increased phosphorus levels could be driving its 

growth. To-date little effort has been made to assess 

the impact of eutrophication on the coral reef 

communities. Because the background nutrient 

levels are relatively high both run-off and sewage 

discharges could have serious impacts on nearby 

coral reef communities. Tertiary treatment (i.e. 

nutrient removal) of sewage should be required for 

all discharges in the vicinity of coral reefs and 

special precautions need to be exercised when 

designing run-off drainage systems. 

Berman, T. and Z. Dubinsky (1985) The 

autoecology of Peridinium cinctum fa. 

westii from Lake Kinneret. Verhandiunsen 

der Internationalen Verlaasbuchhandlune 22:2850- 

2854. 

{autoecology; Peridinium; pH; temperature; 

dinoflagellate; Lake Kinneret; freshwater) 

No abstract 
Berounsky, V. M. and S. W. Nixon (1985) 

Eutrophication and the rate of net 

nitrification in a coastal marine 

ecosystem. Estuarine, Coastal and Shelf Science 

20~773-78 1. 

{nitrification; estuaries; nitrogen; nutrients) 

Rates of nitrification were calculated for four large 

(13 m3) estuarine-based microcosms that had been 

subjected to inorganic nutrient enrichment. 

Calculated rates were based on two years of weekly 

nitrate and nitrite measurements and ranged from a 

maximum of 0-55 umol NO-2+3 produced l-lday-l 

in the control tank (no enrichment) to over 13 

umol NO-2+3 produced l-lday-l in the most 

enriched tank (receiving 18.6 umol NH4 1-lday-l). 

Almost all NO-2+3 production was pelagic, little 

was benthic. Net NO-3 production or net 

production dominated the net nitrification rates 

during different seasons. Good correlations were 

found between various oxidation rates and substrate 

concentrations. The calculated net nitrite production 

rates were 10 to 1000 times higher than previously 

reported rates for open ocean systems, 

demonstrating the potential importance of 

nitrification to estuarine systems. 

Beukema, J. J. (1990) Long-term and recent 

changes in the benthic macrofauna living 

on tidal flat in the western part of the 

Wadden Sea. Proceedings of the 7th 

International Wadden Sea Symposium, Ameland, 

The Netherlands. 

{benthic; macrofauna; tidal flat; Wadden Sea; 

species abundance; biomass) 

The bottom fauna living on tidal flats in the 

westernmost part of the Wadden Sea were followed 

quantitatively by biannual sampling at 15 fixed 

stations throughout the 1970-1990 period. During 

this period, the following long-term changes were 

observed; 

1. biomass doubled and numerical abundance more 

than doubled; as a consequence the size of a 'mean' 

macrobenthic animal declined; 

2. the abundance of more than half of the species 

studied increased significantly; 

3. among these species, small deposit-feeding 

worms were over-represented and some species 

increased more than other species. 

The above changes are attributed,to the recent 

eutrophication of .the area (CADEE 1992). A 

favorable effect is the enhanced stock and 

productivity of the benthos. Detrimental effects 

(caused by prolonged periods of oxygen depletion) 

occurred incidentally and locally. recent (1988- 

1990) changes include a marked increase in the 

species which are known to be sensitive to low 

winter temperatures and are ascribed to the mild 

character of the last three winter. The stock of 

mussels declined during 1990 to unprecedently low 

levels. All intertidal mussel beds were completely 

destroyed by commercial fishermen to stock their 

subtidal plots. It is recommended that extensive 

tidal-flat areas should be closed for such fishery 

activities. 

Beukema, J. J. (1991) Changes in 

composition of bottom fauna of a tidalflat area during a period of 

eutrophication. Marine Biology 111:293-301. 

{benthos; faunal composition; tidal-flat; Dutch 

Wadden Sea; production; abundance; biomass) 

During the last 20 yr the western half of the 

Dutch Wadden Sea has undergone significant 

eutrophication: concentrations of P and N 

compounds and planktonic algae have roughly 

doubled, as has primary production. Though 

oxygen levels are often low in summer, anoxic 

areas are small and rare due to strong tidal mixing. 

During the 1970 and 1990 period, macrozoobenthos 

was sampled annually at 15 stations at Balgzand, a 

50-km2 tidal-flat area in the westernmost part of 

the Wadden Sea. Not only did the estimates of total 

numbers, biomass, and production double during 

these two decades, but significant changes in the 

composition of the benthic community were 

observed, too: (1) the numerical proportion of 

polychaetes increased at the expense of molluscs 

and crustaceans, (2) the overall mean weight per 

individual of the macrozoobenthos decreased 

(numbers of individuals of small-size species 

increased more rapidly than those of large-sized 

species), and (3) though absolute numbers and 

biomass of all feeding types increased, the share of 

carnivores declined and that of deposit feeders 

increased; the proportion of suspension feeders 

showed little change. This study refers to true 

macrobenthos only (I-mm sieve) and further 

excludes two taxa (Corophium spp. and Hydrobia 

ulvae) which occasionaly exercised an undue 

influence on numbers. Mass mortalites caused by 

low oxygen concentrations were of a small-scale 

nature only. Total number of species fluctuated 

without a clear trend. As a consequence of the 

increasing numerical densities, trends in species 

numbers were slightly increasing when expressed 

per unit area and slightly decreasing when estimated 

per 100 individuals (by rerefaction). 

Beukema, J. J. and G. C. Cadte (1986) 

Zoobenthos responses to eutrophication 

of the Dutch Wadden Sea. Ophelia 26:55- 

64. 

{benthos; Dutch Wadden Sea; biomass; secondary 

production; abundance) 

During the last few decades nutrient 

concentrations in the Dutch coastal water have 

increased significantly. Values observed for rates of 
primary production as well as for concentrations of 

chlo rophyll and for particulate organic matter 

have been higher during recent years than estimates 

from earlier periods. Thus more food will have been 

available for herbivorous benthos during recent 

years. 

Both biomass and annual production of the 

macrozoobenthos living on the tidal flats in the 

western part of the Wadden Sea doubled during the 

1970 to 1984 period. More than half of the species 

contributed to-this increase. In the species studied 

in most detail, Macoma balthica, both reproductive 

success and growth rate increased significantly 

during the 15-year period. 

Though a causal relationship between 

simultaneous increases of nutrient levels, primary 

production and secondary production cannot be 

proved, such a relationship seems to be natural. So 

far. obvious detrimental effects such as mass 

mortalities from lack of oxygen appear to be rare in 

the well-mixed Wadden Sea. 

Beukema, J. J. and G. C. Cadte (1991) Growth 

rates of the bivalve Macoma balthica in 

the Wadden Sea during a period of 

eutrophication: relationships with 

concentrations of pelagic diatoms and 

flagellates. Marine Ecology Progress Series 

68:249-256. 

{phytoplankton; bloom; flagellates; Wadden Sea; 

diatom; bivalve; growth; Macoma; secondary 

production} 

Probably as a consequence of eutrophication, both 

the length of the annual phytoplankton blooms and 

the mean concentrations of flagellates (dominated 

by Phaeocystis pouchetii) have increased in the 

western part of the Wadden Sea during the last 

decades. Planktonic diatoms, on the other hand, 

have hardly increased, but fluctuate heavily from 

year to year. Among the benthic tidal-flat fauna, 

which has increased in total numbers and biomass 

during the last decades, the bivalve Macoma 

balthica was studies in detail. The 15 year (1974 to 

1989) patterns of its annual growth and condition 

were compared with abundance patterns in 

phytoplankton components (during the growing 

seasons of M. balthica). Fluctuations in M. 

balthica were found to parallel fluctuations of 

planktonic diatoms but not those of P. pouchetii. It 

is concluded that M. balthica responded by faster 

growth and higher condition to better feeding 

conditions in years of high diatom abundance. 

Diatoms rather than P. pouchetii or other 

flagellates will be the most important food source 

for M. balthica. 

Biggs, R. B., T. B. DeMoss, M. M. Carter, and E. 

L. Beasley (1989) Susceptibility of US 

estuaries to pollution. Review in Aquatic 

Sciences 1: 189-207. 

{estuary; classification; susceptibility) 

No abstract 

Birch, P. B. and J. 0. Gabrielson (1984) 

Cladophora growth in the Peel-Harvey 

estuarine system following blooms of the 

cyanobacterium Nodularia spumigena. 

Botanica Marina 27: 17-21. 

{bloom; cyanobacteria; estuary; macroalgae; 

Cladophora; Nodularia, seaweeds} 

In recent years severe blooms of the nitrogenfixing cyanobacterium Nodularia spumigena have 

occurred during spring and early summer in the 

eutrophic Peel-Harvey Estuary. Following 

decomposition of these blooms there are blooms of 

Cladophora and other macroalgae, and more recently 

of a benthic species of Oscillatoria. Data on 

nitrogen and phosphorus in estuary water and in 

Cladophora tissue, combined with a decomposition 

study of Nodularia, indicate that large amounts of 

nutrients are recycled from Nodularia to other 

algae. Control of Nodularia blooms is now of 

primary importance not only because of serious 

problems caused directly by it, but also because of 

the nutrients made available upon its decay for 

other nuisance algae and bacteria. 

Birch, P. B., J. 0. Gabrielson, and K. S. Hamel 

(1983) Decomposition of Ckdophora. 

Botanica Marina 26: 165- 17 1. 

{algae; Cladophora; estuary; Australia; 

decomposition; nitrogen; phosphorus) 

Cladophora aff. albida is a benthic alga which 

grows in nuisance proportions in the shallow (2m) 

Peel-Harvey estuarine system in Western Australia. 

It grows as small (1-3 cm) ball-like clumps in beds 

1 - 10 cm deep, the lower self-shaded sections of 

which are in various stages of decomposition. 

Samples of Cladophora (half of which were prekilled by freezing) were buried in a bed in nylon 

mesh for up to one year. Initially live Cladophora 

was very resistant to decay; only after two months 

did significant decomposition begin, resulting in 

loss of 50-60% of dry weight, carbon and nitrogen 

losses were slower (20-30% in two weeks, 50% 

after three months). 

It was estimated that up to 60% of the 

Cladophora pool of nitrogen and phosphorus could 

be available for recycling in one year. This reduces 

the dependence of Cladophora on external inputs to 

maintain production. 

Blackburn, S. I. and Y. Oshima (1989) Review 

of culture methods for Pyrodin ium 

bahamense. In: G. M. Hallegraeff and J. L. 

Maclean (ed.) Biology, Epidermiology and 

Management of Pyrodinium Red Tides. Fisheries 

Department, Ministry of Development, Manila. 

The dinoflagellate Pyrodinium bahamese, both 

var. bahamense and var. compressum, has been 
successfully cultured in seawater-based media by 

several workers. These cultures were sometimes 

kept for many years but all of them were eventually 

lost. This tropical dinoflagellate requires 

temperatures of 24-30°C, dilute nutrient media and 

soil extract, while a pH optimum of 8 has also 

been observed. Recommendations for renewed 

culture attempts are provided. 

Boddeke, R. and P. Hagel (1991) 

Eutrophication of the North Sea 

continental zone. Netherlands Institute for 

Fisheries Research, E:7. 

{North Sea; shellfish; fish} 

North Sea landings of demersal fish rose after 

1963 to more than 1 millions tons from a rather 

constant level of about 400,000 tons since 1909. 

Landings of bivalves from the Dutch coastal zone 

increased greatly after 1950. From the data 

presented, a structural change in productivity of the 

Southern and Central North Sea in the period 1950- 

1985 as the result of eutrophication of the 

continental coastal waters emerges as the most 

likely cause of the enhanced commercial 

production. Therefore, the continuing decrease of 

the discharge of phosphate to the southern North 

Sea since the beginning of the eighties, is likely to 

have negative effects on the production of fish and 

shellfish in the S-E North Sea. 

Bodeanu, N. (1992) Algal blooms and 

development of the main phytoplanktonic 

species at the Romanian Black Sea 

littoral in conditions of intensification 

of the eutrophication process. In: R. A. 

Vollenweider, R. Marchetti and R. Viviani (ed.) 

Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{species abundance, species composition, Black 

Sea) 

The growth of mineral salts and organic matter in 

the western Black Sea, caused by intensification in 

the 1980s and 1990s of anthropic activity in the 

pontic basin, the Danube and other tributary rivers, 

determined the increase of the frequency and 

magnitude of algal blooms with their detrimental 

consequences for ecosystems. More than 20 

monospecific algal blooms have occured in 

Romanian coastal waters between 1983 and 1988. 

Of the eight species responsible for the 

phenomenon, five achieved the greatest blooms 

ever known at the Romanian littoral. 

With the exception of algae which produce 

monospecific blooms, remarkable development of 

numerous other species .has been recorded. The 

number of species with high numerical densities 

(over 100,000 cells 1-I), increasing from one 

period to the next, reached 72 in 1983-1988, 

compared with 61 in 1971-1982 and only 38 in 

1960-1970. Of the total mass species which 

developed between 1960-1988, almost 60% reached 

the greatest development during the years 1983- 

1988. The intensity of the blooms, the growth of 

the mass species number, and the increase in 

density induced high levels of global photplankton. 

Continually rising, the mean biomass for the 

Romanian Black Sea areas was ten ti;mes greated in 

1983-1988 than in 1959-1963. 

Boesch, D. F. (1983) Implications of oxygen 

depletion on the continental shelf of the' 

Northern Gulf of Mexico. Coastal Ocean 

Pollution Assessment News 2:26-28. 

{oxygen; hypoxia; Gulf of Mexico; nutrient) 

No abstract 

Boni, L., E. Carpent, D. Wynne, and M. Reti 

(1989) Alkaline phosphatase activity in 

Protogonyaulax tamarensis. Journal of 

Plankton Research 11:879-885. 

{bloom; pH; alkaline phosphatase; indicator; 

dinoflagellates) 

A non-toxic strain of the marine dinoflagellate 

Protogonyaulax tamarensis (= Gonyaulaz 

tamarensis) has been isolated from a bloom in the 

Adriatic Sea, off the Emilia-Romagna coast. 

Cultures of the cells were grown in the laboratory 

in enriched seawater at various initial ambient 

orthophosphate (Pi) concentrations, ranging from 

0.3 to 40 uM. The growth rate varied from 0.3 to 

0.8 divisions day-l, depending on the Pi levels. 

From measurements of kinetic parameters, the 

binding of the artificial substrate p - 

nitrophenylphosphate to the P. tamarensis alkaline 

phosphatase was quite strong (Km = 50 uM). 

Maximal activity was observed at pH 8.4, although 

the pH-activity curve was broad, in contrast to that 

of other alkaline phosphatases. Protogonyaulax 

tarnarensis alkaline phosphatase, measured over a 

24 h period, exhibited an apparent diurnal 

fluctuation in activity, in common with the 

enzyme from other dinoflagellates. 

Boni, L., L. Mancini, A. Milandri, R. Poletti, M. 

Pompei, and R. Viviani (1992) First cases of 

diarrhoetic shellfish poisoning in the 

Northern Adriatic Sea. In: R. A. 

Vollenweider, R. Marchetti and R. Viviani (ed.) 

Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{Adriatic Sea, toxicity} 

Shellfish poisoning occurs when toxic 

photoplankton species are present in their 

environment. Because shellfish such as mussels 

are filter feeders they concentrate toxins. It is also 

well known that sometimes also small amounts of 

toxic algae can produce serious shellfish poisoning. 

In the Adriatic Sea, dinoflagellate red tides are a 

recurring phenomenon, but no shellfish poisoning 
has been described despite the presence of 

potentially toxic species such as Alexandrium spp. 

(Protogonyaulax) and Dinophysis spp. In June 

1989 several cases of food poisoning characterized 

by vomiting, abdominal pain and diarrhoea were 

recorded along the coast of Emilia-Romangna and 

Marche, facing the north-western Adriatic Sea. The 

poisoning was attributed to shellfish ingestion, 

mostly mussels. Microscopic investigations 

revealed the presence of Dinophysis spp. in 

seawater and mussel guts. My ti 1 us 

galloprovincialis, Venus gallina, Tapes 

semidecussatus, and Venus verrucosa have been 

monitored for toxin by mouse bioassay and 

diarrhoetic shellfish poisoning (DSP) was detected 

only in mussels. Shellfish samples from Friuli 

Venezia Giulia, Veneto, Emilia-Romagna and 

Marche were monitored for toxin and from June 22 

a temporary prohibition on mussel fishing for the 

period of DSP was imposed. 

Borum, J. (1985) Development of epiphytic 

communities on eelgrass (Zostera marina) 

along a nutrient gradient in a Danish 

estuary. Marine Biology 87:211-2 18. 

{eelgrass; epiphytes; community; estuary; 

biomass; phytoplankton; nutrients) 

The effect of nutrient enrichment on epiphyte 

development was examined by following the 

seasonal development of epiphyte biomass on 

eelgrass (Zostera marina L.) at four localities along 

a nutrient gradient in Roskilde Fjord, Denmark 

between March and December 1982. In the most 

nutrient-poor area, epiphyte biomass followed a 

distinct bimodal seasonal pattern with maxima in 

spring and early fall. Low nutrient availability and 

a high rate of eelgrass leaf renewal kept epiphyte 

biomass at a low level throughout the summer 

period. Unlike phytoplankton, the epiphytic 

community was not stimulated by nutrient 

enrichment during spring, however from May 

though August, the biomass of both components 

increased exponentially with increasing 

concentrations of total N in the water. Along the 

nutrient gradient, phytoplankton biomass increased 

5- to 10-fold, while epiphyte biomass increased 50- 

to 100-fold. Thus differences in nutrient conditions 

among study sites were more clearly reflected by 

epiphytes than phytoplankton. 

Boynton, W. R., W. M. Kemp, and C. W. Keefe 

(1 9 82) A comparative analysis of 

nutrients and other factors influencing 

estuarine phytoplankton production. In: 

V. S. Kennedy (ed.) Estuarine Comparisons. 

Academic Press, New York. 

{phytoplankton; productivity; estuary; biomass; 

production; N/P ratio; nitrogen; phosphorus} 

We reviewed data concerning phytoplankton 

production, chlorophyll a, and associated physical 

and chemical variables from 63 different estuarine 

systems. Data were analyzed statistically to test 

hypotheses regarding algal productivity and factors 

regulating temporal patterns. Prior to statistical 

analysis, estuarine systems were classified into four 

groups based on criteria of physical circulation and 

geomorphology. Analysis of grouped data indicated 

that algal production and biomass were consistently 

high in warm periods of the year in a broad 

spectrum of estuaries and that ratios of available 

nitrogen to phosphorus were low during periods of 

high production, except in highly eutrophic 

systems. In general, phytoplankton production and 

biomass exhibited weak correlations with a variety 

of physical and chemical state variables, perhaps 

indicating the significance of rate processes as 

opposed to standing stocks in regulating these 

important features of estuarine systems. A six-year 

time series of measurements of algal production and 

chlorophyll a at stations in middle Chesapeake Bay 

exhibited considerable year-to-year variability, with 

a three-fold range in peak values. Summertime 

maxima were strongly related to annual loadings of 

both nitrogen (N) and phosphorus (P) but annual 

production appeared to be sustained primarily on 

recycled nitrogen and phosphorus. To generalize 

from these findings, N and P loading rates were 

estimated for 14 different estuarine systems, and a 

significant positive relationship was obtained 

between phytoplankton production and nitrogen 

(but not phosphorus) inputs. 

Brand, L. E., M. D. Gottfried, C. C. Baylon, and 

N. S. Romer (1991) Spatial and temporal 

distribution of phytoplankton in 

Biscayne Bay, Florida. Bulletin of Marine 

Science 49599-613. 

{phytoplankton; abundance; distribution; Florida; 

pigments) 

Biscayne Bay provides a study in contrasts. North 

Biscayne Bay has undergone substantial changes 

during the twenties century resulting from the 

development of the metropolis of Miami around it 

while south Biscayne Bay has been relatively little 

affected by the dramatic human population increase 

in South Florida. 

The phytoplankton at 24 stations throughout 

Biscayne Bay were examined monthly for 1 year. 

Phytoplankton were found to be 5 to 10 times 

higher in abundance in the north bay compared to 

the south and even more abundant near canal 

mouth. Eutrophication from freshwater runoff is 

the likely cause of the higher abundance of 

phytoplankton in the north bay as well as the 

higher fluorescence response indices and 

chlorophyll to phaeopigment ratios in the 

phytoplankton in the north. Phytoplankton are 

most abundant in the north bay right after the first 

major rains of the rainy season, thought to be the 

result of the large amounts of nutrients flushed into 
the bay at that time. Phytoplankton are least 

abundant at the end of the dry season after a long 

period of little runoff. Phytoplankton abundance 

shows virtually no seasonality in the south bay 

where land runoff is relatively insignificant. The 

south Biscayne Bay ecosystem remains dominated 

by benthic macrophytes while the north bay shifted 

to an ecosystem dominated by a relatively dense 

plankton community with a few macrophytes 

remaining. 

Breuer, G. and W. Schramm (1988) Changes in 

macroalgal vegetation of Kiel Bight 

(Western Baltic Sea) during the past 20 

years. Kieler Meeresforschungen, Sonderheft 

6:241-255. 

{Baltic; Kiel Bight; Furcellaria; Phyllophora; 

Phycodrys; abundance; biomass; coverage) 

A large scale quantitative survey of the benthic 

vegetation of the Kiel Bight (Western Baltic Sea) 

has been performed by analysis of underwater 

television observations and samples obtained by 

SCUBA diving during 1985-86. This investigation 

was compared to a semiquantitative survey carried 

out in 1962-64 by Schwenke (1964, 1964). For the 

total study (2571 km2), distinct changes in 

biomass and species composition have been 

observed. There is an increase of biomass above the 

12 m level (probably with exception of the 6 m 

level) and a decrease below 12 m. Extensive 

Furcellaria lumbricalis populations have 

disappeared. Furcellaria lumbricalis has been 

replaced by Phyllophora truncata and phycodrys 

rubens which are the predominant species at 

present. Among other possible causes for these 

changes, the role of commercial stone fishing and 

effects of increasing eutrophication in the Kiel 

Bight are discussed. 

Brey, T. (1986) Increase in 

macrozoobenthos above the halocline in 

Kiel Bay comparing the 1960s with the 

1980s. Marine Ecology Progress Series 28:299- 

302. 

{macrozoobenthos; Kiel Bay; biomass; mollusc) 

The macrobenthos of sandy sediments in the 9 to 

13 m water depth area of Kiel Bay was investigated. 

Data from 1961 to 1965 were compared with data 

from 1982183. Biomass of total macrobenthos was 

significantly higher in 1982183 (14.6 g ash-free dry 

weight mm2) than in 1961 to 1965 (4.7 g AFDW 

m-2). This increase is mainly caused by mollusc 

species. Eutrophication, changes in predation 

pressure and changes in physical disturbance are 

discussed as possible reasons. 

Brockmann, U., G. Billen, and W. W. C. Gieskes 

(1 9 8 8) North Sea nutrients and 

eutrophication. In: W. Salomons, B. L. 

Bayne, E. K. Duursma and U. Forstner (ed.) 

Pollution of the North Sea. An Assesssment. 

Springer-Verlag, Berlin. 

{North Sea; nutrients; oxygen; effects; bloom; 

review) 

No abstract 

Brockmann, U. and E. Dahl (1990) 

Distribution of organic compounds 

during a bloom of Chrysochromulina 

polylepis in the Skagerrak. In: E. Grandi, 

B. Sundtrom, L. Edler and D. M. Anderson (ed.) 

Toxic Marine Phytoplankton. Elsevier, New York. 

{bloom; Chrysochromulina; phytoplankton; 

inorganic nutrients; organic; Skagerrak; Kattegat} 

Nutrient elements, bound in dissolved organic 

substances, can become a major controlling factor 

for algal growth when inorganic nutrients are 

depleted. During the late phase of the extended 

bloom of Chrysochromulina polylepis in the 

SkagerraklKattegat during spring 1988, 

concentrations of particulate and dissolved organic 

substances (C, N ,P) were estimated together with 

hydrographic measurements, and nutrient and 

phytoplankton analyses. Along a profile through 

the Skagerrak between Skagen and Ha110 a stable 

and most biologically rich pycnocline at 7-25 m 

depth separated two distinct chemical regimes: (i) a 

surface layer where nitrate (<0.2 ug at N dm-3) and 

phosphate (<0.02 ug at P dm-3) were nearly 

depleted and dissolved organic compounds (DOM) 

reached maximum concentrations (>lo ug at N dmand >0.05 ug at P dm-3), (ii) lower layers with 

higher inorganic nutrients and lower DOM. As a 

result of this, the pycnocline exhibited a minimum 

concentration of total dissolved nitrogen (40 ug at 

N dm-3) together with maximum cell densities 

(>16x106 cells dm-3) of C. polylepis. Because of 

these vertical nutrient profiles, the cells could 

utilize both inorganic and organic bound nutrient 

elements from the adjacent layers. The main Nsource was however probably nitrate as seen from 

mixing curves. High concentrations of dissolved 

carbohydrates (>3 umol Glc-eq dm-3) in the mixed 

layer, significantly negative correlated with 

salinity, indicate that the dominating source for 

DOM was the Baltic outflow. 

Brockmann, U. H., E. Dahl, and K. Eberlein 

(1 985) Nutrient dynamics during a 

Gyrodinium aureolum bloom. In: D. M. 

Anderson, A. W. White and D. G. Baden (ed.) 

Toxic Dinoflagellates. Elsevier, New York. 

{nutrient; dinoflagellate; phytoplankton; bloom} 

Monospecies cultures of Gyrodinium aureolum 

were investigated in two 16 and 20 m deep 

enclosures for 20 and 22 days respectively. 

Nutrients were added at the beginning and after 10 

or 12 days, simulating coastal front mixing. In 

spite of nutrient exhaustion in the upper layer 
(nitrate below 0.1 ug at N dmm3) the cells did not 

migrate to the nutrient-sufficient lower layer. After 

two days of nutrient depletion, G. aureolum 

maintained its high cell number. However, the cells 

decreased in the second enclosure during 5 days of 

nutrient deficiency and did not increase following 

the addition of nutrients, so that other species 

become dominant, including diatoms. 

Brown, J. R., R. J. Gowen, and D. S. McLusky 

(1987) The effect of salmon farming on 

the benthos of a Scottish sea loch. 

Journal of Experimental Marine Biology and 

Ecology 109:39-5 1. 

{benthos; abundance; biomass; fish farming; 

diversity) 

The effects of waste from a salmon farm on the 

benthos of a fjordic sea loch on the western coast of 

Scotland have been studied. Within 3 m of the 

floating cages the sediments was highly reducing, 

and dissolved oxygen content of the water overlying 

the sediment ranged from 35 to 75% saturation. At 

215 m from the cages the sediment was 

oxygenated, and the dissolved oxygen content of the 

water overlying the sediment was 50-80% 

saturation. Sedimentary redox potential and 

dissolved oxygen content of bottom water showed a 

seasonal variation. The benthic fauna showed 

marked changes in species number, species 

diversity, faunal abundance, and biomass in the 

region of the fish farm, with four zones of effect 

identified. Directly beneath, and up to the edge of 

the cages, there was an azoic zone. A highly 

enriched zone, dominated by Capitella capitata 

(Fabricius) and Scolelepis fuliginosa (Claparbde), 

occurred from the edge of the cages out to =8 m. A 

slightly enriched "transitional" zone occurred at 125 

m, and a "clean" zone at the distances >25 m. This 

study showed that salmon farming had similar 

effects on the benthos as other forms of organic 

enrichment, but the effects were limited to a small 

area in the immediate vicinity of the cages. 

Buhl-Jensen, L. and J. H. Fossi (1991) 

Hyperbenthic crustacean fauna of the 

Gullmarfjord area (western Sweden): 

species richness, seasonal variation and 

long-term changes. Marine Biology 109:245- 

258. 

{benthic; fauna; change; amphipods; disappearance) 

Distribution of 118 species of Amphipoda, 

Mysidacea and Decapoda, sampled in 1984 and 

1985 with an epibenthic sledge along a transect 

from the Skagerrak to the inner Gullmarfjord 

(Sweden), is described. Amphipods were richest in 

species in the Skagerrak, while most mysid species 

were taken at the sill. Deca-impoverished and the 

Lilljeborgidae, earlier represented by three species, 

had disappeared since 1930s. The changes are most 

likely caused by eutrophication of the fjord and 

deep-basin samples; (111) sill samples; (IV) 

Skagerrak samples. Characteristic species of the 

groups were identified with a pseudo F-test. 

Detrended Correspondence Analysis indicated 

seasonal faunal changes at 42 and 72 m depths. At 

both depths a shift in numerical dominance from 

amphipods to mysids was found in winter. The 

most pronounced changes seemed to occur at 42 m. 

Changes in composition and abundance of species 

during a renewal of the bottom water indicated that 

mysids were influenced by presumed near-bottom 

currents, while amphipods and decapods were not. 

Comparisons of the present .amphipod fauna with 

the fauna in 1933-37 revealed significant 

differences. The fauna in the deep, basin was 

impoverished and the Lilljeborgidae, earlier 

represented by three species are most likely caused 

by eutrophication of the fjord. 

Cadte, G. C. (1986) Increased phytoplankton 

primary production in the Marsdiep area 

(Western Dutch Wadden Sea). Netherlands 

Journal of Sea Research 20:285-290. 

{phytoplankton; primary production; Dutch 

Wadden Sea; phosphate; eutrophication) 

Annual phytoplankton primary production in the 

Marsdiep tidal inlet increased from ca 150 g C m-2 

in the period 1964 to 1976, to ca 300 g C m-2 in 

198111982 and 1985. This increase is considerable, 

but comparable to that observed in the outer Ems 

estuary, eastern Wadden Sea, from 240 g C m-2 a-l 

in 197211973 to 400-500 g C m-2 a-l in 

197611980. Although the increase is most probably 

due to eutrophication, as illustrated by the regular 

increase in phosphate in the Marsdiep area since 

1950, it is difficult to explain why this affected 

primary production no earlier than the late 

seventies. Primary production has probably not 

increased in the more turbid inner parts of the 

Wadden Sea, where light is the limiting factor, and 

P values were already higher than in the inlet areas. 

CadCe, G. C. (1986) Recurrent and changing 

seasonal patterns in phytoplankton of the 

westernmost inlet of the Dutch Wadden 

Sea from 1969 to 1985. Marine Biology 

93:28 1-289. 

{phytoplankton; abundance; Dutch Wadden Sea; 

distoms; flagellates; turbidity) 

Data for phytoplankton composition and 

abundance in the Marsdiep are presented for the 

period from 1969 to 1985 inclusive. Only a few 

species dominated the phytoplankton. A recurrent 

pattern was observed in the seasonal succession: in 

winter, total cell numbers were invariably low, but 

freshwater algae, sluiced into the Wadden Sea from 

IJssel Lake, showed highest densities in winter. A 

diatom spring peak was observed around mid-April, 

followed by a Phaeocystis pouchetii peak about 
three weeks later. Later in summer usually two 

more diatom peaks followed by non-diatom peaks 

were present. The exact timing of the spring peak 

varied from year to year, with the extremes being 

late March and early May. A relatively late spring 

peak usually coincided with a relatively high 

turbidity in the preceding winter. An increase in 

total cell numbers was found over the 17-year 

observation period. Diatoms decreased from 1969 to 

1974 but have increased since then, reaching values 

above those of 1969 during recent years. Flagellates 

showed a consistent increase over the entire 

observation period. 

Cadte, G. C. (1990) Increased bloom. Nature 

346:418. 

{phytoplankton; bloom; North Sea; Phaeocystis; 

nutrient) 

No abstract 

Cadte, G. C. (1992) Trends in Marsdiep 

phytoplankton. Proceedings of the 7th 

International Wadden Sea Symposium, 

{phytoplankton; Marsdiep; diatom; bloom; 

phosphorus; nitrogen; North Sea; primary 

production; composition; biomass) 

Phytoplankton cell-counts indicate an almost 

continuous increase in non-diatom plankton since 

the early 1970s. Particularly Phaeocystis blooms 

increased in duration. Diatoms fluctuated more 

irregularly. Man-induced eutrophication - increased 

in phosphate and nitrogen, but not in silicate - can 

explain such trends. They are comparable to those 

found in Helgoland, but different from (climateinduced) trends observed in the open North Sea 

during the Continuous Plankton Recorder survey. 

Such climatic effects are apparently overruled by 

eutrophication effects in coastal waters. Historical 

data on Phaeocystis for the Marsdiep area confirm 

the changes observed; they indicate blooming 

periods lasting some 50 days in 1897 and 1899, 

before eutrophication started. This blooming period 

has tripled in the late 1980s. 

Annual primary production was measured less 

regularly, but the first 14c production data were 

collected in 1963-1965. Primary production has 

remained c. 150 g carbon (C) m-2 a-l up to the 

early 1970s, but doubled in the early 1980s. 

Production in 1990 might be relatively low due to 

freshwater (including nutrient) discharge from the 

Usselmeer. 

Chlorophyll-a showed an increase parallel with 

the primary production increase in the late 1970s, 

but the annual variation is large. Comparison with 

the 1951-1953 data is hampered because of 

differences in the methods used. 

Cadte, G. C. and J. Hegeman (1991) Historical 

phytoplankton data of the Marsdiep. 

Hydrobiological Bulletin 24: 1 1 1- 1 18. 

{phytoplankton; trend; Marsdiep; Wadden Sea; 

Phaeocystis; bloom) 

Published and unpublished data on phytoplankton 

of the Marsdiep tidal inlet were studied. Most older 

data going back to 1897, are based in netphytoplankton only, the earliest quantitative 

(Utermohl) data being from 1965. Phaeocystis sp. 

bloomed in the Marsdiep after a spring diatom 

peak, at least as long ago as 1897. Summer and 

autumn peaks of Phaeocystis sp., frequent now, 

were also observed in 1898 and 1899. The duration 

of the Phaeocystis blooms in 1897 to 1899 was 

shorter than observed after 1978, but longer than 

in the early 1970s. The recent (1987 to 1989) 

duration of Phaeocystis blooms is 2 to 3 times that 

of 1897- 1899. The increase surpasses normal yearly 

variation and can be related to anthropogenically 

caused increase in nutrient concentrations. A 

number of diatom species, at present numerically 

dominant in the spring peak, are not mentioned as 

dominant in the earlier periods of observation. They 

are small and passed through the nets used. 

Biddulphia sinensis, at present often abundant, is an 

immigrant in the North Sea since 1903, and for 

that reason absent from the earliest Marsdiep 

observations. No clear trend in duration of diatom 

blooms is apparent during 1965 to 1989. 

Anthropogenic eutrophication did not affect diatom 

blooms. Marsdiep records in the literature of 

Phaeocystis globosa, P. pouchetii and P. sp. all 

refer to the same species. 

Caldwell, J. W. (1985) Effects of elevated 

turbidity and nutrients on the net 

production of a tropical seagrass 

community. Dissertation for Ph.D., The 

University of Florida. 

{turbidity; nutrients; seagrass; dredging; 

production; shading) 

Dredging effects on seagrass communities in the 

Florida Keys were examined by (1) comparing 

impacts on net production resulting from dredging 

and natural weather events, (2) determining changes 

in community photosynthetic efficiency, (3) 

evaluating shading and nutrient effects on net 

production, and (4) developing a systems dynamics 

model. 

Net community production was estimated during 

numerous meteorological and dredging events using 

the Odum-Hoskins oxygen technique in flowthough field microcosms. In other experiments, 

shading and nutrients (phosphorus, nitrate, and 

ammonia) were manipulated to simulate dredge 

plume conditions. The model examined the 

relationships between seagrass biomass, water 

column and sediment nutrients, detritus, and 

consumers. 

The greatest depression in net community 

production resulted from severe thunderstorms and 

dredging events, respectively. Net community 
production measured two years after dredging 

showed an approximate four-fold decreased. 

In field microcosm experiments, significant 

interaction occurred between shading and nutrient 

concentration. Significant metabolic reduction 

occurred due to shading , even with higher nutrient 

conditions. All but the lowest concentration 

resulted in significantly increased production in the 

light. Qualitative comparison with a control 

showed an enhancement of production only in the 

light. 

The model of seagrass production was most 

sensitive to changes in nutrient-seagrass 

relationships, seagrass production estimates, and 

seagrass-light interactions. Recovery of seagrass 

biomass following numerous dredging events (3.5 

years) was longer than that from estimated total 

annual thunderstorms encountered (1 year) but 

shorter than recovery from hurricane events (4.1 

years). The effects of short-term dredging on net 

community production as shown by field 

experimentation and the model were less severe 

than some weather events because of differences in 

duration and intensity. Lowered post-dredging 

photosynthetic efficiency, as defined by unit 

biomass production per unit light, was a result of 

degraded system function possibly from sediment 

scour from the dredging cut. The major deleterious 

effect of dredging resulted from shading, although 

some production enhancement could occur from 

nutrient release. The model suggested that dredging 

effects were prolonged because of current scour 

from the dredging cut, that recovery time was 

comparable to hurricane events, and that more 

investigation is needed in seagrass-nutrient and 

light-production relationships. 

Cambridge, M. L. and A. J. McComb (1984) 

The loss of seagrass in Cockburn Sound, 

Western Australia. I. The time, course 

and magnitude of seagrass decline in 

relation to industrial development. 

Aquatic Botany 20:229-243. 

{seagrass; effluents; Australia) 

The areas of seagrass meadows in Cockburn 

Sound, a marine embayment in Western Australia, 

were estimated from historical aerial photographs 

supplemented by ground surveys, studies on 

meadows in adjoining areas, and coring for rhizome 

remains. Ten species of seagrass with different 

habitat tolerances are recorded for the area, with 

Posidonia sinuos Cambridge et Kuo forming the 

most extensive meadows. It is estimated that from 

1954 to 1978 the meadow area was reduced from 

some 4200 to 900 ha., based on leaf detritus 

production from 23000 to 4000 t (dry wt.) y-l. The 

major loss of seagrass occurred during a period of 

industrial development on the shore, and the 

discharge of effluents $ch in plant nutrients. 

Carpenter, E. J. (1970) Effects of phosphorus 

mining wastes on microorganisms of the 

Palmico River estuary, North Carolina. 

Dissertation for Ph.D., North Carolina State 

University at Raleigh. 

{Palmico River estuary; blue-green algae; sewage; 

phosphorus; mining; photosynthesis) 

To test the effects of phosphorus mining wastes 

(SPW) and domestic sewage on microorganisms of 

the Palmico River estuary, six 1400 m3 artificial 

estuaries and eight 15.1 m3 static pools were 

constructed. Concentration, excluding controls, of 

SPW in the pools was 1 percent in estuaries ranged 

from 1 percent to 51 percent of the total water 

volume. The present day concentration of SPW in 

the lower Palmico River is 0.7 percent (vlv) and is 

expected to rise as other companies begin mining 

operations. Three experiments were carried out, two 

(54 and 44 days long) in the estuaries and one (15 

days long) in the plastic pools. 

The experiments showed that the addition of SPW 

to estuarine water does not immediately increase the 

photosynthesis or biomass of phytoplankton. This 

is most likely because phosphorus does not limit 

the growth of phytoplankton in the river. As 

shown in a nutrient limitation study, nitrogen 

limits algal photosynthesis in the Palmico estuary. 

However, blue-green algae cell numbers (Anabaena 

sp. and Spirulina sp.) were 77.2 percent higher 

(.010<p<.025) than controls in estuaries containing 

approximately 13 percent SPW. Although cell 

numbers were 33.2 percent above controls in 

estuaries with 3 percent SPW, the difference was 

not significant at the 5 percent level. Also, bluegreen algae (Anabaena torulosa) numbers were 404 

percent higher (.Ol<p<.025) than controls in 

plastic pools with 1 percent SPW (vlv) and 295 

percent above (.010<p<.025) controls with 1 

percent SPW and 0.1 percent (vlv) artificial sewage. 

The growth of blue-green algae was probably 

stimulated by the addition of phosphorus in the 

SPW and their nitrogen fixing ability allowed then 

to obtain sufficient nitrogen in waters where this 

nutrient is limiting. This increase in blue-green 

algae may be potentially harmful to the estuary by 

changing the algal species composition and by 

adding fixed nitrogen to an estuary where 

photosynthesis is nitrogen limited: 

An interaction occurred in the pools between 

SPW and the artificial sewage mix which resulted 

in photosynthesis 28.8 percent above the controls. 

The existence of this interaction shows the 

importance of sewage treatment facilities for towns 

in the Palmico River basin. Rate of phosphorus 

uptake by microorganisms was 23.1 percent higher 

(p<0.005) than controls in plastic pools with 1 

percent SPW and 74.9 percent higher (pe.005) than 

controls in pools with both SPW and artificial 

sewage. This indicated that microorganisms are 
taking up the added phosphorus in SPW. The 

uptake in pools with both SPW and sewage was 

3.2 x that in the pools with SPW added alone. This 

uptake may reflect the primary productivity 

interaction which occurred in the pools. No changes 

in the Vmax of glucose by bacteria was seen in the 

studies. This was probably because the rate of 

formation of glucose did not increase in an 

experiment of such short duration. 

Carpenter, E. J. and D. G. Capone (ed.) (1983) 

Nitrogen in the Marine Environment. 

Academic Press, New York. 

No abstract 

Caspers, H. (1987) Changes in the benthos 

at a sewage-sludge dumpsite in the Elbe 

Estuary. In: J. M. Capuzzo and D. R. Kester 

(ed.) Oceanic Processes in Marine Pollution. Vol. I. 

Biological Processes and Wastes in the Ocean. 

Robert E. Krieger Publishing Company, Malabar, 

Florida. 

{sewage; fauna; settlement; stability) 

Twenty years of investigations on the benthos in 

the outer part of the Elbe Estuary, which includes a 

sewage-sludge dumpsite, show that the community 

structure was highly unstable. Some changes were 

brought about by hydrological factors and the 

variable pattern of sedimentation. Most of the 

changes, however, resulted primarily from 

biological factors, such as the predominance of one 

or two species, the availability of physical space, 

predation, and the life span of the species. The 

continual arrival of larvae, belonging to various 

taxa, caused a more rapid fluctuation in the benthic 

community structure than is usually encountered in 

freshwater habitats. Communities in the coastal 

marine area of the Elbe Estuary might best be 

designated as astatic and characterized as unstable 

but highly resilient. 

Cederwall, H. and R. Elmgren (1980) Biomass 

increase of benthic macrofauna 

demonstrates eutrophication of the Baltic 

Sea. Ophelia, Supplementum 1:287-304. 

{abundance; macrofauna; benthic; Baltic) 

Twenty-eight stations around the Baltic islands of 

0land and Gotland, sampled by Hessle in 1920 and 

1923 were revisited in 1976-1977. Using methods 

similar to those of Hessle, we demonstrate a 

statistically highly significant increase in 

macrofauna biomass above the halocline (median 

over 4 times higher, based on 23 stations) over the 

intervening 55 years. Neither differences in 

methods, nor interannual fluctuations can explain 

the large increase, which is considered to represent a 

real long-term change, apparent over the entire 

Baltic Sea. The only plausible cause is 

eutrophication of the Baltic, leading to higher food 

supplies for the benthic fauna. These results 

constitute the first statistically confirmed biological 

evidence of this eutrophication, and demonstrate 

that eutrophication is a major process affecting the 

entire Baltic Sea, and not just restricted coastal 

areas. 

Cederwall, H. and R. Elmgren (1990) 

Biological effects of eutrophication in 

the Baltic Sea, particularly the coastal 

zone. Ambio 19: 109-1 12. 

{Baltic; review) 

The reported biological effects of the increased 

nutrient load on the Baltic Sea are summarized, 

with some comparisons with the Kattegat and 

Skagerrak. Interest is focused on the coastal zone, 

where effects are more obvious than in offshore 

areas, but from which results have not often been 

published internationally. Reports demonstrate 

environmental degradation over extensive coastal 

areas of the Baltic Sea. Recorded effects include 

increased nutrient levels; increased algal blooms, 

chlorophyll a concentrations, and primary 

productivity; decreased water transparency and 

decreased depth penetration of Fucus vesculosus; 

increased deposition of organic matter on the 

bottom and increased frequency and severity of 

oxygen deficency in bottom waters; and reduction 

of bottom fauna. It is concluded that for most 

Baltic countries efforts to reduce local 

eutrophication are likely to have important positive 

effects, even when reductions in discharges of 

nutrients are relatively insignificant in comparison 

to the total nutrient load on the Baltic Sea. 

Cerco, C. F. (1981) Nitrification in the 

upper tidal James River. In: B. J. Neilson 

and C. E. Cronin (ed.) Estuaries and Nutrients. 

Humana Press, Clifton, New Jersey. 

{bacteria; nitrification; ammonia; model) 

A field and model study of the nitrification 

process in the upper tidal portion of the James 

River, Virginia, has been completed. Attention was 

devoted to the enumeration of nitrifying bacteria 

and to the determination of the fate of constituents 

involved in the nitrification process. Ammonia and 

nitrite oxidizers are present in the James River 

water column in concentrations of 10-I - 10 

mpmlml and in the bottom sediments in 

concentrations of lo2 - lo5 mpmlml. Elevated 

populations are observed in the vicinity of waste 

discharges suggesting that nitrification of these 

wastes begins immediately upon discharge. 

Examination of three sets of field data supports the 

hypotheses that nitrification occurs. Of particular 

interest is the observation, during two instances, of 

nitrate removal concurrent with nitrification. 

Cheney, D. P. (1992) Project development to 

harvest the beach-fouling alga Pilayella 

littoralis in Nahant Bay, MA. 31st Annual 
Symposium of the Northeast Algal Society, Woods 

Hole. 

{seaweeds; beach; fouling; Pilayella; Nahant Bay) 

The beaches of Nahant Bay, located on the north 

shore of Massachusetts, have been perturbated for 

many years by an unusual, presumably mutant, 

free-living form of the brown alga Pilayella 

littoralis. Normally, P. littoralis grows as a small 

filamentous plant attached to larger algae in the 

littoral zone during late winter and spring months. 

However, ever since at least 1903, large masses of 

free-living, perennial form of P. littoralis have been 

fouling Nahant Bay beaches, often accumulating at 

low tide in amounts of hundreds of tons. In recent 

years, there have been sightings of large casts of P. 

littoralis on beaches south of Nahant Bay, raising 

concerns that the distribution of the free-living 

form may be spreading. Recently, my laboratory in 

collaboration with a fisheries engineer at MIT has 

embarked upon a project aimed at developing the 

necessary harvesting equipment and strategy for 

effecting a solution to Nahant's beach-fouling 

problem. This project will examine the problem 

from a population biology as well as engineering 

viewpoint. The current distribution, quantity and in 

situ growth rate of Pilayella will be evaluated, in 

order to determine if harvesting will prove 

economically-practical. Molecular techniques will 

be used to determine the genetic relationship 

between presumed mutant and attached forms. A 

prototype harvesting system has already been 

developed and is being evaluated. 

Cho, C.-H. (1991) Mariculture and 

eutrophication in Jinhae Bay, Korea. 

Marine Pollution Bulletin 23:275-279. 

{red tide; mortality; diatom; flagellates) 

The contents of organic matters and sulfide in the 

superficial bottom muds, and mass mortality of 

shellfish mainly due to red tides in the Jinhae Bay 

are reviewed. COD, ignition loss, and sulfide 

contents increased for the last decade from 1972 to 

1982. Causative organisms of red tides also 

changed from diatom to flagellates. A decrease of 

shellfish production or mass mortalities of this area 

would be caused by a high level of eutrophication, 

which resulted from both waste discharge from an 

industrial complex and organic deposits from the 

aquaculture. 

Colombo, G., I. Ferrari, V. U. Ceccherelli, and R. 

Rossi (ed.) (1992) Marine Eutrophication 

and Population Dynamics. Olsen & Olsen, 

Fredensborg. 

Contains eighteen papers on various aspects of 

eutrophication. 

Conolly, N. J. and E. A. Drew (1985) 

Physiology of Laminaria. 111. Effects of 

a coastal eutrophication gradient on 

seasonal patterns of growth and tissue 

composition in L. digitata Lamour and L. 

saccharina (L.) Lamour. Marine Ecology 

6:181-195. 

{seaweeds; growth; nutrient; Laminaria) 

Dissolved nitrate, nitrite, ammonium and 

phosphate were monitored for 2.5 years along a 

coastal eutrophication gradient originating at the 

St. Andrews sewer outfall. Linear growth rates and 

tissue composition (phosphorus, various nitrogen 

fractions and storage carbohydrates) were monitored 

at the same stations in lamina tissue of Laminaria 

digitata and L. saccharina. Growth rates were 

considerably enhanced at the eutrophicated stations 

both in spring, when exogenous nutrients were at 

peak values at all stations, and during the summer 

when exogenous nutrients were very low at all 

stations. Enhanced summer growth rates were 

correlated with the increased reserves of N and P 

accumulated during winter and spring, and 

particularly with soluble organic nitrogen reserves. 

Accumulation of storage carbohydrates was 

inversely correlated with growth rate and tissue N 

and P reserves, presumably because fixed carbon 

could be incorporated into new protein and thence 

new tissue only if internal non-protein N reserves 

were available. 

Copeland, B. J. and D. E. Washlschlag (1968) 

Biological responses to nutrients - 

eutrophication: Saline water 

considerations. Advances in water quality 

improvement. Water Resources Symposium, 

Austin, Texas. 

{ecosystem; nutrient; production; zooplankton; 

community; fish) 

1) The addition of a known nutrient source to a 

marine community resulted in alteration of the 

metabolic patterns of the ecosystem. An increase in 

the amount of nutrient material caused an increase 

in photosynthetic production. 2) Species diversity 

of zooplankton is reduced in marine environments 

receiving various types of organic wastes, which 

indicates an interruption of normal community 

structure by the addition of new nutrient materials. 

3) A theoretical case was presented to demonstrate 

that slight toxic effects are pronounced although 

not detectable by conventional means of 

measurement. With just a slight increase in 

mortality rate the biomass of the fishes affected 

would be decreased greatly. 4) Experiments on the 

metabolic rate of fishes revealed that slight 

pollution stresses tend to lower the metabolic rates 

considerably. The depression was greatest when the 

fish were already subjected to regular environmental 

stress, such as low temperature. 5) Most of the 

effects of waste materials in the marine 

environment are subtle. It appears that the 

community approach rather than the organismic 

approach will be more fruitful in evaluating the 
impact of man-made changes in inland saline water 

ecosystems, notwithstanding the fact that the 

effects of stresses on individual populations can be 

physically quantifiable. 

Correll, D. L. (1981) Eutrophication trends 

in the water quality of the Rhode River 

(1971-1978). In: B. J. Neilson and L. E. 

Cronin (ed.) Estuaries and Nutrients. Humana 

Press, Clifton, New Jersey. 

{nutrient; phosphorus) 

Five to eight-year data sets on turbidity and a 

series of nutrient parameters have been taken in the 

Rhode River, a small tidal river tributary to 

Chesapeake Bay. Trends of change at headwater 

stations indicate changes due primarily to local 

watershed runoff, while changes at stations near the 

river's mouth are indicative of changes in 

Chesapeake Bay. 

The data have been analyzed first by summarizing 

monthly and seasonal means and variances, then by 

looking for year-to-year trends. Thus, for example, 

linear regressions of summer and fall turbidity 

versus time for 1971-1978 had very low slopes and 

low coefficients of determination. An interesting 

finding was the pattern or total phosphorus in 

surface waters. At the mouth of Rhode River, 

seasonal mean concentrations increased steadily for 

each season each year from the fall of 1971 to the 

fall of 1976. The most dramatic increases (fourfold) were observed in the summer and fall. A much 

smaller increase occurred in winter and spring 

values. In contrast, year-to-year concentrations of 

total phosphorus in surface waters in upstream 

stations over the same time period showed less 

clear-cut trends. Also, phosphorus loadings from 

local watershed runoff fluctuated widely, but had no 

steady rapid rise with time. Although total 

phosphorus increased dramatically at the mouth of 

Rhode River, dissolved orthophosphate, nitrate, and 

dissolved ammonia remained essentially constant, 

especially in the summer and fall. These data could 

indicate an increasing impact of summertime 

anoxic bottom waters on the phosphorus dynamics 

of the upper western shore of the Bay. 

Cosper, E. M., V. M. Bricelj, and E. J. Carpenter 

(ed.) (1989) Novel Phytoplankton Blooms: 

Causes and Impacts of Recurrent Brown 

Tides and other Unusual Blooms (Papers 

from a conference at Stony Brook, NY, 

27-28 October, 1988). Springer-Verlag, 

Berlin. 

No abstract 

Cosper, E. M., W. Dennison, A. Milligan, E. J. 

Carpenter, C. Lee, J. Holzapfel, and L. Milanese 

(1989) An examination of the 

environmental factors important to 

initiating and sustaining "brown tide" 

blooms. In: E. M. Cosper, V. M. Bricelj and 

E. J. Carpenter (ed.) Novel Phytoplankton Blooms, 

Causes and Impacts of Recurrent Brown Tides and 

other UnusualBlooms. Springer-Verlag, Berlin. 

{brown tide; bloom; Narragansett Bay; Long 

Island) 

No abstract 

Cosper, E. M., C. Lee, and E. J. Carpenter (1990) 

Novel "brown tide" blooms in Long 

Island embayments: A search for the 

causes. In: E. GranBli, B. Sunstrom, L. Edler 

and D. M. Anderson (ed.) Toxic Marine 

Phytoplankton. Elsevier, New York. 

{Aurococcus; brown tide; bloom; phytoplankton; 

Long Island; organic; micronutrients; nutrients; 

precipitation; heterotrophic uptake) 

Unusual blooms of a previously unidentified 

chrysophyte, Aureococcus anophaefferens; have 

occurred in several coastal embayments along the 

northeast coast of the USA. The monospecific 

blooms were termed the "brown tide" due to the 

resulting water color. The first appearance of the 

"brown tide" occurred early in the summer of 1985 

over a wide geographic range in non-contiguous 

bodies of water. The bloom did not return to some 

bays but, on Long Island, New York they recurred 

during the summer of 1986 and in diminishing 

densities during the summers of 1987 and 1988. 

Historically, a diverse group of small microalgal 

species dominate the phytoplankton biomass and 

productivity in Long Island bays during the 

summer. The continued dominance through several 

months at high cell densities (> lo9 cells 1-l) of A. 

anophaefferens was the distinctive feature of 

consequence during these blooms. 

Environmental variables which may contribute to 

the occurrence of the "brown tide" include elevated 

salinities due to drought conditions, pulses or 

rainfall delivering organic and/or micronutrients to 

bay waters, reduced grazing and restricted flushing 

of bays. The "brown tide" species appears to be 

closely related to an open ocean chrysophyte, 

Pelagococcus subviridis, and possibly was seeded 

into northeast coastal bays from offshore when 

conditions during 1985 were particularly favorable 

for its growth. The ability of this species to 

maintain at least minimal populations during the 

winter months seems to allow for its recurrence 

during subsequent summers. Culture studies have 

shown thatt his species has growth requirements for 

trace elements, chelators and organic nutrients some 

of which are different from many common estuarine 

and coastal phytoplankton species. The competitive 

advantage of A. anophaefferens over other 

potentially co-occurring species probably related to 

its heterotrophic and photoadaptive capabilities. 

Costa, J. E. (1988) Eelgrass (Zostera marina 

L.) in Buzzards Bay: Distribution, 
production, and historical changes in 

abundance. Dissertation for Ph.D., Boston 

University. 

{eelgrass; periphyton; abundance; growth; 

recolonization; Buzzard Bay) 

The past and present-day distribution of eelgrass 

(Zostera marina L.) was documented using aerial 

photographs, field surveys, nautical charts, 

sediment cores, and first-had accounts. Eelgrass 

growth correlates with local temperature and 

insolation, and annual production is =350 g C m2yr-1. In Buzzards Bay, eelgrass beds cover 41 

km2 of substrate and account for 12% of primary 

production; in shallow bays, eelgrass equals 40% of 

production. 

Prior to the "wasting disease" of 1931-32, 

eelgrass populations equaled or exceeded present-day 

abundance. Six to 10 years after the disease, 

eelgrass covered less than 10% of the present-day 

habitat area. The process of recolonization was 

similar in many areas: new beds initially appeared 

on bare substrates, beds expanded, new beds 

appeared, and some beds were removed by 

disturbance. A computer simulation modeled these 

events, and showed that rapid recolonization of 

eelgrass populations is highly dependent on new 

bed recruitment, which in nature depends on seed 

dispersal. High disturbance rates slow eelgrass 

colonization and lower peak cover. 

Local changes in eelgrass abundance are driven by 

anthropogenic and natural disturbances which are 

superimposed on the regional pattern of 

catastrophic decline and gradual recovery. 

Hurricanes, ice scour, and freezing periodically 

destroyed eelgrass beds in some areas. Eelgrass 

populations in poorly flushed, developed bays, with 

declining water quality, never recovered from the 

wasting disease or showed new declines in recent 

years. 

The distribution of eelgrass is light limited, and 

eelgrass beds may disappear in enriched areas 

because of increases in algal epiphytes and 

phytoplankton. To identify what levels of nutrient 

loading cause these changes, concentrations and 

inputs of dissolved inorganic nitrogen (DIN) in 

Buttermilk Bay were measured. Periphyton on 

eelgrass leaves and plastic screen strips on floats 

correlated well to mean DIN. Experimental floats 

released nutrients and demonstrated that small 

increases in DIN significantly increased periphyton 

abundance. The depth of eelgrass growth in 

Buttermilk Bay decreased by 9 cm for every 1 uM 

increase in DIN. Periphyton abundance is more 

important than phytoplankton concentrations in 

limiting eelgrass growth in Buttermilk Bay, 

because water in this bay has short residence time, 

and phytoplankton gradients are less important. 

Crema, R., A. Castelli, and D. Prevedelli (1991) 

Long term eutrophication effects on 

macrofaunal communities in northern 

Adriatic Sea. Marine Pollution Bulletin 

22~503-508. 

{Adriatic Sea; macrobenthos; Po River; 

community structure; dystrophic; Italy) 

The macrozoobenthic community in the northern 

Adriatic Sea, south of the Po river, along the 

Emilia-Romagna region coast, was sampled in 

1985. Sampling site was central to a highly 

eutrophicated area with greatly increased intensity 

and frequency of dystrophic events over recent 

decades. The sampled community differs from all 

those described in the same area in a period (1934- 

1936) previous to the actual degree of 

eutrophication. Large abundances of species 

indicative of unstable bottoms, such as the bivalve 

Corbula gibba and the polychaete Lumbrineris 

latreilli were recorded. Moreover, the community 

structural features indicate a state of immaturity, 

such as in early successional stage communities. 

The increased frequency of acute dytrophic events 

and consequent shortening of the time between 

successive disturbances is proposed as the cause of 

bicenosis modification and its current structure and 

composition. 

Crowder, A. and D. S. Painter (1991) 

Submerged macrophytes in Lake Ontario: 

current knowledge, importance, threats 

to stability, and needed studies. Canadian 

Journal of Fisheries and Aquatic Science 48: 1539- 

1545. 

{submerged vegetation; stability; shading; Lake 

Ontario) 

The submerged limnetic community in Lake 

Ontario includes algae, bryophytes, and about 30 

species of angiosperms. Their distribution is 

accurately known in some areas but not lake-wide, 

and a whole-lake survey is recommended. In 

nutrient cycling, submerged vegetation acts as a 

sinks during the summer; metals and metalloids 

occur in high concentrations in tissues from some 

areas. Known herbivores include invertebrates, fish, 

and waterfowl. Stands are necessary for many fish 

taxa as breeding or nursery habitats, and for 

waterfowl, but may be damaged by carp (Cyprinus 

carpio). Stability has been affected by water levels, 

sedimentation, wave and ice movement, invasive 

species, herbivory, eutrophication and turbidity, and 

contaminants. Recovery after control of P loading 

has occurred in Irondequoit Bay but is delayed by 

turbidity in the Bay of Quinte. 

Cruz, A. A. d. I., C. T. Hackney, and J.P.Stout 

( 19 8 1 ) Aboveground net primary 

productivity of three Gulf Coast marsh 

macrophytes in artificially fertilized 

plots. In: B. J. Neilson and L. E. Cronin (ed.) 

Estuaries and Nutrients. Humana Press, Clifton, 

New Jersey. 
{nitrogen; enrichment; tidal marsh; macrophytes) 

Plots (100 m2) of four tidal marsh communities 

(Juncus roernerianus and Spartina alterniflora in 

Alabama, J. roernerianus and Spartina cynosuroides 

in Mississippi) common in the Gulf Coast were 

enriched with commercial NH4N03 (23 percent N). 

The fertilizer was applied once at the beginning of 

1978 growing season to stimulate a farm-plantation 

operation at a dosage (136 g/m2) estimated to 

return to the soil approximately the same amount 

of nitrogen contained in the plants. Six 0.25 m2 

quadrants were harvested monthly from each 

community from April to November. The annual 

net productivity was estimated with a maximum 

minus minimum standing crop technique based on 

a predictive periodic model (PPM). A correction for 

plant mortality during the sampling period is 

provided in the PPM technique. Annual 

aboveground net primary productivity increased by 

59 percent in the Alabama J. roernerianus, 84 

percent in the Mississippi J. roernerianus, 82 

percent in the S. alterniflora and 26 percent in the 

S. cynosuroides. It appears that short form or high 

marsh macrophytes responded more to nitrogen 

enrichment than tall form or low marsh plant. 

Cruz-Kaegi, M. E. and G. T. Rowe (1992) 

Benthic biomass gradients on the TexasLouisiana shelf. In: (ed.) Nutrient Enhanced 

Coastal Ocean Productivity, NECOP Workshop 

Proceedings, October 1991. Texas A&M 

University Sea Grant Program, TAMU-SG-92- 109, 

{benthic abundance) 

Macrofauna and bacterial biomass were studied in 

relation to depth, distance from the Mississippi 

River, primary productivity, and concentration of 

organic carbon in sediments. Preliminary results 

suggest that combined benthic biomass was 

relatively low in the Gulf of Mexico compared to 

that off New England. Low benthic biomass is 

thought to be a function of sediment load, 

unpredictable hypoxic events and trawling pressure, 

counteracting the positive effects of high primary 

productivity, fine grain size and relatively high 

concentrations of organic matter in the sediments. 

Darnell, R. M. and T. M. Soniat (1981) 

Nutrient enrichment and estuarine 

health. In: B. J. Neilson and L. E. Cronin (ed.) 

Estuaries and Nutrients. Hamana Press, Clifton, 

New Jersey. 

{nutrient; estuary; cycle; ecosystem health) 

Ecosystem health may be defined in terms of 

system norms or in terms of human utility. It is 

here defined as that state in which the components 

and processes remain well within specified limits of 

system integrity selected to assure that there is no 

diminution in the capacity of the system to render 

its basic services to society throughout the 

indefinite future. Knowledge of nutrient enrichment 

in freshwater systems is reviewed as a point of 

departure for understanding the response of estuaries 

to enrichment. Mixed and stratified estuaries 

respond in somewhat different fashions. Major 

changes associated with enrichment are changes in 

species succession and oxygen depletion in areas of 

organic accumulation, low mixing, and poor 

flushing. If of temporary and local chronic nature or 

of widespread occurrence, they could lead to 

irreversible loss of species and genetic stocks. 

Measures of ecosystem health are discussed, and 

pertinent management recommendations are put 

forth. Of especial importance are the needs to 

establish local species reserves, to manage total 

system integrity, and to develop better lines of 

communication between scientists and managers. 

Dauer, D. M. and W. G. Conner (1980) Effects 

of moderate sewage input on benthic 

polychaete populations. Estuarine and 

Coastal Marine Science 10:335-346. 

{benthic; sewage; polychaete; abundance; biomass; 

productivity) 

The polychaete fauna of a sandy intertidal habitat 

receiving effluent from a sewage treatment plant 

was compared to that of a control site. The total 

number of individuals, total biomass, and average 

species numbers were significantly greater at the 

sewage-affected site. The response of individual 

species to nutrient enrichment varied. Some species 

showed significantly greater numbers of individuals 

at the sewage-affected site while other species' 

densities showed no difference. All species 

maintained greater biomass at the enriched site, 

most biomass differences were significant. Those 

species which did not show density differences 

between sites tended to have the greater difference 

in average weight per individual. In general, species 

with either benthic development or some form of 

brooding behavior were best able to exploit the 

enriched condition by increased densities, or greater 

average biomass per individual. It was concluded 

that for moderate increasee in nutrient 

concentrations, benthic productivity will be most 

enhanced in those sediments with very low 

concentrations of fine sized particles. 

Dauvin, J. C. and F. Gentil (1989) Long-term 

changes in populations of subtidal 

bivalves (Abra alba and A. prismatica) 

from the Bay of Morlax (Western English 

Channel). Marine Biology 103:63-73. 

{Abra; bivalve; organic; population; reproduction; 

growth; abundance; life history) 

Two populations of Abra alba (Wood) and one of 

A. prisrnatica (Montagu) (Molluscs: Bivalvia) were 

studied over a 10 year period (1977-1987) in two 

muddy fine-sand subtidal communities of the Bay 

of Morlax, France. The survey provided an example 
of long-term changes in the three Abra spp. 

populations, which displayed synchronized changes, 

with a regular annual cycle and increasing densities 

during 1979-1980 related to the higher 

concentration of organic matter resulting from the 

"Amoco Cadiz" oil spill in March 1978. A. alba 

rapidly adapts its demographic strategy to eutrophic 

conditions by increasing its reproductive potential, 

growth, and abundance. During times of 

eutrophication, A. alba has three spawning periods 

and two recruitments per year as opposed to two 

spawning periods and two recruitments per year 

during oligotrophic conditions. Growth of the 

juveniles of this species is insignificant until 

spring for individuals recruited in the autumn, 

whereas individuals which settle during spring or 

summer display immediate rapid growth. A. 

prismatica has a low capacity to adapt to eutrophic 

conditions. It has one annual period of sexual 

maturation at the end of the summer, with 

spawning in September-October and settlement 

beginning in mid-November. Growth of the 

juveniles after settlement is also insignificant until 

April. These results enable comparison of the 

demographic strategies of these two sympatric 

species. 

DeBoer, J. A., H. J. Guigli, T. L. Israel, and C. F. 

D'Elia (1978) Nutritional studies of two red 

algae. I. Growth rate as a function of 

nitrogen source and concentration. Journal 

of Phycology 14:261-266. 

{red algae; growth; nitrogen; culture) 

Gracilaria foliifera (Forsskal) Borgesen and 

Neoagardhiella baileyi (Harvey ex Kiitzing) Wynne 

& Taylor were grown in continuous-flow culture 

under controlled environmental conditions in 15 

liter experimental chambers. Growth rate was 

related to the source and concentration of nitrogen 

enrichment supplied to the plants. Growth rate 

appeared to follow saturation-type nutrient uptake 

kinetics for plants receiving ammonium, nitrate, 

urea or sewage effluent enrichments. Ammonium 

enrichment produced higher growth rates than 

nitrate or sewage enrichment. The lowest growth 

rates occurred in the chambers receiving unenriched 

seawater or urea. Half saturation controls (K) for 

growth were in the range of 0.2 - 0.4 uM N for all 

N enrichments examined. The low estimated values 

of K compared closely with those found for 

microalgae and indicate that both species posses the 

ability to utilize very low concentrations of N. 

Degobbis, D. (1989) I n c re a s e d 

eutrophication of the Northern Adriatic 

Sea. Marine Pollution Bulletin 20:452-457. 

{phytoplankton; Adriatic Sea; oxygen; mucous 

aggregates) 

Very calm and warm weather in late spring and 

early summer 1988 supported intense non-seasonal 

phytoplankton blooms in the western part of the 

northern Adriatic which is under direct influence of 

Po River discharge. Surface salinity decreased to 16 

X and concentration of chlorophyll a 

increased up to 17 mg m-3, with oxygen 

supersaturation up to 17 203 X 

Nanoplankton dominated the blooms (up to 76 X 

lo9 cells m-3, but diatoms, particularly 

Chaetoceros spp. (up to 17 X lo9 cells m-3), also 

occurred in large numbers. Large 'organic' 

aggregates were formed in the water column with 

dimensions (up to 2 m long) and in quantities never 

previously observed. During late summer and early 

fall aggregates were distributed over the entire 

region, and a general contamination of the coasts 

occurred. Oxygen concentration was dramatically 

reduced in the bottom layer of the entire region, an 

event not observed to such a degree since 1977. 

During that year, in contrast to 1988, large 

quantities of freshwater spread over the entire 

northern Adriatic causing extended phytoplankton 

blooms. In 1988 extremely low horizontal 

advection (current velocities usually below 20 cm 

s-l) primarily favoured an accumulation and high 

aggregation of detritus, and greatly reduced oxygen 

concentration in the bottom layer. 

DeGroodt, E. G., H. F. J. Los, T. A. Nauta, A. A. 

Markus, and I. deVries (1992) Modelling 

cause-effect relationships in 

eutrophication of marine systems: an 

integral approach. In: G. Colombo, I. 

Ferrari, V. U. Ceccherelli and R. Rossi (ed.) 

Marine Eutrophication and Population Dynamics. 

Olsen & Olsen, Fredensborg. 

{modelling; nutrients; phytoplankton species 

composition; North Sea) 

There is evidence that the elevated anthropogenic 

nutrient input into the North Sea is related to 

observed changes in primary production, 

phytoplankton biomass as well as species 

composition. Mathematical modelling techniques 

were used in order to analyze the influence of these 

inputs. The overall partitioning of gaines and 

losses has been simulated with a dynamic global 

model. The model simulates the spatial and 

temporal concentrations of nutrients and 

phytoplankton in the entire North Sea. 

A general, more detailed ecological model has 

been developed for the representation of vertical 

inhomogeneities of nutrients and phytoplankton 

species composition. Nutrient and phytoplankton 

levels were computed for different nutrient inputs 

and varying meterological conditions. The model 

results show an interdependence of phytoplankton 

species composition and nutrient input. However, 

there is little effect of nutrient inputs on the 

calculated chlorophyll-a concentrations at deep 

(stratified) areas. These results indicate that 
phytoplankton species composition may be more 

sensitive to changes in the nutrient inputs than the 

chlorophyll-a concentration. 

Dennis, L. and H. H. Seliger (ed.) (1979) Toxic 

Dinoflagellate Blooms (Proceedings of 

the Second International Conference on 

Toxic Dinoflagellate Blooms, Key 

Biscayne, Florida, 31 October - 5 

November, 1978). Elsevier, North Holland. 

No abstract 

Dethlefsen, V. and H. V. Westernhagen (1983) 

Oxygen deficiency and effects on bottom 

fauna in the eastern German Bight in 

1982. Meeresforschung 30:42-53. 

{benthos; fish; oxygen) 

In August and September 1982 two cruises were 

carried out to investigate the extent and effects of 

02-deficiencies in German and Danish coastal 

bottom waters. In two thirds of the more than 15 

000 km2 covered by the survey 02-content was 

less than 4 mln (60% saturation); lowest values 

were near 1 mln. 

In bottom waters with low 02-contents, pH 

values were generally low (7.7). During the first 

cruise water in the area investigated was stratified, a 

thermocline was between 10-15 m above the 

bottom. In 02-deficiency areas (02-saturation 

around 10%) fish catches were low containing dead 

Agonus cataphractus, Pleuronectes platessa and 

Limanda limanda. When operating underwater TV 

dead fish (Agonus cataphractus, Callionymus lara, 

Ammodytes sp. and flatfish) were detected lying 

dead on the bottom. 

Evaluation of underwater photography revealed the 

occurrence of dead benthic organisms in areas with 

low oxygen. The species were Ophiura albida and 

Venus stritula (paired empty valves). Possible 

causes for the occurrence of oxygen deficiencies are 

discussed and it is concluded that eutrophication 

processes in combinations have triggered low 

oxygen conditions in bottom waters. 

Doering, P. H., C. A. Oviatt, L. L. Beatty, V. F. 

Banzon, R. Rice, S. P. Kelly, B. K. Sullivan, and 

J. B. Frithsen (1989) Structure and function 

in a model coastal ecosystem: silicon, 

the benthos and eutrophication. Marine 

Ecology Progress Series 52:287-299. 

Lowering the N:Si ratio (14:l to 1:l) in nutrient 

(N, P, Si) additions to large (13 m3) outdoor 

mesocosms with and without an intact benthic 

community generally resulted in higher standing 

stocks of diatoms. An enriched diatom flora during 

the year-long experiment was not accompanied by a 

concomitant reduction in the extent of 

eutrophication caused by nutrient loading, as judged 

by oxygen concentration and metabolism or by 

chlorophyll a biomass. Enhanced growth of 

Atlantic menhaden Brevoortia tyrannus in the low 

N:Si ration treatment suggested that diatoms 

promote transfer of energy to higher trophic levels. 

Removal of the benthic community and associated 

sediments had a dramatic effect on pelagic 

community structure. Zooplankton were 

holoplanktonic in character and higher trophic 

levels were dominated by ctenophores, medusae, 

chaetognaths and fish. In the presence of an intact 

benthos the zooplankton had a larger 

meroplanktonic component and higher trophic 

levels were dominated by larvae of benthic adults 

(anemones, shrimp) or temporary planktonic adults 

(mysids). 

Doi, T. and A. Nitta (1991) Ecological 

modelling at Osaka Bay related to longterm eutrophication. Marine Pollution 

Bulletin 23:247-252. 

{modelling; biomass; fauna) 

Based upon the ecological model presented by 

Anderson & Ursin (1977), we constructed an 

adequate model which will quantitatively evaluate 

the overall impact of industrial large-scale 

development on fisheries at Osaka Bay. It is 

expected that the patterns of changes in biomass of 

dominant species, which caused by eutrophication, 

will be explained by the model we have established. 

The simulated results for two years, 1956-57, are as 

a whole comparable to the actual variations. 

However, the results for several species, including 

sea bass, flat fish, crab, mantis shrimp, and 

cuttlefish, differ significantly from observed data. It 

was attributed to the lack of accurate parameters of 

anabolism and catabolism for younger-aged stages 

of these species. 

Dortch, Q., D. Milstead, N. N. Rabalais, S. E. 

Lohrenz, D. G. Redalje, M. J. Dagg, M. E. Turner, 

and T. E. Whitledge (1992) Role of silicate 

availablilty in phytoplankton species 

composition and the fate of carbon. In: 

(ed.) Nutrient Enhanced Coastal Ocean 

Productivity, NECOP Workshop Proceedings, 

October 1991. Texas A&M University Sea Grant 

Program, TAMU-SG-92-109, {composition, 

nutrient ratios, silicate, hypoxia) 

It has been hypothesized that hypoxia and the fate 

of carbon are determined in part by pytoplankton 

size and species composition, which are influenced 

by many environmental factors, but especially by 

the supply of N and Si. In particular when N and 

Si concentrations are high, the predominant algal 

species are large, heavily to moderately silicified 

diatoms. While these may be grazed at rapid rates 

and fuel productivity at higher pelagic trophic 

levels, a large portion either sinks below the 

pycnocline directly or in fecal pellets, perhaps 

leading to hypoxia. When N concentrations are not 
limiting and Si concentrations are low, as often 

happens on the Louisiana continental shelf, either 

lightly silicified diatoms or non-diatoms, usually 

small, flagellated algal species, predominate. These 

organisms yield less vertical flux of organic matter 

to the bottom, because most sinking 

phytoplankton is made up of only a few heavily or 

moderately silicified diatom species. Thus, silicate 

availability determines the vertical flux of directly 

sinking phytoplankton and influences the severity 

and extent of hypoxia. Any changes in riverine 

silicate input will affect hypoxia by this 

mechanism. 

Dortch, Q. and T. E. Whitledge (1992) Does 

nitrogen or silicon limit phytoplankton 

production in the Mississippi River 

Plume and nearby regions? Continental 

Shelf Research 12: 12293- 1309. 

{nutrient limitation) 

The Mississippi River carries very high 

concentrations of nutrients into the otherwise 

oligotrophic Gulf of Mexico, resulting in high 

primary production and hypoxia along the 

Louisiana continental shelf. The hypothesis that 

nitrogen availability controls and ultimately limits 

phytoplankton production ont he shelf was tested 

by measuring an indicator of nitrogen deficiency, 

the ratio of intracellular free amino acidslparticulate 

protien (AAIPr) in the area of the Mississippi 

River plume on a spring ans summer cruise. 

Neither AAIPr ratios or nutrients in the water 

showed nitrogen limitation to be widespread. 

Ammonium concentrations were generally quite 

high, so the lack of phytoplankton nitrogen 

deficiency can be explained by rapid regeneration 

rates. Nitrogen limitation was most likely in the 

summer at high salinities. However, ratios of 

dissolved nutrient concentrations suggested that 

silicate was as likely, or sometimes more likely, to 

be limiting than nitrogen. Although silicate 

depletion may not cause a decrease in productivity, 

it could result in major changes in phytoplanakton 

size and species composition, and ultimately 

influence tropodynamics, regeneration, the fate of 

carbon, and severity and extent of hypoxia. 

Doucette, G. J. (1989) Aspects of iron and 

nitrogen nutrition in two red tide 

dinoflagellates, G y m n o d i n i u m 

s a n g u i n e u m Hirasaka and 

Protogonnyaulax tamarensis (Lebour) 

Taylor. Dissertation for Ph.D., The Univeristy of 

British Columbia. 

{red tide; iron; nitrogen; dinoflagellate; 

phytoplankton; ultrastructure) 

Iron stress-mediated effects on growth, 

biogeochemical composition, iron and nitrogen 

uptake, and ultrastructure have been examined in 

the red tide dinoflagellates G y mnodin ium 

sanguineum Hirasaka and Protogonnyaulax 

tamarensis (Lebour) Taylor. The influence of 

nitrogen source (i.e. NO3 or NH4) on certain iron 

stress-mediated effects was studied, and in some 

comparisons were made with nitrogen stressmediated changes in biogeochemical composition. 

The half-saturation constant for iron-limited growth 

(Ku = 1.7.10-~O M) of G. sanguineum was 

estimated to be 10-1000 times greater than for other 

neritic species investigated previously. Also, the 

iron requirement of this dinoflagellate, in terms of 

FelC ratios, exceeded those of certain coastal 

diatoms by one to two orders of magnitude. FeIN 

ratios demonstrated a larger (1.5-fold) minimum 

iron content of NO3- than NH4-grown cells, likely 

reflecting the iron content of NO3 assimilatory - 

enzymes. Acquisition of nitrogen by Fe-deplete, 

N03-grown cells was sufficiently inhibited to yield 

symptoms of N deficiency, revealed by decreased 

(ca. 1.4-fold) N quotas and free amino acidprotein 

ratios compared to Fe-deplete, NH4-grown cells. 

Reductions in chlorophyll a (chl a) quotas (Qchl) 

and photosynthetic electron transport (PET) 

efficiency (as measured by in vivo fluorescence 

indices) occurred under Fe depletion, and are 

consistent with the essential role of iron and chl a 

and PET component (i.e.) cytochrome and Fe-S 

proteins) biosynthesis. Nitrogen depletion affected 

Qchl similarly, but altered PET efficiency to a 

markedly lesser extent than did Fe depletion. Irondepleted G. sanguineum exhibited an enhanced iron 

transport capacity, which failed to be manifested 

following a transition from NHq to NO3 nutrition. - 

This suppression may result from concurrent iron 

and nitrogen stress, due to the inability of Fedeplete, NHq-grown cells to rapidly assimilate 

NO3. The complete initial inhibition of NO3 

uptake when Fe-deplete, NH4-grown cells were 

given saturating iron additions supports this idea. 

Iron stress caused reductions in chloroplast number 

and some degeneration of lamellar organization in 

this species. For P. tamarensis, iron limitation 

induced the formation of temporary (=pellicle) and 

resting (=hypnozygotes) cysts. Degenerative 

changes in organelle (i.e. chloroplasts, 

mitochondira and chromosomes) ultrastructure were 

largely restricted to pellicular cysts., consistent 

with their hypothesized role of maintaining 

variability over brief, rather than extended (c.f. 

hypozygotes) exposure to adverse conditions. 

Dougherrty, J. E. and M. D. Morgan (1991) 

Benthic community response (primarily 

Chironomidae) to nutrient enrichment and 

alkalinization in shallow, soft water 

humic lakes. Hydrobiologia 215:73-82. 

{benthic; community; nutrients; lake; pH; 

chironomonids) 
A comparison of the benthic fauna found in two 

shallow lakes in the New Jersey Pinelands (USA) 

illustrated the impact of elevated pH and nutrients 

caused by residential and agricultural disturbance on 

a naturally acidic, poorly buffered aquatic 

environment. Detailed community analysis 

suggested that change in community composition 

was a better indicator of response to disturbance 

than biological diversity indices. Chironomonidae 

(insecta) were the predominant components of the 

benthic macroinvertebrate assemblage of both 

undisturbed Oswego Lake (low pH, low nutrients) 

and Nescochague Lake (fluctuating pH, elevated 

nutrients). The genera Procladius, Tribelos, and 

Pagastiella dominated Oswego Lake, where as 

Zulutschia zalutschicola, Procladius, Dicrotendipes, 

and Tanytarsus dominated Nescochague Lake. 

Glyptotendipes was a common and unique member 

of the Nescochague Lake assemblage. Cluster 

analysis indicated that the chemical differences 

between lakes were the most important community 

determinants, although, within each lake, depth and 

substrate affected the local communities. Oswego 

Lake exhibited a departure nonchironomonid 

benthic fauna typical of low nutrient, acid lakes. In 

turn, Nescochague Lake exhibited an enriched 

nonchironomonid fauna including mollusks and 

planaria which were not found in Oswego Lake. 

Dzurica, S., C. Lee, E. M. Cosper, and E. J. 

Carpenter (1 989) Role of environmental 

variables, specifically organic compounds 

and micronutrients on the growth of the 

chrysophyte Aureococcus 

anophagefferens. In: E. M. Cosper, V. M. 

Bricelj and E. J. Carpenter (ed.) Novel 

Phytoplankton Blooms, Causes and Impacts of 

Recurrent Brown Tides and other Unusual Blooms. 

Springer-Verlag , Berlin. 

{brown tide; bloom; growth) 

No abstract 

Eadie, B. J., J. A. Robins, P. Blackwelder, S. 

Metz, J. H. Trefy, B. McKee, and T. A. Nelson 

( 1 9 9 2) A retrospective analysis of 

nutrient enhanced coastal ocean 

productivity in sediments from the 

Louisiana continental shelf. In: (ed.) 

Nutrient Enhanced Coastal Ocean Productivity, 

NECOP Workshop Proceedings, October 1991. 

Texas A&M University Sea Grant Program, 

TAMU-SG-92-109, {production, sediments) 

Sediments have been collected and analyzed to 

obtain evidence in support of the argument that 

anthropogenic nutrient loading has led to changes 

in coastal water quality and increased productivity. 

Cores representing approximately 100 years of 

input show unmistakable signs of increased 

accumulation of organic carbon beginning early in 

the 1900s. Organic tracers show that virtually all 

of this increase appears to be of marine origin. At 

two sites within the plumelhypoxia region, 

preliminary estimates are that 50 to 70 percent 

more organic carbon is presently accumulating than 

at the turn of the century. These preliminary 

interpretations provide strong support for the 

central themes of the NECOP program. Analysis 

and interpretation of further supporting information 

is continuing. 

Edgar, R. K. (1970) The effect of nitrogen 

and phosphorus eutrophication on diatom 

community structure. Dissertation for Ph.D., 

Rutgers University, The State University. 

{diatom; community; nitrogen; phosphorus; 

bacteria; phytoplankton) 

An experimental, field test of the hypothesis that 

diatom community structure changes with changes 

in water quality was conducted by partitioning a 

brook with Plexiglass cylinders and maintaining in 

each different concentrations of nitrate (0.1, 1.0 and 

10.0 mg NO3ll) and phosphate (0.3 and 3.0 mg 

P0411) for 18 days. The conclusive stability with 

respect to treatments of all community structural 

parameters - biotic composition, density, species 

diversity (H) and redundancy - strongly suggests 

that this enrichment had no significant effect upon 

the functional interactions occurring among the 

species composing the community. It is suggested 

that diatom communities are not as sensitive 

functions of water quality as they are purported to 

be in that their sensitivity is conditional upon the 

probabilities of environmental changes affecting the 

functional status of the community and our ability 

to detect such changes. Additionally, the results 

indirectly support the idea that products of bacterial 

metabolism other than nitrates and phosphates may 

exert a controlling influence over algal 

communities. 

El-Sayed, M. K. (1982) Effect of sewage 

effluent on the sediment of Northisvatnet 

(a land-locked fjord), Norway. Marine 

Pollution Bulletin 13:85-88. 

{sewage; fjord; Norway; sediments) 

Nordlsvanet is a land-locked anoxic fjord along 

the west coast of Norway which is used as a natural 

recipient of untreated domestic sewage. The study 

of four core sediments collected from the heavily 

polluted basin of the fjord reflects the enrichment of 

surface sediments by Fe, Mn, Cu, Zn and organic 

matter. This surface enrichment is entirely 

attributed to anthropogenic input. 

Elkins, J. W., S. C. Wofsy, M. B. McElroy, and 

W. A. Kaplan (1981) Nitrification and 

production of N20 in the Potomac: 

evidence for variability. In: B. J. Neilson 

and L. E. Cronin (ed.) Estuaries and Nutrients. 

Humana Press, Clifton, New Jersey. 
{nitrification; ammonia; sewage) 

Extensive measurements were carried out during 

the summer of 1977 and 1978 to define 

concentrations of inorganic nitrogen, 02 and N20 

in the Potomac River. The chemistry of the river 

varied significantly between 1977 and 1978, with 

nitrification rates slower near the city of 

Washington D.C. by more than a factor of 10 in 

1978. The nitrification rate was inversely correlated 

with the rate of fresh water flow into the estuary. It 

appears that production of N20 in 1978 occurred 

mainly as a by-product of nitrification. The 

quantity of N20 released to the atmosphere - 

represented approximately 0.3 percent of sewage 

nitrogen. Conversion was more efficient in the 

summer of 1977, about 1-5 percent, reflecting 

either additional mechanisms for production of N20 - 

or larger yields for gas production in nitrification. 

Elmgren, R. (1989) Man's impact on the 

ecosystem of the Baltic Sea: Energy 

flows today and at the turn of the 

century. Ambio 18:326-332. 

(energy flow; eutrophication; phytoplankton; 

benthic; primary; secondary; fish; production; 

ecosystem) 

The impact of man on the Baltic ecosystem in the 

20th century is summarized by estimating changes 

in major energy flows, expressed as organic carbon. 

Recent eutrophication has increased pelagic primary 

production by an estimated 30-70% and 

sedimentation of organic carbon by 70-190%. 

Above the primary halocline, biomass and 

production of benthos have increased, while in the 

deep waters of the Baltic proper, oxygen deficiency 

has led to the loss of formerly important food 

chains over nearly 100 000 km3 of bottom. The 

net. result is an approximate doubling of 

macrobenthic production, but little increase in 

meiobenthic production. Zooplankton production is 

thought to have increased less than primary 

production, perhaps by about a quarter. Fish catches 

in the Baltic have increased more than tenfold, but 

this is considered to be due only partly to increased 

fishing effort and the near extermination of fisheating Baltic seals, though hunting and pollution, 

are likely to be about equally important. Today, the 

Baltic fishery requires about 10% of the primary 

production in the Baltic to produce its catches, 

whereas the remnant seal population needs less than 

0.1%. At the beginning of this century the fishery 

required about 1% and the marine mammals an 

estimated 5% of the primary production. 

Emara, H. I., M. A. Shriadah, T. H. Moustafa, and 

M. S. El-Deek (1992) Effects of sewage and 

industrial wastes on the chemical 

characteristics of the Eastern Harbour and 

El-Mex Bay waters of Alexandria, 

Egypt. In: R. A. Vollenweider, R. Marchetti 

and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

(oxygen, chlorophyll) 

Sea water samples were collected from the Eastern 

Harbour and El-Mex Bay during the period from 

November 1987 to January 1989. The 

concentrations of dissolved oxygen, hydrogen 

sulphide, inorganic phosphae, nitrate, nitrite, 

ammonia, chlorophyll a and oxidizable organic 

matter are scattered in the ranges (0.00-5.11 ml 1- 

I), (0.00-2.94 mg 1-I), (0.35-17.6 ug at 1-l) 

(0.00-103.5 ug at 1-I), (0.00-21.6 ug at 1-I), 

(0.10-88.0 ug at l-l), (0.00-5.56 mg m-3) and 

(0.10-13.8 mg 02 1-l) respectively. 

In general, El-Mex Bay exhibited higher 

concentrations of nutrients and oxidizable organic 

matter, which reveal higher eutrophication levels 

than the Eastern Harbour. Chlorophyll values were 

low and similar for the surface water of both areas, 

while the bottom water of El-Mex Bay reflected 

slightly higher levels. Hydrogen sulphide 

concentrations showed wide variations, reflecting 

higher values for the Eastern Harbour (1.33 mg 1- 

l) compared with El-Mex Bay (0.98 mg 1-I). 

Hydrogren sulphide-ammonia concentrations 

showed an impressive relationship, with estimated 

ratios of H2S/NHc of 1.7 and 1.03 for the surface 

water of the Eastern Harbour and El-Mex Bay 

respectively. 

The NIP ratio reflects higher values for the surface 

and bottom waters of El-Mex Bay (10.2, 13.2) in 

comparison with the Eastern Harbour (7.0, 7.4), 

and the majority of the investigated areas seems to 

be P-limited. The estimated amount of oxygen 

present after complete oxidation of organic matter 

ranged from 50 to 55% throughout 1988, which is 

about 14% lower than the values recorded during 

the last ten years in El-Mex Bay. 

Eng, C. T., J. N. Paw, and F. Y. Guarin (1989) 

The environmental impact of aquaculture 

and the effects of pollution on coastal 

aquaculture development in Southeast 

Asia. Marine Pollution Bulletin 20:335-343. 

{aquaculture; Asia) 

Coastal aquaculture is a traditional practice in 

Southeast Asia. Accelerated development in the last 

three decades has created negative environmental 

impacts, such as extensive mangrove conversion to 

ponds, changes in hydrologic regimes in enclosed 

waters due to proliferation of aquaculture structures, 

and discharge of high levels of organic matter into 

coastal waters. Similarly, the increasing 

deterioration of coastal water quality resulting from 

the discharge of domestic, agricultural and industrial 

wastes into coastal waters has affected aquaculture 

production and profitability. Furthermore, the 
increased frequency of red tides in the region has 

posed serious threats to coastal aquaculture, 

especially to mollusc cultivation. The introduction 

of management measures to mitigate deteriorating 

coastal water quality and the adverse environmental 

impacts of aquaculture development has now 

become a matter of urgency to the region. 

Fabrikant, R. (1984) The effect of sewage 

effluent on the population density and 

size of the clam Parvilucina tenuisculpta. 

Marine Pollution Bulletin 15:249-253. 

{sewage; density; size; bivalves; Parvilucina; 

sediments; nitrogen; California) 

Changes in the population density and size of the 

clam Parvilucina tenuisculpta were related to 

municipal waste effluent (in particular, organic 

nitrogen concentration) off the coast of Palos 

Verdes, California. Average clam size increased as 

sediment organic nitrogen concentration increased, 

but population density increased until organic 

nitrogen concentration reached a critical level when 

population density decreased dramatically. These 

effects were explained in terms of increased 

available nutrition ( in the form of sediment 

organic nitrogen pollutants) enhancing both 

population and individual size growth. However, 

when a critical concentration of organic 

decomposition products was reached, population 

density decreased due to the toxicity of the released 

compounds. It is suggested that Parvilucina 

tenuisculpta would make a good bioindicator of 

organic enrichment pollution. 

Faganeli, J. (1982) Nutrient dynamics in the 

seawater column in the vicinity of Piran 

submarine sewage outfall. Marine Pollution 

Bulletin 13:61-66. 

(nutrient; sewage; Adriatic; nitrogen; phosphorus) 

Variations of nitrogen and phosphorus 

concentrations in the vicinity of Piran (North 

Adriatic) submarine sewage outfall were studies as 

well as the treatment plant operation. On the basis 

of the data it was concluded that the influence of the 

outfall to the nutrient level in that area is 

negligible. In comparison the data from the shallow 

organically polluted water in the inner part of the 

Bay of Kopper are reported. 

Fanuko, N. (1984) The influence of 

experimental sewage pollution on lagoon 

phytoplankton. Marine Pollution Bulletin 

15: 195-198. 

{sewage; Yugoslavia; lagoon; phytoplankton; 

abundance; Adriatic; bloom) 

In a sewage pollution experiment in the lagoon 

system at Strunjan, Yugoslavia, an artificially 

polluted lagoon showed significantly lower 

chlorophyll a biomass and cell density, and the 

relationship between these two variables was 

altered. 

Faust, M. A. and R. J. Chrost (1981) 

Phytosynthesis, extracellular release, and 

heterotrophy of dissolved organic matter 

in Rhode River estuarine plankton. In: 

B. J. Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{excretion; DOM; phytoplankton; primary 

production; bacteria) 

Rates of phytoplankton photosynthesis, 

extracellular release of dissolved organic matter 

(DOM), and the assimilation of DOM by 

bacterioplankton during in situ incubation were 

measured in the Rhode River estuary. 

Phytoplankton photosynthesis was in the range of 

3 to 147 ug C 1-I h-l. The release of DOM 

appeared to be inversely related to the 

photosynthetic activity of phytoplankton. A large 

percentage of the total carbon (C) fixed was released 

as 14c-D0~, ranging from 4 to 66 percent of the 

total. Ultrafiltration was used to fractionate the 

14c-D0~ into size classes from less than 500 to 

300,000 molecular weight (MW). Bacterioplankton 

readily assimilated a wide range of 14c-D0~ MW 

size fractions. Both photosynthesis and phosphorus 

uptake were higher in nanoplankton. A positive 

linear relationship existed between log surface to 

volume ratio and log phosphorus uptake per 

biomass of phytoplankton. Bacterioplankton 

assimilated phosphorus more rapidly than 

phytoplankton. The species composition of 

plankton populations also constantly changed. The 

14c-D0~ represented only 0.4 to 2.3 percent of 

the total DOM in the Rhode River. Its quantity and 

quality appear to be determined by the changing 

phytoplantkon population with produces it, and the 

changing bacterioplankton population which uses 

it. Preliminary data such as these may yield 

important insight into changes occurring in 

metabolic patterns of brackish ecosystems due to 

biological enrichment with nutrients. 

Ferraro, S. P., R. C. Swartz, F. A. Cole, and D. 

W. Schults (1991) Temporal changes in the 

benthos along a pollution gradient: 

Discriminating the effects of natural 

phenomena from sewage-industrial 

wastewater effects. Estuarine, Coastal and 

Shelf Science 33:383-407. 

{benthos; pollution; sewage; toxic; California; 

wastewater; sediment) 

As pollution from the Los Angeles County 

Sanitation Districts (LACSD) outfalls decreased 

between 1980 and 1983, the macrobenthic 

community partially recovered and surficial(0-2 cm 

deep) sediment contamination and toxicity decreased 

at 60 m water depth along a pollution gradient from 
the outfalls. Pollution from the LACSD outfalls 

continued to decrease but macrobenthic conditions 

and surficial sediment quality deteriorated 1 km, 

was unchanged 3 km, and improved 5-15 km from 

the LACSD outfalls between 1983 and 1986. The 

net effect of natural phenomena is indicated when 

ecosystem changes occur in the opposite directions. 

Our data suggest that the net effect of natural 

phenomena (e.g. winter storms, El Niiio) on the 

benthos was greater than LACSD wastewater 

effects 1 krn, about equal to LACSD wastewater 

effect 3 km, and less than the LASCD wastewater 

effects 5-15 km from the outfalls at the LASCD 

1983-86 mass emission rate. Surficial sediment 

samples collected beyond the 1 km station from the 

LASCD outfalls probably represented * 3 years of 

natural+effluent particulates accumulation, and they 

were, therefore, better suited for detecting long-term 

trends than for testing short-term temporal 

variability in surficial sediments significantly 

increased between 1983 and 1986, probably 

primarily reflecting renewed wastewater effects near 

the outfalls and the effect of natural phenomena 

(e.g. storm-induced sediment transport or erosion) 

further from the outfalls. Since natural phenomena 

may have an effect on the benthos 2 3 years of 

LASCD wastewater effects, short-term benthic 

changes must be interpreted cautiously at the study 

site. 

Fiedler, P. C. (1982) Zooplankton avoidance 

and reduced grazing responses to 

Gymnodinium splendens (Dinophyceae). 

Limnology and Oceanography 27:961-965. 

{zooplankton; red tide; production; grazing) 

A dense subsurface layer of Gymnodinium 

splendens, a feature often observed in coastal waters 

off southern California was actively avoided by 

macrozooplankton. Filtration rates, measured by 

uptake of trace amounts of radiocarbon-labeled 

diatoms, and ingestion rates, measured by gut 

fullness, of some herbivorous species were 

significantly reduced within the layer. These 

behavioral responses may help explain the 

formation and persistence of dinoflagellate blooms 

such as red tides in coastal waters often dominated 

by diatoms with higher maximum growth rates. 

Fleischer, S., S. Hamrin, T. Kindt, L. Rydberg, 

and L. Stibe (1987) Coastal eutrophication 

in Sweden: Reducing nitrogen in land 

runoff. Ambio 16:246-25 1. 

(nitrogen; Sweden; runoff) 

Eutrophication caused by excess nitrogen and the 

resulting plankton blooms in Kattegat have led to 

oxygen deficits in the bottom waters. Regional 

input of nitrogen affects southern Kattegat. The 

interdisciplinary project "Land Use-Water Quality" 

has identified the main source and transport routes 

of nitrogen in the drainage basin (10 100 km2) of 

Laholm Bay. Possible measures to decrease 

nitrogen loading in the coastal waters, such as 

increased cultivation of catch crops and increased 

spreading of manure in the springtime, are 

presented. A holistic approach includes measures 

that may also prove beneficial to a number of 

related agricultural problems, e.g. monotonous 

landscape (very few types of crops, decreasing 

grass- and wet-land areas), sensitivity to 

environmental catastrophes (atmospheric loading by 

radioactive materials or by chemicals), surplus 

production, and the threat to almost 200 wild plant 

species. In addition to direct agricultural measures, 

restoration of wetlands and streams is also 

necessary to reduce nitrogen losses to the coastal 

environment. 

Fletcher, R. L., V. Cuomo, and I. Palomba (1990) 

The "Green Tide" problem, with 

particular reference to the Venice 

Lagoon. British Phycological Journal 25:87. 

{green tide; macroalgae; Venice Lagoon; sewage; 

nutrient) 

The excessive growth of macroalgae is a well 

known consequence of the nutrient enrichment of 

coastal waters by increased levels of sewage 

discharge and agricultural "run-off'. These growths 

are characteristically dominated by green algae, in 

particular species of Enteromorpha and Ulva, and 

are appropriately termed "Green Tides". Such 

growth weas particularly evident during the warm 

summer of 1989 in the shallow, muddy harbours of 

the Solent (south coast of England). Problems 

caused by these algal growths include the fouling of 

nets and dredges, changes in the community 

structure of sediment dwelling invertebrates and the 

excessive deposits of rotting driftweed on amenity 

beaches, requiring removal and disposal by local 

Councils. Nowhere is the problem of "Green Tides" 

more dramatically illustrated than in the Venice 

Lagoon, Northern Adriatic. The present paper 

considers the causes and consequences of these 

luxuriant macroalgal growths in the Lagoon, the 

biological characteristics of the dominant 

contributors and the presently used methods of 

removal, treatment and utilization. 

Foster, P. (1984) Nutrient distributions in 

the winter regime of the Northern Irish 

Sea. Marine Environmental Research 133 1-95. 

{nutrient; Irish Sea; nitrogen) 

Quasi-synoptic distribution of salinity and the 

concentrations of dissolved inorganic nutrients in 

the winter regime of the Northern Irish Sea are 

presented. Salinity to nutrient regression analyses 

and the distributions of nutrient ratios show that 

the characteristics of Atlantic water are variously 

modified during their northward passage through the 

Irish Sea. Inherently different chemical 

characteristics of the fresh water sources discharging 
into the Northern Irish Sea generate a 

conglomeration of water types in the area. Marine 

waters along the Irish coast are relatively enriched 

in silicon, while waters adjacent to the eastern 

coastal boundary are relatively enriched by 

anthropogenic nitrogen sources. Possible 

implications of the spatial dichotomy in nutrient 

status for the seasonal production cycle in the 

northern Irish Sea are considered. 

Fransz, H. G. and J. H. G. Verhagen (1985) 

Modelling research on the production 

cycle of phytoplankton in the southern 

bight of the North Sea in relation to 

riverborne nutrient loads. Netherlands 

Journal of Sea Research 19:241-250. 

{modelling; phytoplankton; production; nutrient; 

North Sea) 

The physio-chemical and biological composition 

of North Sea water off the Dutch coast is influenced 

by various factors including meteorological 

conditions, wind and tide induced flows and mixing, 

river pollution loads, waste loads and various 

physical, chemical and biological processes. The 

long term variations (> - 1 month) of the 

biological composition are mainly caused by 

fluctuations in meteorological conditions and 

concentrations of nutrients for algae. 

The loads of phosphorus and nitrogen compounds 

on the North Sea have increased significantly since 

the early 20th century. Especially the discharge of 

phosphorus (P) and nitrogen (N) transported by the 

river Rhine has increased about sixfold since 1930. 

The nutrient rich river water originating from the 

rivers Scheldt, Meuse and Rhine, mixes with sea 

water in such a way that 50% of the river discharge 

is transported in a 15 km wide strip along the 

Dutch and Belgian coast. Especially in this coastal 

area, the increase in nutrient loads has had a marked 

influence on the nutrient concentrations. In fact, the 

increase in N and P and the essentially constant 

discharge of reactive silicate (Si) caused a change in 

nutrient, first depleted by phytoplankton, from N or 

P some 50 years ago to Si nowadays. 

The question, to what extent the increase in 

nutrient enrichment has given rise to enhanced 

phytoplankton growth, or to changes in species 

composition, is difficult to answer because of a 

lack of reliable data from the past. Nevertheless it 

is conceivable that at present primary productivity 

may occasionally be higher than in the past. The 

upper limit to the recurrent diatom blooms in 

Dutch coastal waters is set by the amount of 

available silicate. An increase in the upper limit is 

not to be expected due to the constancy of the Siload during the last 50 years. Some flagellate 

species, however, appear to be phosphorus limited 

in spring and locally dominate diatoms. Nitrogen 

compounds maintain low concentrations 

throughout the summer. Therefore, it is 

conceivable that the abundance of the specific 

flagellate species is closely related to the riverborne 

P- and N-loads. 

The research as described in this paper intends to 

simulate the phytoplankton biomass and 

composition in relation to meteorological and 

hydrophysical conditions, -nutrient loads and 

zooplankton biomass. It can be considered as a 

theoretical contribution towards a better 

understanding of the production and eutrophication 

processes in this polluted and exploited part of 

North Sea. 

Fraser, T. H. and W. H. Wilcox (1981) 

Enrichment of a subtropical estuary with 

nitrogen, phosphorus and silica. In: B. J. 

Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{enrichment; nitrogen; phosphorus; silica; estuary; 

phytoplankton) 

Seasonal pulses of nutrients are delivered to 

Charlotte Harbor from the Peace River and other 

tributaries. Greatest loads occur during the summer 

wet season. Phosphate and nitrate values are much 

higher in the Peace River than in adjacent 

tributaries while ammonium and silica are similar 

among the streams. 

Phosphate dilution curves in the estuary suggest 

that changing concentrations along the chloride 

gradient are the result of mixing processes. 

Inorganic nitrogen and silica dilution curves 

suggest non-conservative processes are occurring, 

since their concentrations decrease faster along the 

chloride gradient than can be explained by dilution 

alone. Thus, both inorganic nitrogen and silica 

could become limiting factors in different parts of 

Charlotte Harbor during portions of the annual 

cycle, while phosphorus is always abundant. 

Phytoplankton populations respond positively to 

seasonal pulses of nutrients with higher 

productivity occumng during or just after high river 

flow. Productivity increases from the lower harbor 

to near the river mouth in all seasons. General 

population levels and productivity are similar to 

other Florida estuaries, with diatoms dominating 

the phytoplankton (blue-green algae less than 1 

percent) community even with the increased 

nutrient levels. 

Frid, C. L. J. and T. S. Mercer (1989) 

Environmental monitoring of caged fish 

farming in macrotidal environments. 

Marine Pollution Bulletin 20:379-383. 

{fish farming; benthos; sedimentation) 

Recent years have seen considerable expansion in 

fish farming using cages. These have until recently 

been located in the sheltered and low current 

environments of sea lochs and fjords. Significant 

benthic enrichment is known to occur at these 

sites, but the effects are highly localized. Fish farm 
development is now occumng in areas with strong 

tidal flows. The sitting of caged fish farming in 

macro-tidal environments may reduce the 

environmental impact of the industry on benthic 

communities at the farm site. However there exists 

the potential for accumulation of farm wastes in 

nearby sedimentary sinks. Additionally the longer 

residence time of the wastes in the water column 

has the potential, especially in regions with already 

high nutrient loadings and/or long flushing times, 

to lead to stimulation of phytoplankton blooms. 

Environmental monitoring of such developments 

therefore needs to consider these addition effects. 

Friligos, N. (1981) Enrichment by inorganic 

nutrients and oxygen utilization rates in 

Elefsis Bay (1973-1976). Marine Pollution 

Bulletin 12:43 1-436. 

{nutrients; oxygen; Elefsis Bay; anoxic) 

This paper concerns the enrichment by nutrients 

and the oxygen utilization rates of an intermittently 

anoxic basin. The concentration of nutrients and 

dissolved oxygen have been determined during 

seasonal surveys of water characteristics. The 

oxygen utilization rates ranged between 1.3 and 1.8 

ml 1-I month-l from March to June and were 

considerably higher than those reported for oceanic 

waters. The nutrients enrichment appears to be 

considerably influenced by eutrophication and 

human activity. During anoxic conditions in 

summer, the mineralization of organic matter was 

associated with consequent accumulation of 

enormous amounts of nitrogen (ammonia), 

phosphates and silicates. Nutrient ratios underwent 

great fluctuation mainly due to anoxic conditions. 

Friligos, N. (1982) Enrichment of inorganic 

nutrients in the inner Saronikos Gulf 

(1973-1976). Marine Pollution Bulletin 

13: 154-158. 

{nutrients; Saronikos Gulf; Greece; nitrogen; 

phosphorus) 

This paper is concerned with the enrichment of 

inorganic nutrients in the Inner Saronikos Gulf near 

the Athens sewage outfall. This Inner Gulf tends to 

accumulate nutrients above the background level, 

expecially ammonia which was about four times 

more than the background value. Comparison of 

nutrient enrichment was made between the Inner 

Gulf and Elefsis Bay. It was found that Elefsis Bay 

contained considerably more nutrients than the 

Inner Gulf. This was mainly due to the different 

sources of nutrients as well as morphology of each 

area and the circulation of the waters. 

Friligos, N. (1985) Compensation effect of 

phytoplankton on nutrients from a 

sewage outfall in summer. Hydrobiologia 

126:53-58. 

(sewage; nutrients; phytoplankton; abundance} 

This paper investigates the relationship between 

nutrients, chlorophyll a and physical variables in 

the upper Saronikos Gulf, an oligotrophic marine 

environment south of the Greater Athens 

Metropolitan Area. Phosphate, silicate, ammonia, 

nitrite, nitrate, salinity and dissolved oxygen were 

determined at eight stations on 9 occasions during 

summer 1982. A thermocline led to the 

stratification of the water column and the 

pycnocline was related to the thermocline. The 

values of oxygen were more or less normal. The 

eutrophication of seawater in the vicinity of the 

sewage outfall was demonstrated by surface levels 

of chlorophyll a being forty to two hundred times 

above background. This parameter provides 

evidence for a high phytoplankton standing stock. 

However, there were no appreciable differences 

between the nutrients in the outfall area and those 

in the background. This suggests rapid uptake of 

nutrients and/or effective dispersal from the outfall. 

Friligos, N. (1985) Impact on phytoplankton 

populations of sewage discharges in the 

Saronikos Gulf (West Aegean). Water 

Research 19: 1 107- 1 1 18. 

{sewage; nutrients; phytoplankton; standing stock) 

Temperature, salinity and concentrations of 

dissolved oxygen, nutrients, chlorophyll a, 

particulate carbon and nitrogen concentrations in 

sea water near the ~eratsini sewage outfall are 

reported. Suggestions are made on the proposed 

length of the outfall diffuse and the extent of the 

sewage treatment. Eutrophication in the area around 

the outfall is demonstrated by enhanced chlorophyll 

a, particulate carbon and particulate nitrogen 

concentrations, for they provided evidence for high 

phytoplantkon standing stock. However no 

appreciable differences in the inorganic nutrient 

levels between the outfall area and the normal 

background were observed. This suggests rapid 

uptake of nutrients andlor effective dispersal from 

the outfall. The study indicates that in the case of 

oligotrophic waters and in the absence of the 

dissolved oxygen depletion, primary treatment may 

be preferable to secondary treatment. 

Friligos, N. (1989) Nutrient status on a 

silled embayment in the Ionian Sea (Gulf 

of Patras). Toxicological and Environmental 

Chemistry 20-21:2 1-27. 

{nutrient; Gulf of Patras; Mediterranean; nitrate; 

silicate} 

This paper is concerned with the enrichment of 

inorganic nutrients in the Patraikos Fulg. The Gulf 

of Patras is a silled embayment opening into the 

Ionian Sea on the west, and through the straits of 

Rio onto the Gulf of Corinth, on the east. The 

latter communicates though the Corinth canal with 

the Aegean Sea. Results of the Patraikos Gulf 

show only slighter greater concentrations of 
nutrients than background, in particular nitrate and 

silicate. Also a comparison is made with the 

nutrient concentrations in polluted coastal gulfs of 

the Aegean. The different nutrient levels are due to 

the different sources of nutrients as well as the 

morphology and the circulation of the waters. 

Gast, V. (1985) Bacteria as a food source 

for microzooplankton in the Schlei Fjord 

and Baltic Sea with special reference to 

ciliates. Marine Ecology Progress Series 

22: 107-120. 

{bacteria; microzooplankton; Baltic; ciliates; 

Uronema) 

In situ vestigations revealed that number and 

biomass of microzooplankton increased with 

eutrophication along the length of the Schlei Fjord 

(FRG). The same observation was made for total 

bacterial numbers and biomass. Microzooplankton 

of the Schlei and total bacterial numbers showed a 

minimum in winter and major periods of 

development in late summer/autumn and spring. 

Usually the microzooplankton biomass in the 

Schlei was greater than the bacterial biomass. In 

contrast, the bacterial biomass for 5 of 6 stations in 

the Baltic Sea surpassed that of the 

microzooplankton during the summer. Number and 

biomass of microzooplankton in both bodies of 

water can mostly be attributed to protozoans of the 

3 to 30 um fraction. Determined with the aid of 

radioactivity labeled bacterial cultures, the filtration 

rate of 'natural' microzooplankton populations 

exhibited a distinct dependency on 

microzooplanktonic biomass and water temperature. 

In March 1982 microzooplankton populations in 

the eutrophic Schlei Fjord filtered 5 to 58 % of the 

water per day. In the central Baltic Sea in August 

1982 the rate was 70 % d-l during the late stage of 

a decaying blue-green algae bloom. Laboratory 

experiments with Uronema marinum clearly 

showed that bacteria concentration exert a 

considerable imjluence on the development of 

ciliates. Only when a limiting concentration of 

about 1 x lo6 bacteria ml-l is attained does a 

proliferation of ciliates commence. Hence, bacteria 

can represent an important food source for 

microzooplankton, especially in biotopes with a 

high bacterial number and biomass. 

Gerlach, S. A. (1988) Eutrophication of 

Kieler Bucht. Kieler Meeresforschungen, 

Sonderheft 654-63. 

{Baltic; nitrogen; phosphorus; nutrients) 

Prior to 1975, average winter water nutrient 

concentrations at station Bokniseck in Kieler Bucht 

(Western Baltic) were 1.23 mmol/m3 total 

phosphorus and 12.7 mmol/m3 dissolved inorganic 

nitrogen. Nitrogen concentrations did not change 

until 1984. The mean of a few total phosphorus 

data from 1980-1984 is 1.94 mmol/m3. Mean 

nutrient concentrations in the area between the 

southern entrance of the Great Belt and Darss Sill 

increased between 1975 and 1984 from 0.8 to 1.6 

mmol/m3 total phosphorus and from about 6 to 10 

mmol/m3 inorganic nitrogen. Comparatively 

higher nutrient concentrations at Bokniseck station 

are correlated to higher salinity. However, no data 

are available on nutrient imports into Kieler Bucht 

with saline deep water. Unchanged nutrient 

concentrations at Bokniseck prior to 1975 are an 

argument against any dominant influence of 

anthropogenic inputs which until 1974 increased to 

annually 1500 t of total phosphorus and 12 700 t 

of total nitrogen, plus 5000 t of nitrogen from the 

atmosphere. Higher phosphorus concentrations 

1980-1984 are correlated to severe oxygen 

deficiency in the deep water. Mobilization of 

phosphorus from sediments becoming anoxic could 

be the reason. Biota could be such sources and 

sinks of nutrients that influence the nutrient 

balance of Kieler Bucht. To sum up, there are 

several processes besides anthropogenic inputs that 

influence nutrient concentrations in water. An 

assessment of winter water nutrient concentrations 

alone is no adequate tool for the evaluation of the 

effects of anthropogenic nutrient inputs via rivers, 

sewage, land runoff, groundwater and from the 

atmosphere. 

Gerlach, S. A. (1990) Nitrogen, phosphorus, 

plankton and oxygen deficiency in the 

German Bight and in the Kiel Bay. Kieler 

Meeresforschungen, Sonderheft 7: 1-341. 

{German Bight; Kiel Bay; Baltic; phosphate; 

nitrate; nitrite; ammonium; nitrogen; phosphorus; 

nutrients; oxygen; hydrogen sulphide; bloom; 

phytoplankton; biomass; abundance; sediments; 

turbidity; atmosphere; review; assemblage) 

No abstract 

Giesen, W. B. J. T., M. M. v. Katwijk, and C. d. 

Hartog (1990) Eelgrass condition and 

turbidity in the Dutch Wadden Sea. 

Aquatic Botany 37:71-85. 

{eelgrass; Zostera; Dutch Wadden Sea; turbidity; 

submerged vegetation; light) 

Populations of eelgrass (Zostera marina L.) in the 

Dutch Wadden Sea have witnessed two major 

phases of decline this century. The first was the 

total disappearance of sublittoral beds during the 

wasting disease epidemic of the early 1930s, and 

their subsequent failure to recover. The second was 

the gradual disappearace of littoral eelgrass after the 

mid-1960s. It is argued here that both the lack of 

recovery of the sublittoral beds, and the 

disappearance of the littoral population, may have 

been effected, to a large extent, by changes in 

turbidity. The increasing turbidity can be linked 
with progressively increasing eutrophication as 

well as deposit extracting and dredging activities. 

Gieskes, W. W. C. and G. W. Kraay (1977) 

Contin~lous plankton records: Changes in 

the plankton of the North Sea and its 

eutrophic southern bight from 1948 to 

1975. Netherlands Journal of Sea Research 

11:334-364. 

{plankton; eutrophication) 

No abstract 

Gilmartin, M. and N. Revelante (1980) Nutrient 

input and the summer nanoplankton 

bloom in the Northern Adriatic Sea. 

P.S.Z.N.1: Marine Ecology 1: 169- 180. 

{nutrient; phytoplankton; nanoplankton; 

microplankton; eutrophicated) 

A four-year oceanographic data set from the 

Northern Adriatic Sea identified the factors favoring 

development of the nanoplankton or microplankton 

components of the phytoplankton crop. 

Latitudinally atypical summer nanoplankton 

blooms, perhaps of recent origin, develop under 

vertically stratified conditions in the presence of 

high nutrient concentrations (including silicates). 

Including the summer blooms, the nanoplankton 

were responsible for more than half the total 

phytoplankton production about 90% of the time. 

No single mechanism governed phytoplankton cell 

size or size succession, but rather a number of 

mechanisms which were not mutually exclusive. 

Giovanardi, F. and E. Tromellini (1992) 

Statistical assessment of trophic 

conditions. Application of the OECD 

methodology to the marine 

environment. In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{chlorophyll, phosphorus, Adriatic Sea) 

In order to study the time-space variability of the 

Po-Adriatic system, a multivariate statistical 

method has been used. This approach allows the 

determination of the source of variability and the 

parameters that mainly maintain the system in a 

perturbed state. The OECD classification of 

trophic state for inland waters has therefore been 

applied to the marine environment. This trophic 

classification criterion has been interpreted in terms 

of probability of assignment to the different trophic 

categories. Functional inter-relationships between 

total phosphorus (TP) and trophic indicators 

(chlorophyll and peak chlorophyll) have been 

determined. Based on the regression of TP versus 

chlorophyll and on the log-normal distribution of 

the chlorophyll data, an empirical model (Monte 

Carlo method) has been developed, thus determing 

the probabilities associated with the trophic level in 

the Emilia-Romagna coastal waters. The 

application of this model is quite useful for testing 

the environmental consequences depending on 

different interventions for phosphorus loading 

reductions. 

Goldman, C. R. (1988) Primary productivity, 

nutrients, and transparency during the 

early onset of eutrophication in ultraoligotrophic Lake Tahoe, CaliforniaNevada. Limnology and Oceanography 33: 132 1 - 

1333. 

{primary productivity; transparency; turbidity) 

For more than half a century, the trophic status of 

water bodies has been of interest to limnologists 

and oceanographers alike. This report demonstrates 

the close, inverse relationship between 14cestimated primary productivity and transparency 

during the earliest stages of cultural eutrophication. 

As the population in the Tahoe basin has rapidly 

increased, Lake Tahoe has been characterized by an 

increase in primary productivity that has averaged 

5.6% yr-l for the last 28 yr. There has been a 

concomitant decline in transparency of 0.37 m yr-l. 

During winter months when transparency is 

highest, the average annual loss has been slightly 

greater (0.40 m yr-l). The average annual Secchi 

depth has decreased by 7 m during the last 19 yr of 

intensive monitoring. There has also been a 

significant increase in the light extinction 

coefficient. Photosynthetic efficiency has increased 

while there has been a gradual shrinkage of the 

euphotic zone. During the same period the total 

NO3--N content of the lake has increased 

significantly, but total P content has not. Lake 

Tahoe primary production has become increasingly 

P sensitive during the last decade as N has 

accumulated in the system. A gradual increase in 

the N:P may prove to be a general evolutionary 

characteristic of oligotrophic lakes during the 

earliest stages of eutrophication. The importance of 

variability is particularly evident from this study. 

Goldman, J. C. (1976) Identification of 

nitrogen as a growth-limiting nutrient in 

wastewaters and coastal marine waters 

through continuous culture algal assays. 

Water Research 10:97-104. 

{wastewater; nitrogen; nutrients; phytoplankton; 

assay; N/P ratio) 

Two series of continuous culture algal assays 

conducted in 1973 and 1974, totaling 63 

experiments, were performed on mixtures of 

seawater and wastewater - both treated and untreated 

- from five locations along the coasts of MA and 

RI. The results conclusively show that nitrogen 

was the growth-limiting nutrient in these 

wastewaters and in coastal marine environments 

receiving such wastes. There was a linear 

relationship between total inorganic nitrogen (EN) 
in the influent and particulate nitrogen (PN) 

representing algal biomass up to a EN 

concentration of about 10 mg 1-l. In addition, the 

N:P ratios in the test alga, Phaeodactylurn 

tricornuturn, varied between 10 and 20 (by atoms), 

whereas the N:P ratios in the wastewater-seawter 

mixtures were between 4 and 12, thus providing 

additional evidence that nitrogen was limiting. 

Enrichment with nitrogen and/or phosphorus to 

span a range of N:P ratios from 3.9 to 20 resulted 

in a linear response in PN concentration to (sum)N 

additions. Additions of P had no effect on algal 

growth. Removal of detergent-P in communities 

practicing marine waste disposal will hence have 

little impact on the control of eutrophication in 

coastal waters. 

Goldman, J. C., Y. Azov, C. B. Riley, and M. D. 

Dennett (1982) The effect of pH in 

intensive microalgal cultures. I. Biomass 

regulation. Journal of Experimental Marine 

Biology and Ecology 57: 1 - 13. 

{pH; phytoplankton; biomass; Phaeodactylurn; 

Dunaliella; carbon) 

Two freshwater and two marine algal species were 

grown in intensive continuous cultures at a fixed 

dilution rate of 0.5 day-l, but a varying pH levels 

in the range 7.6 to 10.6. Both freshwater species 

Scenedesmus obliquus (Turp.) Kutz. and Chlorella 

vufgaris Beij., grew up in pH 10.6 although C. 

vulgaris was more adversely affected by alkaline pH 

than was Scenedesmus obliquus . Of the marine 

species, Phaeodactylum tricornutum (TFX- 1) 

Bohlin was hardly affected by varying pH up to its 

maximum tolerable level of 10.3, whereas growth 

of Dunaliella tertiolecta (Dun) Butcher was 

adversely affected by increasing pH and ceased when 

the pH exceeded 9.3. These results are consistent 

with the general observations that many marine 

species cannot tolerate alkaline pH values much 

above 9.5. Moreover, the unique ability of 

Phaeodactylurn tricornutum to grow at pH > 10 

probably is a major factor contributing to its well 

documented success in large-scale outdoor cultures 

that are poorly buffered. It is difficult to separate 

metabolic from purely chemical factors that 

influence the pH tolerance limits of the individual 

species. The lower pH limits were, however, 

distinctly controlled by the production of alkalinity 

concomitant with NO3- uptake, whereas the upper 

pH limits in the case of Scenedesrnus obliquus and 

Phaeodactylurn tricornuturn seemed to be regulated 

primarily by metabolic control. In no case was the 

availability of inorganic carbon an influencing 

factor in setting the maximum attained pH. 

Goldman, J. C., C. B. Riley, and M. R. Dennett 

(1982) The effect of pH in intensive 

microalgal cultures. 11. Species 

competition. Journal of Experimental Marine 

Biology and Ecology 57: 15-24. 

{pH; compsitition; Phaeodactylum; Dunaliella; 

phytoplankton) 

The results of a series of competition experiments 

between the chlorophyte Dunaliella tertiolecta 

(Dun) Butcher and the diatom Phaeodactylurn 

tricornutum (TFX-1) Bohlin demonstrate 

conclusively that Phaeodactylurn tricornuturn 

dominates in intensive marine cultures when the 

pH rises above =lo. This dominance results 

because of the diatom's unique ability among 

marine species to tolerate alkaline conditions. 

When the pH is regulated both freshwater and 

marine algae, once firmly established in culture at 

their respective pH optima, can resist invasion 

from competing species. Hence, pH control may be 

a method for maintaining species other than P. 

tricornuturn in mass culture. When Dunaliella 

tertiolecta, however, is grown under even slight pH 

stress it becomes susceptible to invasion by 

Phaeodactylurn tricornuturn. This susceptibility to 

takeover by P. tricornutum increases with 

increasing pH. In contrast, the freshwater 

chlorophyte Chlorella vulgaris Beji., which also is 

sensitive to increasing pH, is capable of remaining 

dominant at any pH within its tolerance range when 

invaded by the pH-insensitive chlorophyte, 

Scenedesmus obliquus (Turp.) Kurz. Although 

allelopathic interactions may be responsible for the 

success of Chlorella vulgaris under seemingly 

stressful conditions, the success of Phaeodactylurn 

tricornuturn at increasingly higher pH seems to be 

related primarily to the alga's pH tolerance 

characteristics and not to any chemical interactions 

with competing species. 

Gowen, R. J. and N. B. Bradbury (1987) The 

ecological impact of salmonid .farming in 

coastal waters: A review. Oceanography and 

Marine Biology: Annual Review 25563-575. 

{fish farming; organic enrichment; sediments; 

review) 

No abstract 

Goldman, C. R. (1988) Primary productivity, 

nutrients, and transparency during the 

early onset of eutrophication in ultraoligotrophic Lake Tahoe, CaliforniaNevada. Lirnnology and Oceanography 33: 1321- 

1333. 

{primary productivity; transparency; turbidity) 

For more than half a century, the trophic status of 

water bodies has been of interest to limnologists 

and oceanographers alike. This report demonstrates 

the close, inverse relationship between 14cestimated primary productivity and transparency 

during the earliest stages of cultural eutrophication. 

As the population in the Tahoe basin has rapidly 

increased, Lake Tahoe has been characterized by an 
increase in primary productivity that has averaged 

5.6% yr-l for the last 28 yr. There has been a 

concomitant decline in transparency of 0.37 m yr-l. 

During winter months when transparency is 

highest, the average annual loss has been slightly 

greater (0.40 m yr-l). The average annual Secchi 

depth has decreased by 7 m during the last 19 yr of 

intensive monitoring. There has also been a 

significant increase in the light extinction 

coefficient. Photosynthetic efficiency has increased 

while there has been a gradual shrinkage of the 

euphotic zone. During the same period the total 

N03--N content of the lake has increased 

significantly, but total P content has not. Lake 

Tahoe primary production has become increasingly 

P sensitive during the last decade as N has 

accumulated in the system. A gradual increase in 

the N:P may prove to be a general evolutionary 

characteristic of oligotrophic lakes during the 

earliest stages of eutrophication. The importance of 

variability is particularly evident from this study. 

Gowen, R. J., P. Tett, and K. J. Jones (1992) 

Predicting marine eutrophication: the 

yield of chlorophyll from nitrogen in 

Scottish coastal waters. Marine Ecology 

Progress Series 85: 153- 161. 

{phytoplankton; chlorophyll; prediction; model; 

Scottish water; nitrogen; nitrite; nitrate} 

There is perceived to be a problem of 

eutrophication in European marine coastal waters 

and hence a need to predict the response in terms of 

enhanced biomass of phytoplankton resulting from 

the input of anthropogenic nitrogen. This response 

was investigated indirectly by studying the 

relationship between nitrate and phytoplankton 

chlorophyll concentrations in waters of the Scottish 

west coast. Two-thirds of the 60 data sets analyzed 

gave significant inverse regression of chlorophyll 

on nitrate concentration. This result is explained by 

interpreting synoptic data as representing variation 

in time, with nitrate decreasing as a result of its 

assimilation by phytoplankton and conversion into 

chlorophyll-containing biomass. Thus the absolute 

value of the slope of each significant regression 

estimates the yield (q) of chlorophyll from nitrate 

and, indeed, from any form of nitrogen assimilable 

by microalgae. The median value for q was 1.05 

mg chl (mmol N)-~; the range from 0.25 to 4.4 

encompassed 95% of values. Some, but not all, of 

the variation in q could be explained by error in 

individual estimates due to chemical-analytical and 

sampling errors or to inhomogeneities in each 

sampled phytoplankton population. The remaining 

variation in q included a seasonal trend, which 

might have resulted from changes in phytoplankton 

species composition, nutrient limitation status, or 

the balance between anthropogenic and 

heterotrophic partitioning of nitrogen. It is 

suggested that an appropriate value of q can be used 

to predict the potential maximum increase in 

phytoplankton which would result from a given 

anthropogenic nitrogen discharge. The sensitivity 

of such predictions to error in determination of q is 

discussed, and the values obtained for q compared 

with observations in algal culture and mesocosms. 

GranBli, E. (1981) Bioassay experiments in 

the Falsterbo Channel- nutrients added 

daily. Kieler Meeresforschungen, Sonderheft 

5:82-90. 

{bioassay; Baltic; nutrients; phytoplankton; 

primary production; composition) 

In situ enrichment bioassays were performed 

during the summers of 1978 and 1979 in the 

Falsterbo Channel, south Baltic Sea. Phosphorus 

andlor nitrogen was added daily for up to two weeks 

to 200 1 polyethylene bags with unfiltered surface 

water. Additions of nitrogen or nitrogen plus 

phosphorus invariably increased the biomass and 

14c fixation of phytoplankton. Phosphorus 

additions had no such effects. Phytoplankton 

species which reacted most strongly to the 

enrichment were Aphanotheca sp., Nodularia 

spumigena, Skeletonema costatum, Nitzschia 

closterium, Chaetoceros wighami and Oocystis sp. 

The mean CIChl a quotient was around 70 for 

chlorophyll a values below 6 mg m-3 but decreased 

to about 30 for chlorophyll a values above 10. 

GranBli, E., P. Carlsson, P. Olsson, B. Sundstrom, 

W. GranBli, and 0. Lindahl (1989) From 

anoxia to fish poisoning: The last ten 

years of phytoplankton blooms in 

Swedish marine waters. In: E. M. Cosper, 

V. M. Bricelj and E. J. Carpenter (ed.) Novel 

Phytoplankton Blooms; Causes and Impacts of 

Recurrent Brown Tides and Other Unusual Blooms. 

Springer-Verlag, Berlin. 

{phytoplankton; bloom; Sweden; effects) 

No abtract 

GranCli, E., W. Graneli, and L. Rydberg (1986) 

Nutrient limitation at the ecosystem and 

the phytoplankton community level in 

the Laholm Bay, south-east Kattegat. 

Ophelia 26: 181-194. 

{nutrient; phytoplankton; abundance; nitrogen; 

phosphorus) 

Nutrient enrichment experiments in the laboratory 

with surface waters from the Laholm Bay, indicate 

that phytoplankton is nitrogen-limited. Phosphorus 

addition to phytoplankton samples rarely caused a 

biomass increase in aggreement with negligible 

alkaline phosphatases activity. Usually the 

inorganic-N/P quotient in surface waters was also 

below the Redfield ratio (16:1, by atoms), 

indicating potential nitrogen limitation. The supply 

ratio (runoff and deep water) of inorganic nutrients 
to the surface water of the bay was 23N: 1P for 

March-October and thus indicates a mean shortage 

of phosphate. During summer when river runoff is 

small, the low Nff-ratio in the deep water (10:l) 

which is mixed into surface water suggests nitrogen 

limitation, in aggreement with the enrichment 

experiments. Benthic denitrification and the low 

NIP-ratio of zooplankton excretion may make 

regenerated primary production nitrogen-limited. 

The recent suggestion by Smith (1984), that 

marine ecosystems, like lakes, are phosphoruslimited because any nitrogen shortage is alleviated 

through nitrogen fixation, does not apply to the 

bay itself, where water exchange is too rapid to 

allow for an adjustment of the NIP-supply quotient. 

Grantli, E., H. Persson, and L. Edler (1986) 

Connection between trace metals, 

chelators and red tide blooms in the 

Laholm Bay, SE Kattegat - An 

experimental approach. Ma r i n e 

Environmental Research 18:61 - 78. 

{copper; dinoflagellates; species composition} 

The influence of the trace metals, copper, 

aluminum and iron, and of the strong complexing 

agents, EDTA and NTA, on phytoplankton growth 

in water from a brackish water bay was investigated 

through bioassay experiments. A diatom 

(Skeletonema costatum (Grev.) Clev) and a 

dinoflagellate (Prorocentrum minimum (Pav.) J. 

Schiller) were used as test organisms. The growth 

of both phytoplankton species was strongly 

inhibited by copper. This inhibition was generally 

inhibited by EDTA and NTA. Both phytoplankton 

species were considerably less inhibited by 

aluminum than by copper at the same metal 

concentration. While S. costatum responded to 

copper and chelator additions in the same way in 

sea water samples from different seasons, the 

growth of P. minimum exhibited pronounced 

seasonal variation. Other parameters than the 

values of DCU must be considered in order to 

account for the experimental results. This work 

supports the theory that alterations in contents of 

trace metals and natural chelators in seawater are 

important factors behind shifts in phytoplankton 

species composition. 

GranBli, E., S. Schulz, U. Schiewer, D. 

Gedziorwska, W. Kaiser, and M. Plinski (1988) Is 

the same nutrient limiting potential 

phytoplankton biomass formation in 

different coastal areas of the Southern 

Baltic? Kieler Meeresforschungen, Sonderheft 

6:191-202. 

{phytoplankton; biomass; Baltic; nutrients; 

chlorophyll) 

During the winter 1984 and summer 1985, a 

series of limiting nutrient experiments (enrichment 

tests) were performed with coastal Baltic water by 

scientists from the German Democratic Republic, 

Poland and Sweden. The water used for the 

experiments was collected outside Warnemiinde 

(GDR); Rostock (GDR), Sopot (P) and Falsterbo 

(S). The bioassays were performed under similar 

laboratory conditions. Nitrogen was the potentially 

most limiting nutrient for phytoplankton biomass 

formation in coastal water from Rostock, Sopot 

and Falsterbo. During winter, before the spring 

phytoplankton bloom had started, phosphorus was 

the "most limiting" nutrient in the Rostock area. 

For the Warnemiinde area, no clear limiting 

nutrient was found, except during July 1984, when 

nitrogen addition doubled the phytoplankton 

biomass. The lowest algal standing stocks were 

found for Warnemiinde and Flasterbo, followed by 

Sopot. Extremely high chlorophyll-a values were 

found outside Rostock. Our results show that these 

coastal areas in the southern Baltic do not differ 

essentially from each other with respect to the most 

limiting nutrient (sensu Liebig), although their 

nutrient levels are quite different. Nutrient 

limitation experiments are discussed in the context 

of assumed eutrophication of the Baltic. 

GranBli, E. and K. Sundback (1985) The 

response of planktonic and microbenthic 

algal assemblages to nutrient enrichment 

in shallow coastal waters, Southwest 

Sweden. Journal of Experimental Marine 

Biology and Ecology 85:253-268. 

{nutrient; nitrogen; phosphorus; microalgae; 

phytoplankton; production; biomass} 

Field and laboratory nutrient (nitrogen and 

phosphorus) enrichment studies were performed 

among natural phytoplankton and 

microphytobenthic assemblages from the brackish 

water ~resund, S.W. Sweden. The response of 

algae from a low-nutrient area (Falsterbo Canal) 

was compared to that of algae from a polluted, 

nutrient-rich area (Lomrna Bay). 

The biomass (measured as chlorophyll a) of both 

phytoplankton and microphytobenthos from the 

Falsterbo Canal increased after the addition of 

nitrogen. Phytoplankton growth was stimulated by 

the addition of phosphorus to the nitrogen-rich 

water of the polluted Lomma Bay. Sediment 

chlorophyll a showed no significant increase after 

the addition of nutrients in the Lomma Bay. In 

containers without sediment, phytoplankton uptake 

was calculated to account for = 90 % of the 

disappearance of inorganic fixed nitrogen from the 

water. In the sediment containers the 

microphytobenthos was estimated to account for = 

20% of the nitrogen uptake. The rest was 

presumably lost mainly through denitrification. 

When containers with microphytobenthos from 

Lomma Bay were kept in the dark, phosphorus was 

released at a rate of up to = 180 uM mm2 day-l. We 

suggest that by producing oxygen microbenthic 
algae keep the sediment surface oxygenated thereby 

decreasing phosphorus transport from the sediment 

to the overlying water. 

Grantli, E., B. Sundstrom, L. Edler, and D. M. 

Anderson (ed.) (1990) Toxic Marine 

Phytoplankton (Proceedings of the 

Fourth International Conference on Toxic 

Marine Phytoplankton, Lund, Sweden, 

26-30 June, 1989). Elsevier, New York. 

No abstract 

Grantli, E., K. Wallstrom, U. Larsson, W. 

Grantli, and R. Elmgren (1990) Nutrient 

limitation of primary production in the 

Baltic Sea area. Ambio 19: 142- 15 1. 

{Baltic; enrichment; primary production; nitrogen; 

phosphorus; nutrient; limitation} 

In the Baltic Sea area, including the Kattegat, the 

external NP loading ratios are generally well above 

the 16:l Redfield ratio for all subareas (the 

Bothnian Bay, The Bothnian Sea, the Baltic proper 

and the Kattegat). During winter, the inorganic NIP 

ratio in surface waters varies. Appreciably higher 

values than the loading ratio are found for the 

northernmost basin, the low-saline Bothnian Bay, 

while lower values than the loading ratio are found 

for the Baltic proper and the Kattegat. Nutrient 

enrichment tests indicate general N limitation in 

the Baltic proper and the Kattegat, although 

stimulation of algal growth after P enrichment has 

been found in the Baltic proper during summer 

blooms of blue-green algae. Blooms of blue-green 

algae are common in the Baltic proper but hardly 

ever occur in the Bothnian Bay and the Kattegat. 

This has been the case for the last century, 

indicating natural summer N limitation. Full-scale 

experimental manipulation of the external NIP 

loading ratio has been carried out in the 

Himmerfjiirdan basin, south of Stockholm. Results 

suggest nitrogen as the most limiting nutrient in 

coastal areas of Baltic proper, uninfluenced by direct 

nutrient discharges. The knowledge of the effects of 

altered external nutrient supplies for nutrient 

limitation in the Baltic Sea system as a whole is 

too limited to allow for reliable predictions. 

However, the Baltic Sea may have developed 

towards a more pronounced N limitation due to a 

twofold historic increase in P supply relative to N 

supply. At present, the situation may be reversed as 

N supply is probably increasing more rapidly than 

P supply. Management of the Baltic Sea area 

cannot be based on removal of either N or P in 

sewage, but must take both elements into 

consideration, as well as differences between subbasin and between polluted and offshore areas. 

Grassle, J. F., J. P. Grassle, L. S. Brown-Leger, 

R. F. Petrecca, and N. J. Copley (1985) 

Subtidal macrobenthos of Narragansett 

Bay. Field and mesocosm studies of the 

effects of eutrophication and organic 

input on benthic population. In: J. S. 

Gray and M. E. Christiansen (ed.) Marine Biology 

of Polar Regions and Effects of Stress on Marine 

Organisms. John Wiley & Sons, New York. 

{MERL; benthic; macrofauna; Narragansett Bay; 

organic; eutrophication; population} 

Benthic macrofaunal population in Narragansett 

Bay and replicate mesocosms representative of 

portions of Narragansett Bay respond rapidly to 

variation in food supply. During five years of 

sampling sharp increases in August mortalities 

occur when estimated carbon demands for benthic 

respiration exceed the amount of organic carbon 

reaching the bottom. Rapid recovery following 

these periods of high mortality indicate a very 

resilient community. Benthic population densities 

in mesocosms rapidly increase in response to 

external additions of plant material, and to within 

system increases in productivity from nutrient 

additions. Larval immigration and predation are also 

important in determining population densities in 

the experimental systems but these variables are 

less easy to control and the effects are less obvious. 

Grassle, J. P. and J. F. Grassle (1984) The 

utility of studying the effects of 

pollutants on single species populations 

in benthos of mesocosms and coastal 

ecosystems. In: H. H. White (ed.) Concepts in 

Marine Pollution Measurements. Maryland Sea 

Grant, College Park. 

{polychaete; bivalve; amphipod; nutrients; 

abundance; Mediomastus ambiseta; Nucula 

annulata; Polydora ligni; Ampelisca abdita) 

No abstract 

Gray, J. S. (1979) Pollution-induced 

changes in populations. Philosophical of the 

Transactions of the Royal Society of London, Part 

B 286:545-561. 

{population; community; pollution; organic 

matter; abundance; benthic fauna; strategy; lognormal; indicator} 

The effects of pollution by organic matter, oil or 

industrial waste on marine communities are 

remarkably similar. Diversity values fall, biomass 

and numbers of organism initially rise and then fall 

as the pollution load is increased. Diversity indices 

are, however, insensitive to pollution-induced 

changes and have to be assessed subjectively. 

Departure from a log-normal distribution of 

individuals among species offers a sensitive and 

objective method of assessing perturbation effects 

on communities. 

Under severe pollution stress, the dominant 

species are those which have a flexible life-history 

ranging from direct development to a planktonic 

larva and the ability to undergo short-term genetic 
selection. Species having a somewhat less flexible 

life-history strategy show increased abundance under 

conditions of slight pollution. The increase in 

abundance of seven or eight neither rare nor 

common species, which gives the departure from 

log-normal distribution, is suggested as being the 

most significant and the earliest detectable change 

caused by pollution in a community. Thus the 

presence of a species in a polluted area may be 

more a question of life-history strategy than the 

tolerance of adverse environmental conditions. If 

this hypothesis is correct, considerable doubt must 

be placed on the ecological relevance of data from 

toxicity tests. 

Gray, J. S. (1992) Eutrophication in the 

sea. In: G. Colombo, I. Ferrari, V. U. 

Ceccherelli and R. Rossi (ed.) Marine 

Eutrophication and Population Dynamics. Olsen & 

Olsen, Fredensborg. 

{review; benthic communities; general model; 

remedial action; research needs) 

Eutrophication begins with incresed growth rates 

of a range of organisms from phytoplankton and 

macroalgae to benthic animals and fish. This is 

followed by a change in the species composition of 

communities and then by a reduction in number of 

species as oxygen concentrationns of water begin to 

fall. Finally, anoxia ensues and few organisms 

survive and only bacterial mats remain. 

Most data show only correlative relationships 

between nutrient input and effects on biological 

organisms. For Example, there is no established 

relationship between nutrient inputs and primary 

production; nor has a quantative relationship 

between organic enrichment and changes in benthic 

communities been established. 

The Vollenweider nutrient loading concept has 

been widely applied to the management of 

freshwater lakes. It is shown that the parameters of 

the model cannot be estimated in marine contexts. 

In the marine environment the most widely 

remedial action for eutrophication is for areas that 

are suspected to be affected to prepare budgets of 

external nutrient loadings and then to find costeffective ways of reducing the amounts of nutrients. 

Yet examples from Norway show that such budgets 

can be widely inaccurate and costly political 

decisions can be taken based on extremly poor data. 

In a research context, for benthic communities 

subject to eutrophication, two models are proposed; 

one for responses to oxygen saturation and the 

other, a more speculative one for changes in C?N 

in sediments. It is suggested that, if verified, the 

latter will allow quantative dose-response 

predictions to be made, rather than the present stateof-the-art of simply simply making correlations 

between nutrient loads and various properties of 

benthic communities. 

Gray, J. S. and E. Paasche (1988) On marine 

eutrophication. Marine Pollution Bulletin 

19:349-350. 

{coast; review) 

No abstract 

Griggs, G. B. and T. S. Hopkins (1976) The 

delineation and growth of a sludge field. 

Water Research 10:501 to 506. 

{sewage; organic carbon; methane; oxygen; 

sediment; anoxic} 

The color, odor, organic carbon, and acoustic 

reflection properties of bottom sediment can be 

used to delineate the area affected by the Athens 

sewage outfall. Both color and organic carbon 

content clearly define the region of high organic 

buildup around the discharge point. Maximum 

organic carbon values of over 6% were determined 

compared to normal regional values of less than 

0.5%. The bottom area covered by black anaerobic 

sediments has increased 7 fold to nearly 9 km2 in 

the past 2.5 yr. A smaller core area around the 

outfall can be easily distinguished by its strong 

seismic reflection characteristics believed to 

coincide with the production of methane gas. 

Haas, L. W., S. J. Hastings, and K. L. Webb 

( 1 98 1 ) Phytoplankton response to a 

stratification-mixing cycle in the York 

River estuary during late summer. In: B. 

J. Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{phytoplankton; hydrographic dynamics; nuisance 

bloom} 

As part of a larger multidisciplinary study of the 

lower York River estuary, phytoplankton response 

to a tidally related cycle of stratificationdestratification was examined during August 1978. 

A "red water bloom" dominated by the 

dinoflagellate Cocchlodinium heterolobatum' 

initially observed in the lower York River 

coincident with the spring tide-induced water 

column destratifiction event. It is proposed that the 

dinoflagellates initiating the red tide were advected 

into the estuary in deep water during the preceding 

period of stratification or were derived from cysts in 

the sediments and that destratification provided 

access to the surface waters. The extent of the red 

water increased during the ensuing restratified period 

in the York River, and several lines of evidence 

indicated that C. heterolobatum migrated diurnally 

between ammonium enriched waters below the 

halocline (8-10 m) and the relatively nutrient-poor 

surface waters. Other estuarine systems in which 

phytoplankton blooms associated with alternating 

periods of stratification-destratification have been 

observed are noted. The results illustrate the close 

relationship between phytoplankton and 

hydrographic dynamics in this estuarine system and 

emphasize the necessity to include the study of 
hydrographic processes in the study of 

phytoplankton dynamics. 

Hagmeier, E. (1978) Variations in 

phytoplankton near Helgoland. Rappots et 

Procds-verbaux des Rbunions 

Counseil International pour L'Exploration de la 

Mer 172:361-363. 

{phytoplankton; abundance; nutrients; phosphate) 

A summary is given of the observations on 

Helgoland Roads between 1962 and 1974. There 

was a tendency of phytoplankton stocks to increase 

during this period, although this was not 

statistically significant. The increase was probably 

related to the significantly rising amounts of 

nutrients available. 

Hallegraeff, G. M. (1993) A review of 

harmful algal blooms and their apparent 

global increase. Phycologia 32:79-99. 

{red tides; nuisance blooms; review) 

no abstract 

Hallegraeff, M. and J. L. Maclean (ed.) (1989) 

Biology Epidemology and Management of 

Pyrodinium Red Tides (Proceedings of 

the Management and Training Workshop, 

Bandar Seri Begawan, Brunei Darussalam, 

Philippines, 23-30 May, 1989). Fisheries 

Department, Ministry of Development, Manila. 

No abstract 

Han, M.-S., K. Furuya, and T. Nemoto (1992) 

Species-specific productivity of 

Skeletonema costatum 

(Bacillariophyceae) in the inner part of 

Tokyo Bay. Marine Ecology Progress Series 

79:267-273. 

{phytoplankton; productivity; Skeletonema; Tokyo 

Bay; photosynthetic rate) 

The composition of red tides in Tokyo Bay varied 

with season; those during summer 1986 and 1987 

were comprised almost entirely of Skeletonema 

costatum. Primary productivity by S. costatum 

ranged from 5.2 to 70.4 % of the total productivity; 

on an annual basis, its contribution was 18.8 %, as 

revealed by species-specific photosynthetic rate 

(SSP), determined by the micromanipulation of 

4~-labelled cells under simulated in situ 

conditions. SSP of S. costatum normalized with 

cell volume, and indicator of growth activity, 

showed temporal variations as the species 

composition of the red tides changed. The volumespecific SSP was high in the initial phase of the 

bloom. However, the high volume-specific SSP 

was rather short-lived. 

Hansson, S. and L. G. Rudstam (1990) 

Eutrophication and Baltic Sea fish 

communities. Ambio 19: 123-125. 

{fish; Ba1tic;increase; decrease) 

The discharge of nutrients to the Baltic Sea has 

increased considerably during the 20th century. 

There are several indications that this 

eutrophication has affected the fish community. 

Changes in species composition has been reported 

from coastal areas, e.g. local increases in the 

abundance of pikeperch and decreased in whitefish. 

Total fish catches, dominated by herring, sprat, and 

cod, have increased tenfold in the past fifty years 

and doubled in the last twenty-five years. This 

increase is mainly due to intensified fishing, but 

possibly also to eutrophication and decreased fish 

predation by seals. 

Hardy, J. T. and Z. Zubayli (1976) 

Phytoplankton standing crop and sewage 

nutrient enrichment along the central 

coast of Lebanon. Environmental Pollution 

11: 195-202. 

{phytoplankton; Lebanon; nutrients; chlorophyll a; 

sewage) 

Surface nearshore water samples from seven 

stations along the central coast of Lebanon were 

analyzed at different seasons for concentrations for 

nitrate, nitrite, ammonia, phosphate and 

chlorophyll-a standing crop. 

Concentrations of nutrients and chlorophyll-a are 

higher within 10 km along coastal zone south of 

Beirut and increase with decreasing distance from 

two major untreated sewage outfall. In some cases, 

chlorophyll-a standing crop near outfalls (e.g. 2.18 

mg m-3) are about 10 times greater than unenriched 

stations further south or than values reported from 

elsewhere in the eastern Mediterranean. Near 

outfalls, concentrations of phosphate, ammonia and 

chlorophyll-a tend to be greatest in summer, while 

nitrate is greatest in winter. 

Preliminary measurements by carbon-14 

assimilation indicate that the enriched coastal 

region has a higher rate of primary production than 

the unenriched area more than 10 km south of 

Beriut. 

Harlin, M. M. and B. Throne-Miller (1981) 

Nutrient enrichment of seagrass beds in a 

Rhode Island coastal lagoon. Ma r i n e 

Biology 65:221-229. 

{lagoon; seagrass; vegetation; algae; seaweeds; 

nutrients; growth) 

Seagrass and algal beds showed a variety of 

responses when the water column was treated with 

low level additions of ammonium, nitrate and 

phosphate. The nutrients were added separately to 3 

uniform seagrass beds of a temperate coastal lagoon 

during 1979 and 1980. (1) Ammonium caused the 

production of dense mats of free-floating green 

algae Enteromorpha plumosa and Ulva lactuca. It 

also stimulated growth in both the leaf and rootrhizome fractions of Zostera marina. This growth 
response in Z. marina was greater in the area where 

current reached 12 cm s-l than in the area with 

little or no current. The concentration of nitrogen 

in the tissue did not change. In contrast, where 

current was lacking, Z. marina growth increase 

with ammonium was small, but the concentration 

of nitrogen in the tissue doubled over that in 

control plots. The growth of Ruppia maritirna was 

inversely related to the growth of green algae in the 

same plots. The red alga Gracilaria tilvahiae did not 

grow better in ammonium, but its tissue reddened. 

(2) Nitrate additions enhanced the growth of the 

green seaweeds Enteromorpha spp. and U. lactuca, 

but not Z. marina or R. maritima. G. tilvahiae, 

when fertilized in isolation from other plants, 

showed a marginal response to this nutrient, and 

the tissue always reddened. (3) Phosphate enhanced 

growth in Z. marina and R. maritima exposed to 

moderate current. G. tilvahiae growing alone 

showed a small growth response to phosphate. The 

phosphate made no difference in the growth of the 

green seaweeds. (4) None of the nutrient 

supplements noticeably altered the species 

composition of either epiphytic or planktonic algae 

associated with the beds, although we did detect 

small increases in their numbers. The rapid and 

dense growth of green algae in nitrogen-enriched 

water probably limited growth of adjacent 

seagrasses and red algae. Because these seaweeds did 

not use the phosphate, it became available to other 

plant components. The overall floral response to 

nutrient addition in seagrass communities depends, 

therefore, upon the particular nutrient supplied, the 

ability of alterate species in the area to compete for 

that nutrient and the velocity of current in the 

specific area. 

Hartig, J. H., J. F. Kitchell, D. Scavia, and S. B. 

Brandt (1991) Rehabilitation of Lake 

Ontario: The role of nutrient reduction 

and food wed dynamics. Canadian Journal of 

Fisheries and Aquatic Science 48: 1574- 1580. 

{Lake Ontario; nutrient; food web; ecosystem} 

The Laurentian Lakes have a complex history of 

changes due to eutrophication, invasion of exotic 

species, and fisheries and phosphorus management 

practices. Remedial actions have reduced nutrient 

loadings and enhanced the role of food web 

interactions in improving water quality. Workshops 

sponsored through the United States - Canada 

International Joint Commission have addressed the 

relative importance of nutrient abatement and/or 

food web manipulation in affecting water quality 

trends. Both controls have combined to enhance 

water clarity in Lake Michigan. Lake Ontario has 

already exhibited the effects of nutrient controls and 

may be on the verge of manifesting food web 

controls. Research and monitoring 

recommendations to elucidate the effects of nutrient 

and food web controls include the following: (1) 

water quality and fisheries agencies must coordinate 

monitoring activities, standardize techniques, and 

establish and maintain long-term data sets to 

evaluate the effects of water quality and fisheries 

programs separately and together; (2) controlled, 

mesoscale, whole system experiments should be 

performed to quantify rates (e.g. growth, predation, 

etc.) of food web interactions; and (3) the scientific 

community should promote research which 

quantifies the impact of changes in food web 

dynamics on changes in toxic substance levels in 

Great Lakes fishes. 

Heinanen, A. P. (1991) Bacterial numbers, 

biomass and productivity in the Baltic 

Sea: A cruise study. Marine Ecology Progress 

Series 70:283-290. 

{bacteria; abundance; biomass; Baltic; 

productivity} 

Bacterial numbers, biomass and productivity were 

studied during 2 cruises in the Baltic Proper, Gulf 

of Finland and the Bothanian Sea. Bacterial 

population in the open sea area had longer turnover 

times and lower productivity than in coastal areas, 

comparable to those found in the open ocean, but 

their biomass and abundance were as high as in the 

coastal areas of the Baltic Sea. High bacterial 

productivity in the aphotic water layer in the Gulf 

of Finland is suggested to indicate eutrophication of 

this area. 

Herbland, A., A. L. Bouteiller, and P. Raimbault 

(1987) Does the nutrient enrichment of 

the equatorial upwelling influence the 

size structure of phytoplankton in the 

Atlantic Ocean? Oceanologica Acta Special 

Volume No. 6: 115- 120. 

{nutrients; phytoplankton; size structure; 

upwelling) 

A study of the size structure of chIorophy1l a 

(Chla) covering the major part of the equatorial 

Atlantic Ocean from 5-N to 54 leads to the 

conclusion that seasonal nutrient enrichment in the 

open Eastern equatorial Atlantic does not drastically 

affect the size distribution of the primary producers: 

90% of the total Chla is everywhere contained in 

the <IOum fraction on the average. In the coastal 

upwelling near Dakar, this percentage is less than 

60%, for the same range of Chla concentrations. In 

the equatorial region the percentage of <I um Chla 

is the same in the deep chlorophyll maximum 

located at the top of the nitracline during the warm 

season and in the subsuperficial maximum of the 

upwelling. Therefore, from an ecological point of 

view, the term "upwelling" is misleading: the 

seasonal equatorial upwelling seems to be nothing 

other than the movement towards the surface of the 

deep chlorophyll maximum, with no appreciable 

increase of its value and some slight modification 

of its trophic organization. 
Herdendorf, C. E. (1986) Rebirth of Lake 

Erie: Recovering from phosphorus 

enrichment. Ohio Sea Grant, Ohio Sea Grant 

Series, OHSU-RS-5 1. 

{lake; Erie; phosphorus; recovery; algae; 

phytoplankton; oxygen; benthos; fish) 

No abstract 

Hesse, K., U. Hentschke, and U. Brockman (1992) 

A synoptic study of nutrient and 

phytoplankton characteristics in the 

German Wadden Sea with respect to 

coastal eutrophication. In: G. Colombo, I. 

Ferrari, V. U. Ceccherelli and R. Rossi (ed.) 

Marine Eutrophication and Population Dynamics. 

Olsen & Olsen, Fredensborg. 

{phytoplankton; nutrients; Wadden Sea) 

For the first time seasonal patterns of nutrient and 

phytoplankton distribution were evaluated in an 

interdisciplinary synoptic study covering the whole 

area (2,500 krn2) of the northern German Wadden 

Sea. Besides the large impact of the river Elbe, 

nutrient patterns revealed the importance of local 

sewage plumes originating from agricultural 

drainage and coastal purification plants. N to P 

ratios were relatively high over all the area in the 

winter and spring, indicating potential phosphorus 

limitation for phytoplankton growth. During the 

spring bloom of Phaeocystis globosa, which 

culminated in biomass concentrationsof 0.5 mg C 

per liter, phosphorus levels decreased to limiting 

concentrations in some of the outer regions of the 

Wadden Sea. In general, however, nutrient stocks 

were never exhausted, emphasizing the importance 

of turbidity as a major factor controlling primary 

production in this area. 

In summer high levels of inorganic phosphate and 

low NIP ratios were observed, reflecting rapid 

remineralization of organic phosphorus 

constituents. It is suggested that the bulk of 

inorganic phosphate in summer results from the 

breakdown of allochthonous phytoplankton blooms 

being advected into the region from sites of intense 

growth such as the frontal region of the Ebe river 

plume and semi-enclosed coastal basins. Red tides 

of decomposing Noctiluca scintillans were observed 

in the Wadden Sea and massive blooms of 

Glenodinium foliaceum in the lagoons, amounting 

to concentrations of 13 mg C per liter and 19 mg C 

per liter, respectively. Mass occurrences of toxic 

forms were not recorded, although several species of 

Dinophysis as well as Alexandrium sp. occurred in 

low cell numbers in spring and summer. 

Allochthonus POC input was estimated to be 

approximately 85 g C per m2 per year, equalling an 

excess remineralization of about 13 tons of 

inorganic phosphate per day for the whole area. 

Owing to inadequate light conditions in the Wadden 

Sea water, it is assumed that this high amount of 

phosphate cannot be recycled in the Wadden Sea 

itself, but is rather transported out of the system, 

thereby enhancing again bloom formation at the 

frontal region of the Elbe River. 

Hesthagen, I. H. (1977) Migrations, 

breeding, and growth in Pomatoschistus 

m in itu s (Pallas) (Pisces, Gobiidae) in 

Oslofjorden, Norway. Sarsia 63: 17-26. 

{growth; sand goby; Pomatoschistus; Oslofjord; 

length) 

Aspects of the biology of the sand goby were 

studied over a period of 21 months. The goby is 

common in the littoral from April (at temperatures 

above 3°C) to July, when breeding takes place, and 

from September to December, when the somatic 

growth occurs. At temperatures above 19°C and 

below 4-5°C the goby is rare in the littoral. 

The growth is very rapid, the 0-group fish 

reaching almost the length of the 1-group fish 

during their first growing season. Growth is faster 

than in other populations from simirar latitudes, 

suggested to be due to eutrophication. Spawning 

occurs even in less than 1 m depth. There is an 

indication that light intensity, together with 

temperature, is important for breeding to start. 

The food consists of any organism of appropriate 

size and texture. Polychaetes are a conspicuous 

component. Feeding occurs intensively all through 

the stay in the littoral, but its lower during the 

breeding season. Various features of the biology of 

the sand goby in Oslofjorden appear to be simple 

and more clearcut when compared to other 

populations studied, probably due to the stable and 

predictable environmental conditions. 

Hinga, K. R. (1990) Alteration of 

phosphorus dynamics during experimental 

eutrophication of enclosed marine 

ecosystems. Marine Pollution Bulletin 21:275- 

280. 

{phosphorus; MERL) 

A 28 month eutrophication experiment was 

conducted in marine mesocosms at Marine 

Ecosystems Research Laboratory of the University 

of Rhode Island. Each mesocosm contained 13 m3 

of seawater and a layer of benthic sediments 

transferred from adjacent Narragansett Bay. 

Nitrogen, phosphorus, and silica were added daily to 

the mesocosms. 

The paper examines net exchange of phosphorus 

between benthic sediments and water column during 

the experiment. At low loading rates the regular 

annual pattern of phosphate concentrations is still 

evident but the amplitude of the pattern is 

magnified. At higher loading rates the annual 

pattern is lost and the effectiveness of the sediments 

to act as a 'buffer' to water column concentrations 

is reduced. In some cases the nutrient loading 

caused a release of phosphorus from the sediments. 
Hinga, K. R. (1992) Co-occurrence of 

dinoflagellate blooms and high pH in 

marine enclosures. Marine Ecology Progress 

Series 86: 18 1- 187. 

{dinoflagellates; diatoms; blooms; pH; 

phytoplankton) 

High abundances of dinoflagellates in mixed 

phytoplankton populations in marine enclosures 

were strongly correlated with high pH during 23 

enclosure-years of weekly samples. Diatom blooms 

were not similarly correlated with high pH. The 

correlation with high pH was not the result of 

dinoflagellate blooms themselves drawing down the 

C02 and driving up the seawater pH. Examination 

of individual blooms of >500 cells ml-l indicates 

that dinoflagellate cell counts increased only after 

the pH was driven high (i.e. >8.5). High pH 

occurred either by natural processes (diatom 

blooms) or, in one case, by an artificial 

manipulation of the pH in the enclosure. There 

were 9 periods in which the seawater pH exceeded 

8.5. Dinoflagellate blooms occurred during 7 of 

those events. A high pH affinity for dinoflagellates 

could help explain reported successional sequences 

of diatom blooms followed by dinoflagellate 

blooms and the association of dinoflagellate 

blooms with eutrophication. Seawater pH should 

probably be included with other environmental 

factors in studies of the mechanisms that control 

the occurrence of field dinoflagellate blooms. 

Ho, Y. B. (1987) Ulva Lactuca 

(Chlorophyta, Ulvales) in Hong Kong 

intertidal waters - Its nitrogen and 

phosphorus contents and its use as a 

bioindicator of eutrophication. Asian 

Marine Biology 4:97-102. 

{green algae; Ulva; sewage; nutrients; tissue; 

nitrogen; phosphorus; bioindicator; Hong Kong) 

The cosmopolitan green alga Ulva lactuca L. is 

often found growing in areas contaminated by 

domestic sewage thus reflecting its ability to thrive 

under such conditions. In order to study the effects 

of eutrophic waters on the nutrient content of the 

alga and its potential as an indicator species, both 

seawater and Ulva samples were collected from 10 

rural and 11 urban intertidal sites around the Hong 

Kong Island over a period of two years. Analysis of 

the water samples showed that the mean dissolved 

inorganic nitrogen (NO3 and NH4) and phosphate 

(PO4) levels in the rural sites were respectively 

49.7 ug N 1-I and 6.7 ug P 1-l. Water samples 

from the urban sites contained very high mean 

levels of ammonia (222.4 ug N 1-l) and phosphate 

(45.3 ug P 1-l) reflecting the extent of sewage 

contamination. The amount of tissue nitrogen and 

phosphorus in Ulva from urban areas were 

respectively 71 and 93% more than those in rural 

sites. Close correlation was found between the 

logarithmic concentration of seawater inorganic 

nitrogen and that of tissue nitrogen (r = 0.949). 

This also applied to seawater phosphate and tissue 

phosphorus (r = 0.927). It is conclude that Ulva is 

a good bioindicator of seawater eutrophication. 

Hobbie, J. E. and J. J. Cole (1984) Response 

of a detrital foodweb to eutrophication. 

Bulletin of Marine Science 35:357-363. 

{bacteria; abundance; activity) 

The response off planktonic bacteria and microprotozoa to an experimental eutrophication gradient 

in coastal marine waters was investigated. Bacterial 

and microflagellate numbers, as well as glutamic 

acid mineralization and H- t h midin e 

incorporation, were measured in a series of 13 m3 

mesocosms for 6 months. The systems received a 

daily addition of sea water (3.7% of total) and 

nutrients so that the nutrients added were 0, 2, 4, 

16 and 32 times the anthropogenic nitrogen, 

phosphorus and silica loading of Narragansett Bay, 

Rhode Island. 

Both the biomass and primary production of the 

planktonic algae increased in direct response to 

nutrient loading through the 16X treatment; the 

response in the 32X treatment equaled that of the 

16X treatment. In general, bacterial numbers and 

growth (calculated from thymidine uptake) were 

higher in the more eutrophic tanks but there were 

lags in the microbial response to the peaks and 

valleys of the algal growth. These lags caused poor 

correlation between instantaneous measurements of 

microbial and algal parameters such as numbers of 

bacteria and algal production. However, when daily 

production was summed for the entire 6 months of 

the study there was excellent correlation between 

the total algal production and the total bacterial 

production (~0.89) for six tanks. It is difficult to 

convert from 3~-thymidine incorporation to 

absolute quantities of bacterial production. We used 

a conservative conversion factor derived from 

laboratory values for the thtmidine content of 

bacterial cells to calculate bacterial production. 

Bacterial production (respiration plus growth) was 

nearly a constant fraction of planktonic primary 

production at all levels of eutrophication and 

averaged 31%. Our results are in good agreement 

with other estimates of the carbon flux through the 

detrital component of pelagic ecosystems. 

Hodgkiss, I. J. and B. S. S. Chan (1987) 

Phytoplankton dynamics in Tolo 

Harbour. Asian Marine Biology 4: 103-1 12. 

{Tolo harbour; phytoplankton; density; abundance; 

composition; nutrients; nitrogen; phosphorus; red 

tide; diversity) 

An intensive study of the Tolo Harbour 

phytoplankton was carried out between January 

1983 and February 1985. The results presented in 
this paper refer to the dynamics of the total 

phytoplankton population at two stations, one in 

the inner harbour and the other in the outer harbour, 

for comparative purposes. 

Results are presented in terms of the total 

phytoplankton cell density in the entire water 

column at each station; the abundance of the 5 

major species of Bacillariophyceae and Dinophyceae 

at each station; and the relative contributions of the 

6 classes of phytoplankton at each station in 1983 

and 1984. 

Increases in total phytoplankton standing crop and 

changes in the percentage composition of the 

component species are related to increasing nutrient 

levels in Tolo Harbour. The increased contribution 

of the Dinophyceae (also reflected in the increase in 

red tide incidents) is particularly stressed, because of 

the implications which such algal blooms have for 

the Tolo Harbour ecosystem. 

Hognestad, P. T. (1987) Assessment of the 

environmental conditions in the 

Skagerrak and Kattegat. ICES, ICES 

Cooperative Research Report, {nitrogen ; 

phosphorus; Kattegat; Skagerrak; priamry 

production; oxygen) 

No abstract 

Hosie, D. A., R. F. Uglow, L. Hagerman, T. 

Sondergaard, and W. Weile (1991) Some effects 

of hypoxia and medium ammonia 

enrichment on efflux rates and circulating 

levels of ammonia in N ep h r op s 

norvegicus. Marine Biology 110:273-279. 

{hypoxia; ammonia; Kattegat; Skagerrak; decapod; 

Nephrops norvegicus) 

Eutrophication has been reported for autumn 

months in regions of the KattegatISkagerrak, 

causing stress to bottom-living organisms. The 

present studies, undertaken in April (1989), 

investigated the effects of hypoxia and high 

ammonia levels in the burrowing decapod Nephrops 

norvegicus (L.). The net ammonia efflux rates and 

circulating ammonia levels at 6 and 12 "C, at 

normoxia [partial] pressure of 02 in the water 

(torr), Pw02 = 155 ton)] and hypoxia Pw02 = 24 

tom) in normal seawater and ammonia-enriched (300 

umol ammonia 1-l) seawater were examined. The 

hourly weight-specific efflux rates were very 

variable and in all groups included some individuals 

which showed periods of no net efflux, or even a 

net uptake of ammonia. At each temperature, net 

efflux-rate differences due to treatments were not 

significant (P>0.05; ANOVA, in all cases) and 

only the differences between the net efflux rates of 

the normoxic groups were significantly affected by 

temperature (P<0.05; ANOVA). Circulating 

ammonia levels were also variable, and at 6°C the 

ammonia-enriched groups had sigilificantly higher 

weight-specific blood ammonia content values than 

the normoxic group (P<0.05 in both cases). A net 

uptake of ammonia occurred in ammonia-enriched 

conditions - probably along a reversed NH4+ 

gradient, as downhill pNH3 gradients were 

maintained in all groups - and may represent the 

only means by which some brachial efflux of 

ammonia could proceed. 

Howells, G. P. (1972) The estuary of the 

Hudson River, U.S.A. Proceedings of the 

Royal Society of London, Part B 180:521-534. 

{Hudson River estuary; phytoplankton; nutrients; 

bloom) 

The Hudson River is a major waterway of the 

eastern seaboard of the United States. Its two major 

tributaries, the Mohawk and Hudson, combine to 

form the long narrow estuary of the Lower Hudson. 

The area is populated and industrially developed, 

and it is predicted that both domestic and industrial 

demands for water will increase, as well as the 

demand for waste disposal and for the maintanence 

of its recreational resources. 

Recent studies of the ecology of the Estuary, 

survey of the levels of pollutants in water, and their 

distribution between water, muds and the biota 

indicate that water quality may need control if 

problems of maintenance of fisheries, 

eutrophication, and drinking water quality are to be 

avoided. There is clearly a case for relevant research 

into the processes and pathways of pollutants 

within this aquatic ecosystem, and for the study of 

effects of pollutants on biological communities in 

relation to concentration and time of exposure. 

Estuaries are areas of great natural variations in 

environmental conditions and their flora and fauna 

is selected and restricted in its variety by the need to 

tolerate or regulate in the variable conditions. For 

this reason, the capacity of estuaries to accept 

pollutants which enhance natural variations is 

relatively great. The limit of environmental 

acceptance need to be determined by achieving the 

best reconciliation between industrial development 

and the maintenance of amenity. 

Hull, S. C. (1987) Macroalgal mats and 

species abundance: A field experiment. 

Estuarine, Coastal and ShelfScience 25519-532. 

{macroalgae; mats; benthos; infauna; abundance; 

Ythan estuary) 

A field experiment was carried out whereby the 

density of macroalgae (Enteromorpha spp.) was 

manipulated and the resultant changes in sediment 

infaunal density were monitored. Four densities of 

Enteromorpha spp. were used: 0, 0.3, 1 and 3 kg 

FW m-2, corresponding to control, low-, medium-, 

and high-density plots. The experiment ran from 

May to October 1985 and was sampled on three 

occasions. By July, the density of Corophium 

volutator was reduced at all weed levels when 
compared to control plots, whereas densities of 

Hydrobia ulvae, Macoma balthica, Nereis 

diversicolor, and Capitella capitata, all increased. 

Samples taken in October when the weed mats were 

buried in the sediment showed fewer differences 

than in July. Macoma, Nereis, and Capitella were 

still significantly more abundant at medium and 

high weed densities. Corophuim showed no 

significant treatment effect. There was, however, a 

highly significant difference in population size 

structure for Corophium. Measurements of 

sediment redox potential and silt content under 

medium- and high-density plots revealed rapid 

anoxia with a significant increase in siltation. 

Hungspreugs, M., W. Utoomprkporn, S. 

Dharmvanij, and P. Sompongchaiyakul (1989) 

The present status of the aquatic 

environment of Thailand. Marine Pollution 

Bulletin 20:327-3332. 

{Thailand; bloom; red tide) 

The rapidly increasing population and developing 

agro-industrial activities in Thailand exert 

considerable stress on the aquatic environment. 

Untreated municipal wastes are discharged directly 

or indirectly to canals and rivers, causing high 

BOD5 values and bacterial contamination close to 

populated areas. Improvements in domestic sewage 

treatment are likely to be prohibitively expensive 

in the near term at least. Eutrophication of coastal 

waters is an emerging problem. By contrast, few 

problems have been documented from trace metals 

discharged by industries, and the public health 

threat from seafood contamination does not appear 

to be significant. Uses, of chlorinated hydrocarbon 

pesticides are declining sharply, these being 

replaced by organophosphate compounds which 

degrade rapidly in aquatic environments. Oil 

pollution has not been a problem, although fears of 

a major spill exists. 

Imai, I., S. Itakura, and K. Itoh (1991) Life 

cycle strategies of the red tide causing 

flagellates Chattonella (Raphidophyceae) 

in the Seto Inland Sea. Marine Pollution 

Bulletin 23: 165-170. 

{red tide; flagellates; cyst; life cycle; bloom) 

Chattonella antiqua (Hada) Ono and Chattonella 

marina (Subrahmanyan) Hara et Chihara are the 

most noxious red tide flagellates which cause 

serious damage to fish farming, especially to 

yellowtail culture, in Japanese coastal waters such 

as the Seto Inland Sea during summer. Cysts of 

Chattonella were identified from sediments of the 

Seto Inland Sea. These cysts overwinter in sea 

bottom and play an important role in initiating the 

summer red tides. Most of the cysts adhere to solid 

surfaces such as diatom frustules and sand grains, 

which may aid in keeping the cyst populations 

within seed beds for red tides. Temperature is a 

principal factor affecting the physiology of cysts of 

Chattonella. No cysts germinate at 10 4. 

Optimum temperature range is between 20 and 25 

-C for germination. For maturation (acquisition of 

germinability) of the cysts, low storage temperature 

of 11 WC or below for more than four months is 

essential, whereas no significant maturation is 

observed at 20 -C or more. In freshly collected 

sediments, marked seasonality of germinability was 

confirmed in Suo-Nada. The cysts have 

germinability between spring and early summer. 

The cysts spend a period of spontaneous dormancy 

between autumn and the next spring, and they 

mature during the winter season. The life cycle of 

Chattonella is therefore well adapted to seasonal 

temperature fluctuation in the ~eto-Inland Sea. And 

further, alteration between benthic and planktonic 

stage is presumably unconstrained by virtue of 

shallowness of the Seto Inland Sea. The life cycle 

of Chattonella can be regarded as superior strategies 

in conclusion for the occurrences of the red tides in 

temperate waters such as the Seto Inland Sea. 

Jaworski, N. A. (1981) Sources of nutrients 

and the scale of eutrophication problems 

in estuaries. In: B. J. Neilson and L. E. 

Cronin (ed.) Estuaries and Nutrients. Humana 

Press, Clifton, New Jersey. 

{estuary, freshwater, nutrient, budget, comparison) 

A comprehensive analysis of external sources of 

nutrients is presented including an impact, 

comparison of external loadings, and the resulting 

scale of eutrophication. The major emphasis of the 

analysis is on nitrogen and phosphorus. 

The relative contribution of various external 

sources for five major ecosystems is delineated. 

Discussion of seasonal and long-term trends of 

external sources is presented. The impact of 

external sources on the eutrophication process is 

evaluated. A scale of eutrophication for estuarine 

ecosystem is suggested for comparing the impact of 

nutrient enrichment. 

Comparisons of external nutrient loadings and the 

scale of eutrophication for 13 estuarine and 

freshwater ecosystems are made. A detailed 

comparison of the five estuaries of the Chesapeake 

Bay is presented from which relationships between 

external nutrient loadings (glm21yr) and eutrophic 

conditions are suggested. 

Analysis of the relationship between external 

nutrient loadings, nitrogenlphosphorus ratio, and 

eutrophic state suggests that, for the East Coast 

estuarine ecosystems, if the phosphorus loading is 

1.0 glm21yr or less, excessive eutrophic conditions 

can be prevented. The favorable response of the 

Potomac estuary to phosphorus control 

demonstrates that excessive eutrophic conditions 

may be alleviated with advanced wastewater 

treatment, depending on many factors including 

nutrient loadings. 
Jaworski, N. A. and J. O.Villa (1981) A 

suggested approach for developing 

estuarine water quality criteria for 

management of eutrophication. In: B. J. 

Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{criteria; management; framework) 

A conceptual approach for developing water 

quality criteria for eutrophication management is 

suggested. The three basic components of the 

framework include source ambient relationships, 

effects, and impact analyses. The approach focuses 

on a conceptual method for developing decisionmaking criteria as opposed to the classical water 

quality criteria for a single value of limitation. The 

approach to developing framework of response 

relationships which can be readily incorporated into 

water quality standard-setting processes that include 

environmental considerations and technological and 

economic factors. 

Jingzhong, Z., D. Liping, and Q. Baoping (1985) 

Preliminary studies on eutrophication and 

red tide problems in Bohai Bay. 

Hydrobiologia 127:27-30. 

{red tide; Bohai Bay; nutrients; phytoplantkon; 

community; composition; productivity; sewage; 

trophic state index) 

A study was carried out in 1978-1981 on the 

relationship between eutrophication and the 

occurrence of red tides in Bohai Bay. 

An assessment method using a single parameter 

as well as the multiparameter trophic State Index 

was adopted. DIN, DIP, COD loading and 

concentrations and phytoplankton community 

(species composition, diversity index, standing 

crop, chlorophyll a and primary productivity) were 

used as parameters. The western part of the bay and 

two sewage outfall sites showed the highest values 

of chlorophyll a, nitrate, phosphate, standing crop, 

primary productivity and the greatest population 

instability. Therefore, Bohai Bay, can be regarded as 

mesotrophic, but its estuary eutrophic. 

Anthropogenic factors are mainly responsible for 

its eutrophication. A relationship between 

environmental factors and red tide species was 

found. Noctiluca miliaris, Prorocentrum minimum 

and Skeletonema coastatum have their bloom 

periods from July to September and are the most 

significant species. 

Jochem, F. and B. Babenerd (1989) Naked 

Dictyocha speculum - a new type of 

phytoplankton bloom in the Western 

Baltic. Marine Biology 103:373-379. 

{Dictyoc ha ; phytoplankton; bloom; 

silicoflagellate; Baltic) 

We report on the bloom incidence of a curious 

new type of phytoplankton organism: the naked 

form of the silicoflagellate Dictyocha speculum. 

This special form does not develop the siliceous 

skeletons which normally characterize the genus 

Dictyocha, its growth is apparently independent of 

the availability of dissolved silicate. The first 

massive bloom of this organism was observed in 

the Kiel Bight area, FRG, in May 1983. We 

suggest that the occurrence of this new type of 

bloom is related to changes in nutrient conditions 

due to increased eutrophication. 

Johansson, J. 0. R. and I. R.R. Lewis (1992) 

Recent improvements of water quality 

and biological indicators in Hillsborough 

Bay, a highly impacted subdivision of 

Tampa Bay, Florida, USA. In: R. A. 

Vollenweider, R. Marchetti and R. Viviani (ed.) 

Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{chlorophyll, seagrass, blue-green algae, 

transparency) 

Hillsborough Bay, the eastern uppermost section 

of the Tampa Bay system, is surrounded by a large 

metropolitan complex, supports extensive 

industrial activity, and serves as a major shipping 

port of fertilizer products. The Bay was determined 

to be highly eutrophic during the late 1960's. The 

City of Tampa's primary sewage treatment plant 

and runoff from fertilizer industry activities were 

considered as the major sources of excessive 

nutrient loading. Loadings from both these sources 

have been reduced during the last 10 years, which 

has probably been the leading cause of improved 

water quality parameters such as water clarity, 

dissolved oxygen, and chlorophyll. The 

improvements recorded in these parameters may, in 

part, be related to large biomass reductions of a 

planktonic blue-green alga, which used to dominate 

the fall and early winter phytoplankton population. 

Coincidental with improved water quality, seagrass 

and an attached macro-alga have vegetated shallow 

areas around the bay, which had been barren of 

attached vegetation for several decades. 

Jones, R. A. and G. F. Lee (1986) 

Eutrophication modeling for water 

quality management: An update of the 

Vollenweider OECD model. Water Quality 

Bulletin 11:118-. 

{model; fish; yield; phosphorus; biomass; 

production) 

No abstract 

Jonge, V. N. d. and H. Postma (1974) 

Phosphorus compounds in the Dutch 

Wadden Sea. Netherlands Journal of Sea 

Research 8: 139-153. 

{phosphorus; Dutch Wadden Sea; Rhine) 

During the period of July 1970 to July 1971 

phosphorus concentrations (phosphate, dissolved 
organic phosphorus and particulate phosphorus) 

have been measured over the whole Dutch Wadden 

Sea including the Ems-Dollard area and the northern 

part of the IJsselmeer. 

Phosphate values were in general higher than in 

the North Sea. Highest concentrations occur in 

summer in the inner parts as a result of 

accumulation and mineralization of organic 

material. In this way steep gradients are built up 

between these inner parts and the tidal inlets, with 

maxima of 2.5 ugst PA. In winter differences are 

small and the phosphate distribution is more 

homogeneous. Very low phosphate values were 

only found in spring and summer in the tidal inlet 

between the islands Vlieland and Terschelling where 

phosphorus probably becomes limiting for 

plankton growth. 

Since 1950 phosphate concentrations increased 

three times (0.5 to 1.7 ugat PA average). Over the 

same period the river Rhine shows a similar 

increase. However, particulate phosphorus 

concentrations only doubled in the Wadden Sea (2.4 

to 5.1 ugat PI100 mg carbonate-free suspended 

matter, on the average). It has been assumed that 

the difference between the two factors is due to a 

considerable increase in microbial activity. 

Josefson, A. B. (1987) Large-scale patterns 

of dynamics in subtidal macrozoobenthic 

assemblages in the Skagerrak: Effects of 

a production-related factor. Marine Ecology 

Progress Series 38: 13-23. 

(macrobenthos; assemblage; organic; sediment; 

abundance) 

Macrobenthic infauna at water depths between 10 

and 300 m was monitored over 5 yr in an 80 km 

wide area off the Swedish west coast. Year-to-year 

variations in number of individuals were analyzed 

for total fauna, trophic and reproductive groups and 

dominating species. Results demonstrated 

significant variations in total abundance 

synchronized between stations suggesting the 

importance of some extrinsic factor. Relative 

positions of dominants seemed largely unaltered, 

suggesting a similar response among species to the 

common factor. Much of the variability was 

accounted for by short-lived predatory and surfacedeposit-feeding polychaetes and surface-depositfeeding cumaceans. A comparison of the variability 

between species with planktotrophic and 

lecithotrophic larval development showed no 

difference between the categories, suggesting that 

the common factor operated on the bottom. 

Temperature and salinity data suggest only small 

differences between years. It is argued that the 

major cause of the common variability pattern is a 

factor related to production in the sea such as 

sedimentation of organic matter, which affects 

either of, or a combination of, settlement, somatic 

growth and survival of the bottom. Sediment 

organic matter over 3 yr changed in a way that 

would be predicted from the faunal changes and 

consistent with the hypothesis that sedimentation 

had caused the variation. Increased abundance were 

accompanied by increased organic content. 

Josefson, A. B. and R. Rosenberg (1988) Longterm soft-bottom faunal changes in three 

shallow fjords, West Sweden. Netherlands 

Journal of Sea Research 22: 149-159. 

{benthic; fauna; abundance; biomass; composition; 

diversity) 

Three West Swedish fjords were investigated at 

the same 14 stations in 1976 and 1986, and 8 of 

these had been investigated in the 1920's. The 

stations are situated at 7 to 27 m water depths in 

protected areas without any significant local 

pollution input, but now with organically enriched 

sediments. Comparisons of benthic fauna between 

1976 and 1986 showed the following significant 

reductions: in total mean abundance and biomass 

(excluding some large and rare species); in 

abundance and biomass of molluscs; in abundance 

of suspension feeders and carnivores. Similarity 

indices gave significant differences between 1976 

and 1986. The greatest changes had occurred in the 

two fjords with the most restricted water 

circulation. Although these faunal changes may be 

attributed to several factors, recently increased 

periods of hypoxia in the bottom water, which 

could be a result of large-scale eutrophication, are 

suggested as the main impact on the benthos. 

Justi'c, D. (1987) Long-term eutrophication 

of the Northern Adriatic Sea. Marine 

Pollution Bulletin 18:28 1-284. 

{Adriatic Sea; anoxia; oxygen; primary production; 

nutrients) 

In the Northern Adriatic Sea, the occurrence of 

anoxic events near the bottom has considerably 

increased in frequency during the last 15 years. 

Although it has been suggested that increasing 

nutrient inflow from land-based sources has caused 

the phenomenon, there has been no direct evidence 

for this. An analysis of the oxygen distribution in 

the Northern Adriatic Sea between 191 1 and 1982 

clearly demonstrates that the probability of anoxic 

events near the bottom has increased in time. This 

process is related to the increasing primary 

production near the surface. There is strong 

evidence that the long-term nutrient enrichment of 

the Northern Adriatic Sea forces the above changes. 

Justi'c, D. (1988) Trend in the transparency 

of the Northern Adriatic Sea 1911-1982. 

Marine Pollution Bulletin 19:32-35. 

{transparency; Adriatic; primary production; 

nutrients). 

In the northern Adriatic Sea, the Secchi disk has 

been used in oceanographic studies since 191 1. An 
analysis of the data collected during summers 

between 1911 and 1982 demonstrates that, on the 

average, the Secchi disk depth has decreased in 

time. It is likely that a decrease in light penetration 

has reduced the benthic primary production and thus 

has increased the probability of an occurrence of 

anoxic events near the bottom. The long-term 

nutrient enrichment of the freshwaters discharging 

into the northern Adriatic Sea appears to be the 

main factor which has caused the above changes. 

Justi'c, D. (1991) Hypoxic conditions in 

the northern Adriatic Sea: Historical 

development and ecological 

significance. In: R. V. Tyson and T. H. 

Pearson (ed.) Modern and Ancient Continental 

Shelf Anoxia. The Geological Society, London. 

{oxygen; Adriatic; Po River; nitrogen; phosphorus; 

hydromedusae; meroplankton) 

In the northern Adriatic Sea the occurrence of 

anoxic events and related benthic mortalities has 

considerably increased in frequency during the past 

15 years. This paper summarizes recent findings 

and presents new evidence on the causes and 

consequences of this phenomenon. Trends in the 

oxygen content 1911-1984, trends in the Secchi 

disk depth 191 1-1982 and long-term changes in the 

communities of benthos and meroplankton are 

described. The relative importance of a set of factors 

which create favorable conditions for the 

development of hypoxia is discussed. 

Kaiser, W., A. Irmisch, D. Nehring, F. Georgi, and 

G. Bruel (1990) Ecological investigation in 

the onshore pelagic zone near 

Warnemiinde from March 1985 to March 

1986. Limnologica 20:33-36. 

{nutrients; chlorophyll; pelagic) 

From March 1985 to March 1986 ecological 

investigations were carried out in the pelagic zone 

near Warnemiinde with the aim of analyzing the 

annual variations in some oceanological parameters 

and ascertaining the productivity and eutrophication 

of this region. Main results are as follows: 

- All parameters studied show annual variations, 

which are greatest for temperature, nutrients and 

biological parameters. 

- Short term variations (few days) were strong, 

except in the case of temperature. 

- Close connections exist between the biological 

and chemical parameters. 

- The influence of the discharge from the River 

Warnow on the ecological conditions could not be 

clarified. 

- As comparisons of chlorophyll and seston 

concentrations with those of the Arkona Basin and 

the Liibeck Bay show, the area studied can be 

characterized as a slightly eutrophic region of the 

Mecklenburg Bay. 

Kat, M. (1987) Toxic and non-toxic 

dinoflagellate blooms on the Dutch 

coast. In: T. Okaichi, D. M. Anderson and T. 

Nemoto (ed.) Red Tides: Biology, Environmental 

Science, and Toxicology. Elsevier, New York. 

{bloom; dino; abundance; toxic) 

The water of the Dutch coastal area is a mixture 

of water entering the North Sea in the south 

through the Straits of Dover, fresh water from the 

rivers Rhine, Meuse and Scheldt and British coastal 

water (figure 2.). The seasonal occurrence and yearto-year frequency for the period 1973-1984 will be 

described of 9 dinoflagellates occurring in a 70 km 

wide area along the coast. Special attention has 

been paid to the presence of toxin producing 

dinoflagellates and the results of this program for 

1986 and 1987. 

Kautsky, N., H. Kautsky, U. Kautsky, and M. 

Waern (1986) Decreased depth penetration 

of Fucus vesiculosus (L.) since the 

1940's indicates eutrophication of the 

Baltic Sea. Marine Ecology Progress Series 

28:l-8. 

{light; depth penetration; seaweeds; chlorophyll; 

Baltic) 

Although nutrient inputs to the Baltic Sea have 

increased drastically since the end of the last 

century, there is still little hard biological evidence 

of a general eutrophication of the Baltic Sea outside 

locally polluted areas. A revisit after 40 yr to some 

well-documented divicg stations in the outer 

archipelago of the Aland Sea gave us an 

opportunity to register any changes in benthic 

vegetation that could be linked to eutrophication. 

By mapping the vertical distribution in coverage of 

bladder wrack Fucus vesiculosus (L.) at 11 stations 

we observed that the lower limit of this alga had 

moved upwards at 10 stations from maximally 11.5 

m in 1943144 to 8.5 m in 1984. Also the depth of 

maximum development had withdrawn from 5 to 6 

m in 1943144 to 3 to 4 m in 1984, while coverage 

at these depths was about the same: 58% and 51 % 

respectively. The deeper specimens today at 8.5 m 

had the same dwarfed appearance as those found at 

11.5 m in the 1940's; at that time growth at 8.5 m 

was luxuriant. During both studies the decrease in 

F.vesiculosus coverage with depth towards the 

lower limit could be approximately fitted to an 

exponentially decreasing light attenuation curve. 

Since the structure of Baltic hardbottom 

communities is almost totally governed by abiotic 

factors, the changes in depth penetration are 

probably caused by decreased transparency of the 

water column due to eutrophication. Results 

indicate that the decreased water transparency arises 

from a 40 to 50% increase in summer values of 

chlorophyll a and nutrients in the offshore surface 

water of the Baltic Sea since the 1940's. 
Keller, A. A. (1988) An empirical model of 

primary productivity (l C) using 

mesocosm data along a nutrient gradient. 

Journal of Plankton Research 10:813-834. 

{primary production; MERL) 

The two parameters of the hyperbolic tangent 

equation, Prn and a, were estimated from in situ 

vertical profiles of primary production using 

mesocosm data along a nutrient gradient. The 

parameters, derived from 4-h (around noon) 14c 

incubations, were used together with the 

photosynthesis-light curve and hourly solar 

radiation data to calculate daily primary production 

rates (PA). Approximately 40% of the daily - production occurred in the 4h around noon. 

Considering parameter uncertainty, there was no 

indication of an increase in variation in production 

with increased nutrient loading, nor did biomassspecific P-I parameters increase. Annual production 

ranged from 82 to 901 g~m-2 year-1 and was 

highest in the highest nutrient treatment tank. 

Daily productivity ranged from 0.02 to 9.1 gCm2day-1 and was significantly correlated, in all 

treatments, with a composite parameter Blolk 

(where B is phytoplankton biomass; 10 is daily 

radiation and k is the extinction coefficient). Linear 

regression of Pd against BIdk indicated that much 

of the variability (86%) in productivity was 

explained by light availability and phytoplankton 

biomass. Two approaches for predicting 

productivity were compared: (i) predicting 

production directly from environmental variables 

(i.e. BIdk ) and (ii) predicting the parameter of the 

P-I curve from environmental variables and using 

these to calculate daily production. 

Keller, A. A. (1989) Modeling the effects of 

temperature, light, and nutrients on 

primary productivity: An empirical and a 

mechanistic approach compared. 

Limnology and Oceanography 4332-95. 

{chl a; 14c production; primary productivity; 

MERL) 

The prospect of reliably predicting production 

from environmental variables in heterogeneous 

estuarine environments offers clear logistic 

advantages over measurement with the 14c 

technique, particularly if detailed coverage (in space 

and time) is desired. To that end, two different 

techniques for predicting phytoplankton production 

were tested and compared using data from 

Narragansett Bay, R.I., and experimental estuarine 

ecosystems. Primary productivity was first 

estimated via an empirical technique (principal 

component regression), which incorporated 

temperature, light, and nutrients as independent 

variables. This approach was compared with a 

traditional mechanistic formulation which included 

the same environmental variables as the major 

limiting factors. Both models provided good fits to 

the data (explaining 77-82% of the total variance). 

The predictive ability of the mechanistic model was 

significantly greater (p<0.05) than the principal 

component regression when tested on two 

independent data sets. Predictive ability between the 

two models was statistically compared by F - tests 

on the residual variability from each method. 

Keller, A. A., P. H. Doering, S. P. Kelly, and B. 

K. Sullivan (1990) Growth of juvenile 

Atlantic menhaden, Brevoortia tyrannus 

(Pisces: Clupeidae) in MERL 

mesocosms: Effects of eutrophication. 

Lirnnology and Oceanography 35: 109-122. 

{secondary production; growth; Atlantic menhaden; 

nutrient; MERL) 

Growth and mortality of recently hatched Atlantic 

menhaden (Brevoortia tyrannus ) were examined for 

7 months under two levels of nutrient enrichment 

(with and without silicate enhancement) and control 

conditions in experimental marine mesocosms (13 

m3). Results from six plankton-dominated systems 

confirmed the often hypothesized but rarely 

demonstrated link between nutrient loading, food 

quantity, and growth in marine fish. Weekly 

abundance and mortality were not significantly 

different (P 2 0.05) between controls and treatment 

mesocosms. On the other hand, von Bertalanffy's 

growth coefficients and mean asymptotic sizes of 

juvenile Atlantic menhaden were significantly 

different (P < 0.05) and positively correlated (r2 = 

0.98) with mean food availability (measured as the 

sum of phytoplankton plus zooplankton carbon), 

which in turn was significantly (P < 0.05) and 

positively correlated (r2 = 0.94) with nutrient 

loading (measured as the input of N+P+Si). 

Keller, A. A. and R. L. Rice (1989) Effects of 

nutrient enrichment on natural 

populations of the brown tide 

phytoplankton A u r e o c o c c u s 

anophagefferens (Chrysophyceae). Journal 

of Phycology 25:636-646. 

{brown tide bloom; nutrient; picoalgae; MERL) 

The brown tide picoalgae A u reococcus 

anophagefSerens Hargraves et Sieburth was present 

in approximately equal numbers in 12 large scale 

(13000 L) mesocosms at the start of a nutrient 

addition experiment in June 1985. Increases in 

abundance in untreated systems mimicked the 

pattern of bloom development in Narragansett Bay, 

Rhode Island, the seawater source for the 

experiment. Aureococcus increased to maximal 

values of 2.6 X 109cells L-~ and persisted at high 

numbers (>lo8 cells L-l) for 7-8 weeks. In nutrient 

addition tanks, the picoalgae bloomed briefly (1-3 

weeks) but rapidly declined to the usual level (-lo7 
cells L-~ for eukaryotic algae in Narragansett Bay). loading. Average summer fluxes for treatments 

The decline in picoalgae abundance was followed by ranged from about 150 to 1200 umols NHd+ m-2 

an increase in total diatoms in all nutrient treated 

tanks. Mean picoalgae abundance in the mesocosms 

and the bay was significantly (P<0.05) and 

inversely correlated (r=-0.93) with mean 

concentration of dissolved inorganic nitrogen. The 

persistence of the brown tide species in control 

mesocosms and Narragansett Bay appears related to 

its ability to grow at very low concentrations of 

dissolved inorganic nitrogen, levels previously 

shown to limit diatom growth. 

h-l, with highest rates being recorded at the 2 

highest loading levels. Benthic nutrient 

regeneration did not increase in direct proportion to 

loading, but appeared strongly related to net 

primary production rates across the enrichment 

gradient. Comparison, across the experimental 

gradient, of the external nutrient supply rate with 

the benthic feedback of dissolved nutrients to the 

water column indicated that beyond a loading of 

about 2000 to 5000 mmols N m-2 vr-l the water 

Keller, A. A. and R. L. Rice (1990) Variation column nutrient dynamics became dominated by the 

in DCMU-enhanced fluorescence relative external supply. Results suggest that the 

to chlorophyll a : Correlation with the importance of benthic-pelagic interactions to 

brown tide bloom. Journal of Phycology biogeochemical and ecological cycles of coastal 

26:202-205. areas may decrease markedly with nutrient 

{brown tide blooms; fluorescence; nutrients; enrichments which exceed the needs of the 

MERL) autotrophic components of the ecosystem. 

DCMU-enhanced fluorescence and extracted 

chlorophyll a were simultaneously measured in 

Narragansett Bay, Rhode Island and the MERL 

(Marine Ecosystems Research Laboratory) 

mesocosms during the 1985 brown tide bloom. 

Marked differences in the relationship between these 

variables were observed as the phytoplankton 

community shifted from dominance by picoalgae to 

diatoms. The fluorescence to chlorophyll a ration 

was significantly (P<0.05) higher in the 

mesocosms and the bay when the brown tide 

species (Aureococcus anophagefferens Hargraves et 

Sieburth) dominated the phytoplankton community 

compared with other taxa. Although several factors 

could have affected the relationship we believe the 

high ratios are related to the pigment composition 

and 1 or small size of the brown tide organisms. 

Kelly, J. R., V. M. Berounsky, S. W. Nixon, and 

C. A. Oviatt (1985) Benthic-pelagic 

coupling and nutrient cycling across an 

experimental eutrophication gradient. 

Marine Ecology Progress Series 26:207-2 19. 

{nutrient; flux; benthic; pelagic; nitrogen; 

phosphorus; MERL) 

Sediment-water exchange rates of dissoloved 

inorganic nitrogen (NH4+, NO3-, N02-) and 

phosphorus (~0~~3, water column concentrations 

of both dissolved and particulate forms of N and P, 

and net primary production (l 4~) were measured 

during a summer period in large (13 m3) 

experimental mesocosms that had been subjected 

to continuous daily nutrient additions (N, P, Si) for 

over 1 yr. The concentrations of combined 

dissolved inorganic plus particulate nutrient forms 

(N, P) were linearly related to the nutrient input 

rate across a loading range from 596 to 34100 

mmols N m-2 yr-l. Benthic nutrient regeneration 

generally increased, although not uniformly, with 

Kelly, M. and M. Naguib (1984) 

Eutrophication in Coastal Marine Areas 

and Lagoons: A Case Study of 'Lac de 

Tuins'. UNESCO, UNESCO reports in marine 

science, 29. 

{coast; lagoon; review; concept; processes) 

During the period from September 6 through 18, 

1982 a workshop was conducted in Tunis, Tunisia 

to discuss and teach about problems of 

eutrophication in coastal marine areas. The history 

of the concept of eutrophication, as developed for 

north-temperate lakes, was discussed. The major 

processes involved, including nutrient enrichment, 

excess algal growth, oxygen depletion, and the 

microbial processes in anaerobic environments were 

described. Participants described eutrophic 

conditions found in coastal areas of their respective 

countries (all bordering the ~editerranean). The 

eutrophication of Lac de Tuins, perhaps the most 

eutrophic embayments in the world, was discussed 

and presented as a case study. During the workshop 

we realized that the concepts of cultural 

eutrophication as originally described for northtemperate lakes apply imperFectly to the marine 

environment. But the processes in both 

environments are similar, with the addition of large 

quantities of nutrients, excessive algal growth, 

depletion of oxygen with consequent microbial and 

chemical changes, and the death of large numbers of 

animals and complete change of ecosystem 

structure. 

Kemp, W. M., R. R. Twilley, J. C. Stevenson, 

W. R. Boynton, and J. C. Means (1983) The 

decline of submerged vascular plants in 

the upper Chesapeake Bay: Summary of 

results concerning possible causes. 

Marine Technology Society Journal 17:78-89. 
{vascular plants; submerged; Chesapeake Bay; 

vegeation; nutrient; epiphytic; phytoplankton; 

algae; biomass) 

This paper provides a summary and synthesis of 

research conducted to investigate possible causes of 

the decline in abundance of submerged aquatic 

vegetation (SAV) in the upper Chesapeake Bay 

beginning in the late 1960s. Three factors were 

emphasized in this study; runoff of agricultural 

herbicides; erosional inputs of fine-grain sediments; 

nutrient enrichment and associated algal growth. 

Widespread use of herbicides in the estuarine 

watershed occurred contemporaneous with the SAV 

loss; however, extensive sampling of estuarine 

water and sediments during 1980-81 revealed that 

typical bay concentrations of herbicides (primarily 

atrazine) rarely exceeded 2 ppb. On two occasions 

relatively high values (20-45 ppb) were observed 

for brief (2-4 h) periods in a small cove following 

runoff events. Short (2-6 h) and long (4-6 wk) term 

experiments indicated that ephemeral phytoxic 

effects would be expected in response to these 

highest herbicide concentrations followed by rapid 

recovery. However, normal concentrations (4 ppb) 

had little measurable effects on plants. Historical 

increases in turbidity have been documented for 

some bay tributaries since the 1940s. During our 

study light (PAR) attenuated by suspended finegrain sediments contributed more to total turbidity 

in bay shallows (< 1.5 m) than did phytoplankton 

chlorophyll a. Die1 cycles of PAR available in 

SAV beds indicated that plant photosynthesis was 

light-limited for much of the day, and PAR often 

fell below the compensation level (Ic) needed for 

minimal plant growth. Although some SAV 

species exhibited considerable ability to adapt to 

reduce light by such mechanisms as increased 

pigmentation and stem elongation, increased 

turbidity has probably reduced overall depth 

distribution of SAV markedly. Effects of the 

continual increase in nutrient enrichment of the bay 

(documented since 1930) were tested by 

experimentally fertilizing pond mesocosms at 

levels common to the upper estuary. Moderate to 

high nutrient loadings resulted in significant 

increases in growth of epiphytic and planktonic 

algae and decreases in SAV production, as well as 

premature seasonal senescence of fertilized plant 

populations. Direct measurements demonstrated the 

inhibitory effect of epiphytic growth on SAV 

photosynthesis, due largely to light attenuation. 

The results of these various ex~eriments were 

synthesized into an ecosystem simulation model 

which demonstrated the relative potential 

contributions of the 3 factors to SAV declines, 

where nutrients > sediments > herbicides. Other 

factors and mechanisms are also discussed along 

with possible resource management options. 

Kennedy, V. S. (ed.) (1984) The Estuary as a 

Filter. Academic Press, Orlando. 

No abstract 

Kerr, S. R. and R. A. Ryder (1992) Effects of 

cultural eutrophication on coastal marine 

fisheries: a comparative approach. In: R. 

A. Vollenweider, R. Marchetti and R. Viviani (ed.) 

Marine Coastal Eutrophication. Elsevier, 

Amsterdam, London, New York, Tokyo. 

{fish yield) 

Several generalities are common to the effects of 

cultural eutrhopication on fisheries in a variety of 

marine and freshwater ecosystems. Foremost is the 

absence of definitive diagnostic features to permit 

discrimination between direct eutrophication effects, 

and those engendered by other factors, often 

associated with anthropogenic change. It is also 

clear that euthrophication may profoundly modify 

the structure, function and species composition of 

fish production systems, with attendant 

consequences for both ecosystem function and the 

economic value of the yield. Eutrophication and its 

associated effects fall into one of four categories: 

direct effects on essential environmental factors; 

diminished suitability of the habitat factors 

necessary to sustain fisheries yeirld; transmission 

of effects accelerated by the rapid transfer of 

inimical biota among affected ecosystems; potential 

for elaboration of fisheries yield in terms of 

products that are not acceptable for human 

comsumption. Taken together, the patterns of 

change accompanying cultural euthrophication of 

fish production systems are generally recognizable, 

but are not always amenable to differentiation from 

the effects of associated factors, or to precise 

quantification. 

Kohler, A. and F. Holzel (1980) Investigation 

on health conditions of flounder and 

smelt in the Elbe estuary. Helgolander 

Meeresuntersuchungen 33:401-414. 

{disease; health; flounder; smelt; River Elbe; North 

Sea; oxygen; nitrite) 

Since 1958 the occurrence of flounder (Platichthys 

JZesus L.) and smelt (Osmerus eperlanus L.) in the 

lower River Elbe has shifted gradually toward the 

mouth of the river into the North Sea: this may be 

due to periodical deprivation of oxygen and 

fluctuating peaks of toxic nitrite concentrations. 

The interior organs of both fish species caught in 

the Elbe estuary were examined. Macroscopic 

inspections revealed discolouring of the liver to be 

most pronounced in specimens 2-3 years of age. 

Cross-sections of the liver demonstrated 

histopathological symptoms of oedematous 

degeneration in smelt, and extreme lipoid 

vacuolation in flounder increasing with size and age 

of the fishes. The intestine of both species 

investigated showed massive desquamation defects 
of the mucosa, occurring in smelts shortly after 

hatching, and in flounders after immigration into 

the estuary. The observations made suggest the 

influence of toxic compounds distribution in the 

aquatic environment on the health of both species. 

Kononen, K. and A. Niemi (1984) Long-term 

variation of the phytoplankton 

composition at the entrance to the Gulf 

of Finland. Ophelia, supplementum 3: 101-1 10. 

{phytoplankton abundance; assemblage; 

composition; blue-green algae; NIP ratio) 

Quantitative data on the phytoplankton in the sea 

zone off Tvarminne in 1968-1981 have been 

analyzed. The area is characterized by strong 

maxima of diatoms and dinoflagellates in spring 

and occasional maxima of blue-green algae in late 

summer. Rapid changes in hydrography, including 

interactions of different water masses, are frequent 

in the area. The year-to-year fluctuations of different 

phytoplankton species and groups are discussed 

against the background of the hydrographical and 

chemical changes in the Baltic Proper in the 1970s. 

Rises in the surface salinity and nutrient level were 

reflected by the increased total biomass, especially 

in summer, during the late seventies, and 

heterocystic blue-green algae became more 

abundant. The vernal marine diatoms were also 

more abundant in the late seventies than in the late 

sixties, but this was partly caused by variations in 

weather conditions in spring. Diatoma elongatum, 

which favours low salinity, was more abundant in 

the late sixties. However, many dominant marine 

species showed considerable year-to-year fluctuation 

not directly connected with the changes observed in 

the environment during the study period. 

Kroncke, I. (1990) Macrofauna standing 

stock of the Dogger Bank. A comparison: 

11. 1951-1952 versus 1985-1987 Are 

changes in the community of the 

northeastern part of the Dogger Bank due 

to environmental changes? Netherlands 

Journal of Sea Research 25: 189- 198. 

{macrofauna; benthic; opportunistic; polychaetes; 

bivalves; species composition; abundance; 

biomass; Dogger Bank} 

During Aprilhlay 1985-87 some of Ursin's 

(Ursin, 1960) stations on the Dogger Bank from 

Aprilhlay 1951-52 were revisited. This paper 

concentrates on results from the 'Tail End', the 

northeastern part of the Dogger Bank. The 

polychaete species Ophelia borealis and Goniada 

maculata, which were dominant species in 1951-52, 

showed similar distributions in 1985-87 on the 

whole of the Dogger Bank. Changes in the 

macrofauna distribution are obvious in the increase 

of opportunistic small polychaete species like 

Spiophanes bombyx, Scoloplos armiger and 

Magelona spp. In 1985-87 these species occurred in 

high individual numbers per m2 all over the 

Dogger Bank in contrast to 1951-52, when they 

were found only in few individuals. Extensive 

patches of the bivalve species Spisula subtruncata 

discovered at the northeastern border of the Dogger 

Bank in the fifties were not found. Only small, 

fast-growing species like Abra prismatica, Tellina 

fabula and Montacuta bidentata occurred in high 

individual numbers per m2. In 1987 the total 

biomass shows a reduction of about 30% in the 

area compared with 1950-54. 

Some hypotheses are given to explain the changes 

in macrofauna distribution on the Dogger Bank 

between 1950-1954 and 1985-1987, taking into 

account the increasing eutrophication and pollution. 

Lam, C. W. Y. and K. C. Ho (1987) Red tides 

in Tolo Harbour, Hong Kong. In: T. 

Okaichi, D. M. Anderson and T. Nemoto (ed.) Red 

Tides: Biology, Environmental Science and 

Toxicology. Elsevier, New York. 

{red tide; oxygen) 

Tolo harbour, a 15 km long landlocked inlet in 

the northeastern part of Hong Kong, has been 

severely affected by red tides over recent years. The 

frequency of occurences has markedly increased 

since 1980 with 69 incidents recorded from 1980- 

1986. A corresponding increase in dinoflagellate 

numbers in the inlet was found. More than twenty 

causative organisms were identified. Prorocentmm. 

tridtinum. P. dentatum. P. si~moides, 

Gvmnodinium sp., Noctiluca scintillans and a 

variety of small flagellates occurred most 

frequently. Red tides were found throughout the 

whole year with peak occurrence in spring and 

autumn. The water quality, oceanography and 

meteorological conditions characterizing red tide 

occurrence during the period from 1976-1986 were 

analyzed and reported. The increasing red tides are a 

consequence of accelerated eutrophication in Tolo 

Harbour following intensive urban development in 

the catchment. The nutrient loadings of nitrogen 

and phosphorus have increased more than two-fold 

in the past decade. Concurrent increase in the 

nutrient levels in the inlet water has been found. 

The major impact of red tides at present is causing 

fishkills by anoxia. 

Lara, R. J., E. A. Gomez, and A. E. Pucci (1985) 

Organic matter, sediment particle size 

and nutrient distributions in a sewage 

affected shallow channel. Marine Pollution 

Bulletin 16:360-364. 

{nitrogen; phosphorus; Blanca Bay; sediment; 

protein) 

Sediment and seawater samples were taken in a 

sewage affected channel in Blanca Bay, Argentina. 

Protein content, 'humic substances' and silt-clay 

percentage were evaluated in the sediments; MI4+, 
~0~~+ and 'dissolved organic matter' in seawater. 

A highly significant correlation (r=0.747, n=59) 

was obtained for the relation between protein and % 

silt-clay. The 'humic substances' distribution 

depends more on the distance from the outfall than 

does the protein content, probably because of the 

important relation of the latter with the sediment 

particle size. The dissolved components showed a 

dilution pattern similar to that of a contamination 

plume in a moving water mass, reaching normal 

values at 1700 m from the outfall. 

Larsson, U., R. Elmgren, and F. Wulff (1985) 

Eutrophication and the Baltic Sea: 

Causes and consequences. Ambio 14:9- 14. 

{nitrogen; Phosphorus; Baltic) 

New estimates of the total nutrient (N and P) load 

to the Baltic Sea are considerably higher than earlier 

calculations. Estimated total nitrogen load is 1.2 X 

lo6 tonslyear, total phosphorus load 77 X lo3 

tonslyear. These figures may still be too low. The 

load may have increased about 4x for total nitrogen 

and about 8x for total phosphorus, due to man's 

activities. 

Available data indicate that phosphorus is 

deposited in large amounts in the sediment, and that 

nitrogen is lost from the sediment through 

denitrification. 

Larsson, U. and A. Hagstrom (1982) 

Fractionated phytoplankton primary 

production, exudate release and bacterial 

production in a Baltic eutrophication 

gradient. Marine Biology 67:57-70. 

{phytoplankton; primary production; bacteria; 

Baltic) 

The distribution of phytoplankton primary 

production into four size fractions (>lOum, 10- 

3um, 3-0.2 um and <0.2 um), the utilization of 

algal exudates by bacteria and the bacterial 

production were studied in a eutrophication gradient 

in the northern Baltic proper. The polluted area 

exhibits substantially increased nutrient, especially 

nitrogen, levels while only minor differences occur 

in salinity and temperature regimes. Total primary 

production was 160 g C m-2 yr-l at the control 

station and about 275 g C m2 yr-l at the 

eutrophicated stations. The estimated total exudate 

release was 16% of the totally fixed 14c02 in the 

control area and 12% in the eutrophicated area 

(including the estimated bacterial uptake of 

exudates). The difference in 14c02 uptake rates 

between incubation of previously filtered water (<3, 

<2, <1 um) and unfiltered water was used to 

estimated bacterial uptake of phytoplankton 

exudated which were found to contribute about half 

of the estimated bacterial carbon requirement in 

both areas. Bacterial production was estimated by 

the frequency of dividing cells (FDC) method as 

being 38 g C m-2 yr-l at the control station and 

50 g C m-2 yr-l at the eutrophication stations. To 

estimate the mean in situ bacterial cell volume a 

correlation between FDC and cell volume was used. 

The increased annual primary production in the 

eutrophicated area was due mainly to higher 

production during spring and autumn, largely by 

phytoplankton cells (mainly diatoms) retaind by a 

10 um filter. Primary production during summer 

was similar in the two areas, as was the 

distribution on different size fractions. This could 

possibly explain the similar bacterial production in 

the trophic layers at all stations since the bulk of 

bacterial production occurs during summer. It was 

demonstrated that selective filtration does not 

quantitatively separate photoautotrophs and 

bacteria. A substantial fraction of the primary 

production occurs in the size fraction <3um. The 

primary production encountered in the 3-0.2 um 

fraction was due to abundant picoplankton (0.5 to 

8x10~ ind 1-I), easily passing 3 um filter. The 

picoplankton was estimated to constitute up to 

25% of the total phytoplankton biomass in the 

control area and up to 10% in the eutrophication 

area. 

Laws, E. A. (1983) Man's impact on the 

marine nitrogen cycle. In: E. J. Carpenter 

and D. G. Capone (ed.) Nitrogen in the Marine 

Environment. Academic Press, New York. 

{nitrogen; cycle; budget; New York Bight; sewage; 

Kanaohe Bay) 

No abstract 

Laws, E. A. and D. G. Redalje (1979) Effect of 

sewage enrichment on the phytoplankton 

population of a subtropical estuary. 

Pacific Science 33: 129- 148. 

{sewage; phytoplankton; estuary; primary 

production; Kanaeohe Bay; chlorophyll a; biomass; 

nutrients; light; growth rate) 

Phytoplankton primary production; concentrations 

of chlorophyll a, particulate carbon and nitrogen, 

adenosine triphosphate, inorganic nitrogen and 

phosphorus; and secchi depths were measured at 

four stations in Kanaeohe Bay, Oahu, on a 

biweekly basis for 20 months prior to diversion of 

sewage discharges from the bay. Nutrient 

enrichment experiments designed to determine 

biomass limitation indicated that phytoplankton 

biomass, as measured by chlorophyll a, was 

nitrogen-limited in all parts of the bay, and that 

phosphorus was simultaneously limiting in the 

sector of the bay furthest from the sewer outfalls. 

Mean light-saturated productivity indices in all 

parts of the bay were about 11-12 mg C mg-l chl a 

hr-l, values close to the maximum reported for 

phytoplankton in eutrophic marine environments. 
Based on the results of dawn-to dusk C-14 

incubations and an estimated phytoplankton C: chl 

a ratio of 50 by weight, phytoplankton growth 

rates were estimated to fall in the range of 4-6 

percent per hour in all parts of the bay. Such 

growth rates are close to the maximum growth 

rates reported for marine phytoplankton grown on 

light-dark cycles in continuous culture, suggesting 

that phytoplankton growth rates (as opposed to 

biomass) were limited primarily by suboptimal or 

supraoptimal light intensities rather than by 

nutrients. Based on these growth rates and an 

assumed phytoplankton C:N ratio of 5.68 by 

weight, nitrogen recycling was estimated to account 

for 80 percent of phytoplankton nitrogen uptake in 

the part of the bay receiving direct sewage inputs, 

and for over 90 percent of phytoplankton nitrogen 

uptake in the other sectors of the bay. Estimates of 

living and detrital particulate carbon were made 

based on an assumed C: ATP ratio in living 

organisms of 285 by weight. From this 

partitioning, living carbon was found to vary by a 

factor of 3-4 between the sewage-enriched and 

unenriched sectors of the bay. However, estimated 

detrital carbon concentrations were uniform 

throughout the bay, as were the measured 

concentrations of inorganic nitrogen. These results 

are consistent with the interpretation that the 

population of microorganisms, both bacteria and 

phytoplankton, are substrate-limited in all sectors 

of the bay. 

Laws, E. A. and D. G. Redalje (1982) Sewage 

diversion effects on the water column of 

a subtropical estuary. Marine Environmental 

Research 6:265-279. 

{phytoplankton; Kaneohe Bay; sewage; estuary; 

growth) 

A study - of the phytoplankton community and 

water column chemistry in Kaneohe Bay, Oahu, 

Hawaii, before and after the diversion of secondary 

treated sewage from the bay has shown that changes 

in total nutrient concentrations in the water column 

cannot be accurately predicted without taking into 

account water column-benthos interactions. During 

the first year after sewage diversion, the 

decomposition of about 400 tonnes of benthic 

organisms, primarily filter feeders, resulted in water 

column dissolved organic nitrogen and phosphorus 

concentrations roughly an order of magnitude 

higher than those expected in the absence of such 

interactions. The biomass of phytoplankton appears 

to have been N limited both before and after sewage 

diversion, with internal nutrient recycling 

accounting for 70-99% of phytoplankton nutrient 

uptake. Both the biomass and growth rate of the 

phytoplankton declined as a result of the sewage 

diversion, with post-diversion growth rates 

evidently well below nutrient-saturated values. 

Since the principal stresses on the bay's coral reef 

community as a result of the sewage discharges 

appear to have resulted from the elevated 

concentrations of plankton in the water, various 

measures of seston concentration appear to be the 

most ecologically significant indicators of nutrient 

enrichment in the system. 

Lee, G. F. and R. A. Jones (1981) Application 

of the OECD eutrophication modeling 

approach to estuaries. In: B. J. Neilson and 

L. E. Cronin (ed.) Estuaries and Nutrients. Humana 

Press, Clifton, New Jersey. 

{modeling; lake; estuary) 

Approximately five years ago, the Organization 

for Economic Cooperation and Development 

(OECD) initiated a 22 country, 200 lake and 

impoundment study of nutrient load-eutrophication 

response relationships. Emphasis in the study is 

being given to the evaluation of the Vollenweider 

models for correlating nutrients load with 

eutrophication response. The U.S. part of this 

study included the investigation of about 40 

waterbodies or parts thereof. The results of the U.S. 

and the other studies all show a strong correlation 

between the phosphorus load to a water body as 

normalized by mean depth and hydraulic residence 

time, and the planktonic algal chlorophyll, Secchi 

depth (water clarity), and the hypolimnetic oxygen 

depletion rate. These relationships have been 

developed to a sufficient degree of sophistication so 

that they are the method of choice for estimating 

the impact of altering the phosphorus load to a Plimited waterbody on eutrophication-related water 

quality. 

While the U.S. OECD waterbodies were primarily 

lakes and impoundments, included as part of the 

data base upon which the statistical correlataions 

were made were three parts of the Potomac estuary. 

It appears that in general, the OECD eutrophication 

modeling approach is applicable to estuarine 

systems as well as lakes and impoundments. In 

addition to reviewing the U.S. OECD study results, 

this paper presents a discussion of the 

modifications that may need to be made in the 

OECD-Vollenweider eutrophication modeling 

approach in order to apply it to some estuarine 

systems. 

Lee, V. and S. Olsen (1985) Eutrophication 

and management initiatives for the 

control of nutrient inputs to Rhode 

Island coastal lagoons. Estuaries 8: 19 1-202. 

{lagoon; management; macroalgae; nutrient; salt 

pond; seaweeds; macroalgae) 

An assessment of developing eutrophic conditions 

in small temperate lagoons along the coast of 

Rhode Island suggests that in such shallow, 

macrophyte based systems the response to nutrient 

enrichment differs from that described for 

planktonic based systems. The nitrogen loadings 
per unit area of the salt ponds are 240-770 mrnol N 

per m2 per year. Instead of the high nutrient 

concentrations, increased phytoplankton biomass 

and turbidity, leading to eventual loss of benthic 

macrophytes described for such systems as the 

Chesapeake, Patuxent and Appalachicola Bay, 

nutrient enrichment of the Rhode Island lagoons 

has led to increased growth of marine macroalgae. 

The increased macroalgal growth appears to alter 

the benthic habitat and a shift from a grazing to 

detrital food chain appears to be impacting 

important shellfisheries. As more extensive areas of 

organic sediments develop, geochemical cycling 

changes, resulting in higher rates of nitrogen 

re mineralization and accelerated eutrophication. 

The major sources of nitrogen inputs to the salt 

ponds have been identified and a series of 

management initiatives have been designed to limit 

inputs from present and potential development 

within the watersheds of the lagoons. 

Lenzi, M. (1992) Experiences for the 

management of Orbetello Lagoon: 

eutrophication and fishing. In: R. A. 

Vollenweider, R. Marchetti and R. Viviani (ed.) 

Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{seaweeds, fish yield) 

The development of human activity has caused 

hypertrophic conditions in Orbetello Lagoon. 

Decisions are based on increasing water circulation 

and on removal of part of the seaweed biomass. 

Consequently, important changes in the trophic 

structure of the ecosystem have taken place, with a 

significant increase in animal bomass. This fact 

has persuaded us that management of a lagoon 

environment with high production is possible. 

Leppakoski, E. (1980) Man's impact on the 

Baltic ecosystem. Ambio 9: 174- 18 1. 

{Baltic; oxygen; review; benthos; abundance; 

composition) 

The drainage from about 250 river systems and 

the municipal and industrial wastes of some 17 

million people sooner or later end up in the Baltic, 

which has been described as one of the most 

severely polluted sea areas in the world. Human 

activities have caused excessive eutrophication and 

resulted in damaging oil spills, and the chronic 

discharge of heavy metals and other toxic 

substances like PCBs which are seriously affecting 

the biomass of the Baltic. Even though the Baltic 

is one of the most thoroughly studied seas, much 

remains to be done and the author concludes that we 

will probably never fully understand it as a "natural 

and man-influenced system". 

Levin, L. A. (1986) Effects of enrichment 

on reproduction in the opportunistic 

polychaete Streblospio benedicti 

(Webster): A mesocosm study. Biological 

Bulletin 171: 143-160. 

{nutrients; reproduction; polychaete; MERL) 

The influence of organic enrichment on growth 

and planktontrophic development of the spinoid 

polychaete Streblospio benedicti Webster was 

examined in two mesocosm experiments conducted 

at the MERL facility, University of Rhode Island. 

Specimens of S. benedicti were collected and their 

reproductive traits monitored near the conclusion of 

a two-year eutrophication experiment. Nutrient (N, 

P, and Si) enrichment at 8X and 32X the average 

aerial input into Narragansett Bay, Rhode Island, 

resulted in increases in body length, segment 

number, and length per segment, and a doubling of 

brood size in S. benedicti females. These increases 

were substantially higher during May (12OC) than 

August (20°C). Enrichment effects were stronger in 

the 8X than 32X nutrient treatment. In the sewage 

sludge experiment body size increased 20% over 

control values at the highest (8X) sludge treatment 

level (nitrogen loading equivalent to the 8X 

nutrient treatment) but no significant increase was 

noted at the 4X sludge level, which received half as 

much nitrogen as the 8X sludge treatment. Mean 

brood size increased by a factor of 4.6 over controls 

in the 8X sludge treatment and by a factor of 2.3 in 

the 4X sludge treatment. Within the range of adult 

body sizes observed, brood size enhancement 

occurred independent of increased length or segment 

in both nutrient and sludge enrichment treatments. 

The ability to translate elevated food supply directly 

into increased reproductive output may underly 

opportunistic dynamics in macrobenthos. Brood 

size enhancement of the magnitude observed 

probably contributes to the high S. benedicti 

densities found in polluted or organically enriched 

settings. 

Liaci, L. S. (1980) Pollution effects on 

marine biota. Memorie di Biologie Marina e di 

Oceanographia Supp1.10:85-94. 

{organic; colored tides; dinoflagellates) 

Effects of organic substance pollution on marine 

organisms have been studied and related to 

eutrophication. The study refers effects of coloured 

tides due to Dinoflagellates on mollusca and fishes. 

The pattern of uptake and accumulation of some 

heavy metals was taken into consideration. 

In fact, some specimens clearly showed that their 

uptake was directly proportional to the 

-concentration of the heavy metal in the sea. Other 

factor such as salinity, temperature etc. and their 

role on uptake, accumulation and elimination of the 

metal are reported. Furthermore, the effects of 

pesticide pollution has been examined with special 

reference to the episode occurring in the harbour of 

Brindsi where methaldehyde was spilt. 

As a consequence of pollution, skeletal and gill 

deformities, as also defects of reproduction, growth, 
number of eggs and epidermal papillomas and 

hepatic tumors were observed. Alteration of the 

biocenosis and involutions of ecological series due 

to organic slimes have been noted. 

Likens, G. E. (ed.) (1972) Nutrients and 

Eutrophication: The Limiting Nutrient 

Controversy (Proceedings of the 

Symposium on Nutrients and 

Eutrophication: The Limiting Nutrient 

Controversy, W.K. Kellog Biological 

Station, Michigan State University, 11- 

12 February, 1971). American Society of 

Limnology and Oceanography and the Allen Press, 

Lawrence, Kansas. 

No abstract 

LISS (1990) Status report and interim 

actions for hypoxia management. Long 

Island Sound Study, U.S. Environmental 

Protection Agency, {oxygen; hypoxia; modeling; 

impacts; sewage; runoff; management; nitrogen; 

Long Island Sound} 

No abstract 

Littler, M. M., D. S. Littler, and B. E. Lapointe 

(1986) Baseline studies of herbivory and 

eutrophication on dominant reef 

communities of Looe Key National 

Marine Sanctuary. NOAA, NOAA Technical 

Memorandum, NOS MEMD 1. 

{algae; community; nutrient; limitation} 

Presently, only limited knowledge is available 

regarding the dominant algal assemblages at Looe 

Key National Marine Sanctuary and the effects of 

herbivory and nutrient limitation in controlling the 

structure of these communities. The goals of this 

research are to: (1) provide a preliminary inventory 

and general distributional assessment of dominant 

algal communities, (2) initiate pilot studies of the 

effects of nutrient enrichment and grazing on 

structure of the reef communities and (3) contrast 

findings with other reef habitats for which 

comparable data are available. This research 

accomplishes these goals. Increases our 

understanding of ecological processes that influence 

dominant reef communities and provides data of use 

in management decisions directed towards 

preserving Looe Key National Marine Sanctuary in 

a natural state. 

LoCicero, V. R. (ed.) (1975) Toxic 

Dinoflagellate Blooms (Proceedings of 

the First International Conference on 

Toxic Dinoflagellate Blooms, Boston, 4- 

6 November, 1974) (MZT Sea Grant 

Report MZTSG 75-8). Massachusetts Science 

& Technology Foundation, Wakefield, 

Massachusetts. 

No abstract 

Lucht, F. and M. Gillbricht (1978) Long-term 

observations on nutrient content near 

Helgoland in relation to nutrient input of 

the River Elbe. Rappots et Procds-verbaux des 

Re'unions 

Counseil International pour LIExploration de la 

Mer 172:358-360. 

{nutrients; Helgoland; River Elbe; phosphate; 

nitrogen; discharge; freshwater) 

No abstract 

Maclean, J. L. (1984) Indo-Pacific toxic redtide occurrences, 1972-1984. In: A. W. 

White, M. Anraku and K.-K. Hooi (ed.) Toxic Red 

Tides and Shellfish Toxicity in Southeast Asia. 

Southeast Asian Fisheries Development Center and 

the International Development Research Center, 

{toxic; red tide; bloom) 

No abstract 

Maclean, J. L. (1989) Indo-Pacific red tides, 

1958-1988. Marine Pollution Bulletin 20:304- 

310. 

{red tides; Indo-Pacific; dinoflagellate; bloom) 

A summary of recent toxic red tides in the 

southwestern Pacific region is given, showing that 

the dinoflagellate Pytodinium bahamense var. 

compressa remains the major causative organism of 

paralytic shellfish poisoning and is apparently 

continuing to spread around the region. Noticluca 

scintillans blooms occur in some countries and 

have been associated with mass mortality of fish. 

The raphidophyte Chattonella (=Hornellia) has 

been implicated in both fish and shrimp kills. A 

relationship with El Nino-Southern Oscillation 

(ENSO) events and some similarities with recent 

plagues of the crown-of-thorns starfish (Acanthaster 

planci) are discussed. 

Maestrini, S. Y. and E. Graneli (1991) 

Environmental conditions and 

ecophysiological mechanisms which led 

to the 1988 Chrysochromulina polyepis 

bloom: An hypothesis. Oceanologica Acta 

14:397-413. 

{bloom; nuisance; nitrogen; Scandinavian) 

Information available on the 1988 

Chrysochromulina polyepis bloom in Scandinavian 

coastal waters is evaluated. Special attention is paid 

to the early bloom stage and the sequence of both 

environmental and physiological mechanisms 

leading to the bloom, and a tentative, integrated 

scheme is given of causes which led to the observed 

events. 

Environmental conditions favourable for 

Chrysochromulina polyepis are considered to have 

resulted from: a) an exceptionally high runoff of 

nitrogen-rich water in winter and early spring; b) 

strong mixing of the water column, immediately 
followed by a long period of vertical stability and 

stratification; c) a diatom bloom 30% larger than 

usual, peaking around 20 March; and d) a second 

diatom bloom in mid April, largely dominated by 

Skeletonema costatum. These events combined to 

cause the euphotic layer to become silicateexhausted, phosphorus-poor and fairly nitrogenrich. The lack of silicate prevented diatom growth. 

Weak turbulence and strong stratification favoured 

growth of non-siliceous, motile nanoplankton. 

C. polyepis is believed to have outcompeted coexisting algal species by producing a toxin acting 

first as a grazer repellent, while its cell density was 

still below lo4 individuals 1-l. At the conjectured 

location of first appearance of the bloom (or at one 

of several locations), in the Skagerrak near Gullmar 

Fjord, the grazing repellent forced a shift of grazing 

to other algae around the latter part of April. With 

increased population density of C. polyepis (lo6 

cells 1-I), the toxin then became sufficiently 

concentrated to affect directly not only by grazers, 

but all other organisms. From late April, the C. 

polyepis population increased free from grazing 

and other losses. In conditions of light and nutrient 

sufficiency, a high growth rate (corresponding to 

0.8 div. d-l) was achieved. Actively growing cells 

were buoyant and remained in the upper part of the 

water column. 

With increasing population density (lo7 cells 1- 

I), nutrient became exhausted and self-shading 

decreased available light. By mid May, the growth 

rate had decreased to zero, and the population started 

to become senescent. C. polyepis deteriorated in 

physiological state, perhaps party because of selfpoisoning, and this led to sinking. No further 

growth occurred, but cell densities showed higher 

values (several lo7 cells 1-l, with maximum 

density between 5 and 10 m) because distribution 

became patchy both horizontally and vertically. At 

the end of May a significant part of the population 

had died, and cell leakage produced an increase in 

dissolved organic matter. The resultant lower 

turbidity and regenerated nutrients then allowed 

other flagellates and some diatoms to resume 

growth. The remaining C. polyepis cells sunk to 

the pycnocline (15-20 m), which acted as a barrier 

and greatly retarded this sinking, by which means 

the cells became concentrated, and produced the 

highest reported cell densities at the pycnocline. It 

is assumed that the bloom later sedimented. 

Since relevant environmental conditions for the 

initiation of the bloom occurred over a very large 

area (75. lo3 lun2) and the dominant currents were 

not altogether consistent with the spreading of the 

bloom, polygenesis of the bloom should be 

considered a possibility. 

To what extent long-term global change and 

eutrophication might have contributed to this 

exceptional bloom remains unclear. Since the 

production of the toxin seems to be favoured by 

phosphorus deficiency, however, a large input of 

nitrogen in combination with a reduction in the 

phosphorus loading is believed to have contributed 

indirectly to the bloom, by changing the nutrient 

status of the coastal waters from being nitrogen- to 

phosphorus-limited. 

Makinen, A., I. Haahtela, H. Ilvessalo, and J. 

Lehto (1984) Changes in the littoral rocky 

shore vegetation in the Seili area, SW 

Archipelago of Finland. Ophelia Suppl. 

31157-166. 

{macroalgae; seaweeds; species composition; rocky 

shore; Archipelago) 

The littoral macrophyte communities were studied 

in 1968-69 by Ravanko at 23 rocky shores around 

Seili, SW archipelago of Finland. Four of these 

shores were revisited in 1981-82. The most 

noticeable finding was the altered species 

composition in the area. The number of green algae 

and phanerograms had increased and the number of 

brown and red algae decreased. The macrophyte 

biomass (fresh and dry weight m-2) had drastically 

decreased because of the total lack of Fucus 

vesiculosus. 

Makinen, A., J. Kaaria, and M. Rajasilta (1988) 

Factors controlling the occurrence of 

Furcellaria lumbricalis (Huds.) Lamour 

and Phyllophora truncata (Pallas) Zinova 

in the upper littoral of the Archipelago 

of SW Finland. Kieler Meeresforschungen, 

Sonderheji 6: 140- 146. 

{red algae; Furcellaria; Phyllophora; plankton; 

light; Mytilus; archipelago) 

The influence of eutrophication on the occurrence 

of the red algae Furcellaria lumbricalis and 

Phyllophora truncata was studied in the sea area of 

Turku. Due to increased sediment load and 

planktonic production, light penetration in the 

water has decreased in the northern parts of the 

study area. Furthermore, the quality of the sea 

bottom has changed. Plots of 1 m2 (N=100) in the 

upper littoral zone (0.5-6m) were studied by 

SCUBA diving, in order to investigate the factors 

controlling in the occurrence of the red algae. Two 

transects were situated in the eutrophicated area, and 

two in the reference area. In the plots, the 

percentage cover of each plant species and of 

Mytilus edulis was documented, and the Secchi disc 

visibility and depth was measured. 

The numerical data were analyzed by correlation 

analysis and stepwise multiple regression analysis 

(BMDP2R). In the eutrophicated area, abiotic 

factors (percentage cover of plain bottom and 

bottom quality) explained 81.28% of the variation 

of red algae cover. In the reference area, the biotic 

factors (total number of species and Mytilus cover 
in %) were the most important factors, explaining 

66.4% of red algae cover. 

Malone, T. C. (1984) Anthropogenic 

nitrogen loading and assimilation 

capacity of the Hudson River estuarine 

system, USA. In: V. S. Kennedy (ed.) The 

Estuarv as a Filter. Academic Press. Orlando. 

{nitrogkn; phytoplankton; productidn; regeneration; 

Hudson River; sewage; capacity) 

The coastal plume of the Hudson River estuary 

receives inputs of new nitrogen of sewage origin 

from the lower estuary and of offshore origin from 

adjacent coastal water. As a consequence of these 

inputs, the plume is one of the most productive 

coastal systems in the world's ocean. The extent to 

which such high production reflects the input of 

sewage-nitrogen depends on the capacity of 

phytoplankton to assimilate new nitrogen input to 

the plume and on the magnitude of sewage-nitrogen 

input relative to coastal inputs, i.e., on the capacity 

of the plume to "filter" inputs of new nitrogen. 

The importance of new nitrogen varies seasonally 

relative to nitrogen recycled within the plume. 

Regenerated production increases from 10% of total 

phytoplankton production during winter to a 

maximum of 80% during summer. Sewagenitrogen supports an average of 54% of new 

production during the spring bloom period (MarchMay) compared to 121% during the subsequent 

period of stratification (June-October) and 221% 

during winter (November-February). On an annual 

basis, these estimates indicate that phytoplankton 

production has increased by ca. 30% in response to 

the input of sewage-nitrogen. 

~hese patterns have important implications in 

terms of cross-shelf transport and exchange. The 

supply of new nitrogen from offshore is highest 

during March-May and, along with sewagenitrogen, appears to support a large seaward export 

of nitrogen in the form of phytoplankton biomass. 

In contrast, most new nitrogen is exported via 

plankton fraction of phytoilankton food webs 

during summer. Consequently, a relatively small 

fraction of phytoplankton production is available to 

fuel oxygen demand below the pycnocline during 

summer when assimilation capacity is greatest. 

Malone, T. C. (1991) River flow, 

phytoplankton production and oxygen 

depletion in Chesapeake Bay. In: R. V. 

Tyson and T. H. Pearson (ed.) Modem and Ancient 

Continental Shelf Anoxia. The Geological Society, 

London, England. 

{oxygen; phytoplankton; Chesapeake Bay; climate; 

anoxia) 

Oxygen depletion leading to a summer anoxia in 

subpycnocline waters of the mesohaline reach of 

Chesapeake Bay has been a annual event for at least 

five decades. It is generally assumed that the spatial 

and temporal extent of bottom water anoxia has 

increased in response to higher phytoplankton 

productivity due to anthropogenic nutrient loading. 

However, synoptic observations of environmental 

conditions (river flow, temperature, salinity, 

dissolved oxygen, and nutrients) and phytoplankton 

from 1984 to 1988 indicate that oxygen depletion 

is not related to phytoplankton productivity on 

seasonal to interannual scales. Rather, nutrient 

loading is linked to oxygen depletion and summer 

anoxia via the spring accumulation of 

phytoplankton biomass which is more than 

sufficient to fuel oxygen depletion. Thus, 

interannual variability in the magnitude of summer 

anoxia is primarily driven by climate factors which 

govern vertical density stratification (mixing). 

Mambeke, F. v., M. A. Bianchi, and G. Cahet 

(1984) Short-term bacterial reactivity of 

nitrogen-enriched seabater of a eutrophic 

lagoon. Estuarine, Coastal and Shelf Science 

19:291-301. 

{bacteria; nitrogen; lagoon; growth; respiration) 

We carried out, in 

to determine the 

communities were defined in terms of tlyir 

catabolic potentialities and 'specific' diversity,. 

Bacterial heterotrophic activity was established by 

measuring heterotrophic uptake and the , 

mineralization percentage of labelled glucose or 

amino acids. From these data, ratios of AMPIATP 

and PIB were calculated where P is the uptake of 

labelled substrate and B is the biomass. 

After the nitrogen enrichment an increase of the 

respiration was seen at first. After 6 h, a biomass 

peak appeared associated with a continuous increase 

in the number of cells. Bacterial growth was 

concurrent with a qualitative modification of the 

community: UAI increased, and diversity dropped. 

A less diversified community resulted suggesting 

an immature ecological state (zymogenous 

bacteria). Low values of respiration percentage 

(20%) characterized high growth yields. 

Marchetti, R. (1992) The problems of the 

Emilia Romagna coastal waters: facts and 

interpretations. In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{gels, Adraitic Sea) 

In this work, the two main problems concerning 

the upper Adriatic coastal waters are examined: the 

so-called algal blooms and the gels or excretions. 
The former comprises the uncontrolled reporduction 

of microalgae causing an enormous mass of living 

matter which is incompatible with bathing, the 

decomposition of which causes oxygenc consuption 

and the death of the animals living on the sea 

bottom. This phenomenon is not new and is 

strongly connected with the polluting loads from 

the Po and other minor rivers along the coast. The 

latter - in respect to the upper Adriatic coatal waters 

during the summers of 1988 and 1989 - is due to 

the excretion of products which are normally 

emitted by algal cells with various functions and 

which, under circumstances still not completely 

understood, can reach enormous amounts as 

occurred in the summer of 1989. In the July of 

that year, it was calculated that about 10,000 km2 

of sea were covered by a layer of gel. The nature of 

the compounds of which the gel is formed hs been 

individuated in polysaccharides forming a structure 

incorporating 98-99% of water and which is 

completely harmless for humans. The most 

probable cause of the formation of these gels has 

been identified with the strong meteorological and 

climatic anomalies which characterized that period, 

traces of which have not been found in the data of 

the preceding 100 years. 

The conclusions of this analysis are that, 

although for the first of the two events adequate 

action on the sources of contamination would have 

some probability of success, for the second there is 

no reasonable counter-action to be proposed at 

present. 

Matsuoka, K. (1989) Morphological features 

of the cyst of Pyrodinium bahamense var. 

compressum. In: G. M. Hallegraeff and J. L. 

Maclean (ed.) Biology, Epidemology and 

Management of Pyrodinium Red Tides. Fisheries 

Department, Ministry of Development, Manila. 

{dino; Pyrodinium; cyst) 

Cysts of Pyrodinium bahamense var. compressum 

and several similar species are compared with 

special attention to differences in process characters. 

Keys to these species with and without references 

to archeophyle features are provided. 

Mattson, J. and 0. LindBn (1983) Benthic 

macrofauna succession under mussels, 

Mytilus edulis L. (Bivalvia), cultured on 

hanging long-lines. Sarsia 68:97- 102. 

{benthic; macrofauna; succession; culture; 

sediments; Capitella; Nucula; opportunistic; 

abundance; biomass) 

Cultures of the mussel Mytilus edulis produce 

accumulations of feces as well as mussels on the 

sea floor below the cultures. Changes in the 

sediment composition and benthic community 

structure under such cultures were studied over a 

three-year period on the Swedish west coast. 

Deposition of organic matter created several 

centimeters of sediment each year, resulting in 

production of H2S in the uppermost layer. 

Originally, the benthic fauna was dominated (as per 

numbers of individuals) by Nucula nitidosa and (as 

per biomass) by Echinocardium cordatum and 

Ophiura spp. After 6-15 months of mussel 

culturing these and other species disappeared and 

were replaced by opportunistic polychaetes 

(Capitella capitata, Sclelepis fuliginosa, and 

Microphthalmus sczelkowskii). The abundance of 

these species fluctuated due to changes in the 

position of the redoxcline and their annual life 

cycle. The anaerobic sediments and mass occurrence 

of opportunistic polychaetes were localized to a 

zone under and 5-20m around the cultures. After 

harvesting of the mussels only limited recovery 

was observed after one and a half year. 

McComb, A. J., R. P. Atkins, P. B. Birch, D. M. 

Gordon, and R. J. Lukatelich (1981) 

Eutrophication in the Peel-Harvey 

estuarine system, Western Australia. In: 

B. J. Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{green algae; Cladophora; phytoplankton; nutrient; 

Australia) 

The most obvious symptom of eutrophication in 

this estuarine system is a green alga, Cladophora, 

which was sparse in 1966 but now accumulates and 

rots on the shores. The work is part of a continuing 

study designed to asses the relationships between 

nutrient input and the growth of Cladophora and 

phytoplankton. 

The system consists of two shallow basins, 

interconnected and linked to the ocean by a narrow 

channel. It is fed by three rivers; 90 percent of river 

flow occurs during four winter months. 

Phytoplankton and water nutrient levels are low 

in summer, but high during and after an input of 

river nutrients in winter. Nitrogen:phosphorus 

ratios, regression analyses, and nutrient limitation 

assays suggest that nitrogen is potentially limiting 

in summer and autumn, phosphorus in winter and 

spring. The Harvey typically supports higher 

phytoplankton levels that the Peel. Diatom 

populations may be replaced by blue-green in 

summer in the Harvey. 

Cladophora forms detached sphere of branched 

filaments and is only prominent in the Peel. 

Changes in biomass and growth of confined 

populations in the field, together with laboratory 

experiments, show that growth occurs when 

temperatures and light intensities are high, not in 

winter when water column nutrient levels are high. 

Water from between the algal spheres has increased 

levels of phosphorus compared with the water 

column above, emphasizing the possible 

importance of nutrient release from decaying 

material below. It is suggested that phytoplankton 

are important in trapping water-column nutrients 
during and after river nutrient input, and that 

subsequent Cladophora and phytoplankton growth 

depends on nutrient recycling. 

McErlean, A. J. and G. Reed (1981) Indicators 

and indices of estuarine overenrichment. 

In: B. J. Neilson and L. E. Cronin (ed.) Estuaries 

and Nutrients. Humana Press, Clifton, New Jersey. 

{indicator; index; estuary; eutrophication) 

Indicators and indices of nutrient overenrichment, 

which might be applied to the problems of 

estuarine overenrichment, are reviewed from the 

literature from the time of the national symposium 

on eutrophication (1968) to the present. While 

many existing methods have been applied to 

estuarine work and some new ones have been 

developed, there do not appear to be a large number 

of usable techniques for estuarine investigation. 

McLusky, D. S., M. Teare, and P. Phizacklea 

(1980) Effects of domestic and industrial 

pollution on distribution and abundance 

of aquatic oligochaetes in the Forth 

estuary. Helgolander Meeresuntersuchungen 

331384-392. 

{distribution; abundance; Oligochaetes; Forth 

estuary; macrofauna; diversity) 

The Forth estuary is a major estuary on the east 

coast of Scotland; it receives effluents from 

domestic and industrial (petro-chemical and 

distilling) sources. Following a study on the 

distribution of macrofauna of the intertidal areas in 

relation to pollution (McLusky et al., 1978), this 

paper is concerned with the distribution and 

abundance of aquatic oligochaetes and the small 

polychaete Manayunkia aestuarina in relation to 

estuarine salinity, organic enrichment, and 

industrial effluent. In the most polluted parts of the 

estuary oligochaetes are the sole inhabitants of the 

mudflats; in other less polluted flats they are very 

abundant. In the least polluted parts the numbers of 

oligochaetes diminish as the numbers and diversity 

of macrofauna increases. Estimates of the 

production of oligochaetes are given. 

Menesguen, A. (1992) Modelling coastal 

eutrophication: the case of French Ulva 

mass blooms. In: R.A.Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

(Ulva, macroalgae) 

During the last decade, the excessive proliferation 

of Ulva sp. (Chlorophyceae) during spring and 

summer has been detrimental to recreational use of 

several beaches of Brittany (France). These 

worldwide mass blooms are well known in 

eutrophicated lagoons (e.g. Venice, Tunis), but are 

paradoxical along open beaches with a large tidal 

range and low terrestrial nutrient inputs. Numerical 

modelling of the coupled physical and biological 

aspects clearly explains the Ulva mass blooms by 

the dynamic trapping of water masses in bays 

where tidal residual drift vanishes, and shows the 

main responsibility to be the increase of nitrogen 

inputs (from agricultural sources) in triggering the 

recent magnification of a natural phenomenon. 

Messner, U. and J.-A. v. Oertzen (1990) Recent 

changes in the phytal zone of Greifswald 

Bay. Limnologica 20: 183-1 86. 

{Greifswald Bay; submerged vegetation; coverage; 

plankton; light; red algae) 

The phytal zone of Greifswald Bay, a coastal 

water in the northeast of the GDR was studied from 

spring to autumn in 1985. The submersal 

vegetation was mapped, and it was found that the 

coverage of the bottom had decreased from 90% 

(Seifert, 1938) to about 10%. The authors consider 

increasing plankton production and the deterioration 

of light conditions at the bottom owing to 

eutrophication are the main causes of this decline. 

Quantitative studies of the macrofauna of the phytal 

zone show that the zone covered by red algae 

[mainly Furcellaria fastigiata (L.) Lamour in the 

investigation area] is populated by a large number 

of species in various growth stages. The fauna 

inhabiting the less ramified Potamogeton 

pectinatus L. and Zostera marina L. stands was less 

diversified. It can be assumed that the decline of the 

red algae stocks to insignificant remains has led to 

major changes in the food web and energy flow. 

Although no quantitative data can yet be offered, it 

must be feared that loss of this habitat threatens the 

continued existence of a hemng population there. 

The dense bluegreen blooms in mid-summer may 

restrict the use of these waters by holiday makers 

for bathing. The changes that have taken place thus 

seem to jeopardize many of the uses to which 

Greifwald Bay has been put. 

Millner, R. S. (1980) Pulp and paper mill 

waste pollution in the Swale, a tidal 

channel on the east coast of England. 

Helgoliinder Meeresuntersuchungen 33:366-376. 

{pulp; paper; oxygen; benthos; abundance) 

The effect of pulp and paper mill waste discharges 

in the Swale, a tidal channel on the southeast coast 

of England, has been studied. A pulsing tidal 

movement in the Swale results in effluent being 

trapped within the estuary for up to 20 days. This 

has resulted in adverse effects on the quality of the 

water and sediment along a substantial part of the 

estuary with the greatest effect occurring east of the 

mill in the direction of the residual current. A 

reduction in the dissolved oxygen concentration 

occurred along a 13-km stretch of the estuary with 

mean values falling to 53% saturation near the 

mill. High levels of organic matter in the sediment 

reflect the deposition of material from the mill 

effluent stream and loss on ignition values of up to 
12 % were found east of the mill, falling to 5 % 

within 4 km northwest of the mill. The macrofauna 

of the intertidal mudflats was typical of a stressed 

environment with a low number of species and 

with a few being represented by large numbers of 

individuals. The dominant species was the sabellid 

polychaete Manayunkina aestuarina which reached 

peak numbers of over 1.5 million m-2 1.5 krn east 

of the mill. An increase in the total biomass near 

the mill is thought to be associated with organic 

enrichment from the mill wastes. 

Mills, C. A., S. I. Heaney, C. Butterwick, J. E. 

Corry, and J. M. Elliott (1990) Lake 

enrichment and the status of Windermere 

charr, Salvelinus alpinus (L.). Journal of 

Fish Biology 37 (Supplement A): 167- 174. 

{fish; lake; trophic status; nutrient; enrichment; 

deoxygenation; feeding) 

All English population of Artic charr, Salvelinus 

alpinus (L.), are found in the Lake District 

(northwest England). There are at least four races of 

charr in Windermere, the largest lake in England; 

the North and South basins of the lake each contain 

two distinct races that spawn in autumn and spring 

respectively. The spring spawners in both basins 

probably represent less than 15% of the total 

population in the lake. 

Changes in the population density of charr in the 

lake are described briefly and examined in relation 

to the trophic status of the lake. Other factors that 

could possibly affect the charr population are 

reviewed, especially the influence of climate 

change. 

Mingazzini, M., A. Rinaldi, and G. Montanari 

(1 992) Multi-level nutrient enrichment 

bioassays on Northern Adriatic coastal 

waters. In: R. A. Vollenweider, R. Marchetti 

and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{species abundance, nutrient enrichment, Adriatic 

Sea, limitation) 

In order to evaluate the role of nutrients on 

phytoplankton growth in Northern Adriatic coastal 

waters, nutrient enrichment bioassays at three 

different complexity levels of the experimental 

system were undertaken. The algal growth 

responses both to inorganic (P and N) and to 

organic (group B vitamins) added nutrients were 

measured on Skeletonema costatum an d 

Prorocentrum micans, which are considereed the 

most representative species in local blooms within 

the two groups of diatoms and dinoflagellates, 

respectively. The results obtained from the 

simplest experimental level, in which the cultured 

algae were added to filtered sea water, are compared 

with the superior levels in which the same species 

were naturally abundance in phytoplankton 

communities of tested samples. A good agreement 

has been found in data sets, indicating phosphorus 

as the primary stimulating algal growth nutrient 

while, from a compositional point of view, 

diatoms always accounted for most of the 

phytoplankton increase. A similar effect of diatom 

dominance has also been observed in autumn assavs 

conducted on a differently shaped community 

(represented by Chaetoceros spp. and Gymnodinium 

sp.) while nitrogen and phosphorus together 

stimulated phytoplankton growth. 

Montgomery, R. T., B. F. McPherson, and E. E. 

Emmons (1991) Effects of nitrogen and 

phosphorus additions on phytoplankton 

productivity and chlorophyll a in a 

subtropical estuary, Charlotte Harbor, 

Florida. U.S. Geological Survey, WaterResources Investigations Report 91-4077. 

{nitrogen; phosphorus; phytoplankton; 

productivity; estuary; Charlotte Harbor) 

The response of natural phytoplankton 

assemblages in a subtropical coastal plain estuary, 

Charlotte Harbor, Florida, to inorganic nitrogen and 

phosphorus additions was determined from 

measurements of relative changes in both the 

uptake of carbon-14 and concentrations of 

chlorophyll a. The effects of nitrate plus nitrite 

nitrogen, ammonia nitrogen, and orthophorus 

additions over a series of concentrations were 

evaluate through in situ experiments conducted 

during periods of seasonally low and high river 

inflows. The responses to nutrient additions were 

evaluated for three different size fractions of 

phytoplankton. Relative changes of phytoplankton 

carbon uptake and chlorophyll-a concentrations 

were highly variable with regard to season, 

location. nutrient. and size fractions. 

withinareas of the estuary characterized by lower 

salinities, phytoplankton exhibited a distinct 

seasonal pattern to additions of inorganic nitrogen. 

Under seasonally high freshwater inflow, 

phytoplankton show little response to inorganic 

nitrogen additions, whereas under seasonally low 

inflow, phytoplankton responded to the inorganic 

nitrogen additions. The seasonally high freshwater 

inflow increased ambient inorganic nitrogen 

concentrations and water color. The high water 

color greatly reduced light penetration in the water 

column and limited phytoplankton productivity. 

The effect of nutrient additions in the higher 

salinity reaches of the estuary indicates that, under 

normal conditions, these areas are continually 

nitrogen limited. During periods of high freshwater 

inflow during the summer months, the estuary can 

be divided conceptually into a low-salinity zone 

where phytoplankton production is mediated by 

light availability, as determined by high water 

color, abd a high-salinity zone where 

phytoplankton production is nitrogen limited. 

Seasonal nutrient concentration data and 
comparisons among inorganic nitrogen inputs and 

observed phytoplankton productivity support the 

experimental bioassay results. Each of these lines 

of evidence indicates that, exclusive of seasonal 

riverine influences that affect light penetration of 

the water column, nitrogen availability normally 

limits phytoplankton production within the 

Charlotte Harbor estuarine system. 

Moore, C. G. and T. H. Pearson (1984) 

Response of a marine benthic copepod 

assemblage to organic enrichment. 

Proceedings of the Second International Conference 

on Copepod, Ottawa, Canada. 

{benthic; copepods; assemblage; sewage; Scottish 

coast; density) 

No abstract 

Mori, K. (1979) Effects of artificial 

eutrophication on the metabolism of the 

Japanese oyster Crassostrea gigas. Marine 

Biology 53:361-369. 

{oyster; Crassostrea; gonads; development; 

spawning; fats; mortality; Matsushima Bay; Japan; 

growth} 

Matsushima Bay, one of the richest oyster-culture 

areas in Japan, is subject to heavy artificial 

eutrophication, mainly from the inflow of city and 

factory sewages. The physiological activity in the 

oyster Crassostrea gigas in hanging cultures (i.e. 

suspended in wire-mesh cages) in this embayment 

declined markedly with progressive development of 

the gonads; this decline in activity coincides with 

eutrophication-induced accumulation of fatty 

material in the epithelia of the digestive respiratory 

substrates in order to maintain its increased 

physiological needs. However, the fats are 

inefficient energy sources for the oyster and hence, 

during each spawning season, 50% mortality 

occurs. In addition, such phenomena as 

overmaturation of the gonad and disturbance of the 

lipid and steroid metabolism seem to accelerate this 

mass mortality. On the basis of the results, the 

author has compiled a schematic diagram which 

illustrates the possible causes of mass mortality 

among C. gigas in an eutrophic environment such 

as Matsushima Bay. 

Morton, B. (1989) Pollution of the coastal 

waters of Hong Kong. Marine Pollution 

Bulletin 2O:3 10-318. 

{Hong Kong; Victoria Harbour; Tolo Harbour; 

oxygen; phytoplankton; algae; Ulva; red tides; 

benthos; abundance; benthos; fish} 

This paper reviews the available literature on 

Hong Kong's marine coastal pollution. The highly 

urbanized and industrial cities of Victoria and 

Kowloon pollute Victoria Harbour with untreated 

domestic sewage and industrial effluents. The 

development of new cities, Shatin and Tai Po, on 

the shores of the partially enclosed Tolo Harbour 

have created Hong Kong's first critical marine 

environmental problem, secondary sewage 

treatment schemes having been inadequate to 

prevent this. Agricultural wastes (notably pig 

manure) compound the problem in Tolo Harbour, 

but virtually everywhere in the New Territories 

lowland water courses are polluted from this source 

leading to the degradation of most enclosed bays. 

Sufficient evidence is at hand to understand local 

long-term trends in environmental degradation. 

Hong Kong's coastal water are polluted generally; 

Tolo Harbour, grossly so. Concern exists regarding 

the safety of bathing beach water quality, 

contaminated sea food, and the overall quality of 

other enclosed bodies of water in Hong Kong. 

Moshiri, G. A., N. G. Aumen, and W. G. 

Crumpton (1981) Reversal of the 

eutrophication process: A case study. 

In: B. J. Neilson and L. E. Cronin (ed.) Estuaries 

and Nutrients. Humana Press, Clifton, New Jersey. 

{bloom; dinoflagellates; fishkill) 

Bayou Texar, Pensacola, Florida, is a bayou 

estuary which was advancing toward eutrophy, due 

to nutrient loadings from various sources. The 

occurrence of characteristic symptoms of 

eutrophication, coupled with the closing of the 

bayou for water contact recreation, led to the 

initiation of an intensive seven-year study. Initial 

results indicated that Carpenter's Creek is a prime 

source of most nitrogen species and that an 

exchange exists between dissolved phosphates and 

those adsorbed onto sediments. Carbon fixation 

rates varied, with the stations closest to Carpenters 

Creek exhibiting the most productivity. It also 

appeared that nitrates may be more important in 

controlling rates of carbon fixation than 

phosphates. An ensuing study directed at phosphate 

exchange suggested that reducing conditions in the 

muds cause substantial release of iron-bound PO4 

from the sediment-water interface followed by 

adsorption onto particles in the aerobic flocculent 

layer above the interface. This mechanism could 

allow sufficiently rapid exchange of PO4 between 

sediment and water resulting in a low, stable 

concentration in the water as observed. Other 

investigations involving bacterioplankton, 

dissolved glucose, and heterotrophic production 

indicated that algal primary productivity is the 

major source of dissolved glucose. A final study 

demonstrated a paucity of benthic 

macroinvertebrates, probably due to a graded 

suspension of particles at the sediment-water 

interface and the absence of a sharply defined 

bottom. The zooplankton community exhibited a 

low diversity while numbers of individual species 

were high. The phytoplankton community was 

composed primarily of dinoflagellates, with 

diatoms, cryptophytes, chlorophytes and 
microflagellates occurring in lesser quantities. It 

was also suggested that the importance of toxins 

from algal blooms to fish kills may be greater than 

previously indicated. Specific recommendations 

made after the fourth year of study were 

implemented in most part and led to a substantial 

improvement of water quality and the subsequent 

opening of the bayou to the public for recreational 

use. 

Murakami, A. (1973) Red tide in the Seto 

Inland Sea. Information Bulletin on 

Planktology in Japan 19: 13 1- 132. 

{Japan; nitrogen; phosphorus; red tide; bloom) 

In the past 10 yr, a serious situation of red tide 

has developed in the Inland Sea. The probable cause 

is excessive eutrophication. In the Inland Sea, the 

concentrations of soluble inorganic N and P in 

seawater have increased greatly compared with 

concentrations found 20 yr ago. The only 

fundamental countermeasure of such red tide is to 

eliminate the source of excessive eutrophication. 

Musayeva, E. I. (1985) Mesoplankton near 

the Bulgarian Coast. Oceanology of the 

Academy of the Sciences of the USSR 25:647- 

652. 

{zooplankton; biomass; mesoplankton) 

The distribution of mesoplankton in Burgas Bay 

in 53 bottle samples taken in October-November 

1982 is discussed. A drop in the total biomass of 

zooplankton from north to south can be traced on a 

northern meridianal section (Cape Krotiriya to Cape 

Kaliakra), probably resulting from a decrease in the 

eutrophication effect of the Danube in this 

direction. The plankton off the Bulgarian coast was 

in typical autumn condition. In the southern part of 

Burgas Bay, where there is a discharge current 

carrying eutrophicated sewage from the city of 

Burgas, various stages in the development of the 

community, from a young community in the inner 

end of the bay to a mature one at its outlet, were 

observed. 

Myers, V. B. and R. I. Iverson (1981) 

Phosphorus and nitrogen limited 

phytoplankton productivity in 

northeastern Gulf of Mexico coastal 

estuaries. In: B. J. Neilson and L. E. Cronin 

(ed.) Estuaries and Nutrients. Humana Press, 

Clifton, New Jersey. 

{nitrogen; phosphorus; limitation; phytoplankton; 

productivity; nutrients) 

An understanding of nutrient limitation of 

estuarine phytoplantkon growth is important in 

making environmentally sound decisions 

concerning watershed development and the use of 

aquatic environments for waste disposal. 

Experiments to determine nutrient limitation of 

phytoplankton productivity were conducted 

monthly during the summers of 1975 and 1976 in 

several shallow north Florida coastal and estuarine 

systems by inorganic carbon-14 uptake and 

phosphorus-32 bioassays. The results of these 

nutrient enrichments experiments suggest that 

phosphorus is frequently more important than 

nitrogen in limiting phytoplankton productivity in 

nearshore northern Gulf of Mexico. A multiple 

regression model was constructed to determine 

which combinations of environmental and nutrient 

variables could explain the most variation of 

phytoplankton productivity in these coastal 

systems. The final regression model was: 

P.P. = 32.1 + 48.4 S.R.P. = 0.54 Salinity 

where: P.P. is phytoplankton productivity in ugC1- 

hr-l, S.R.P. is soluble reactive phosphate in ugatom PO4 1-l, and salinity is part-per thousand. 

This model explained 64 percent of the variation in 

phytoplantkon productivity. 

Nitrogen has been identified as the primary 

limiting nutrient for phytoplankton in coastal 

waters and it has been proposed that the removal of 

phosphorus from marine waste discharges will have 

little impact on the control of eutrophication. The 

observation that phosphorus is important in 

limiting phytoplankton productivity in these 

coastal and estuarine areas suggests that water 

quality planning for the coastal zone is best done 

on a regional basis, with consideration given to 

local nutrient cycling processes. 

Nakanishi, H., M. Ukita, M. Sekine, M. 

Fukagawa, and S. Murakami (1992) 

Eutrophication control in the Seto Inland 

Sea. In: R. A. Vollenweider, R. Marchetti and 

R. Viviani (ed.) Marine Coastal Eutrophication. 

Elsevier, Amsterdam. 

{fish yield, Japan) 

The Seto Inland Sea, surrounded by the four large 

islands of Honshu, Shikoku and Kyushu, is famous 

for its scienic beauty and coastal fishery. However, 

the beauty of this areas has been threatened by a 

high concentration of human population and 

industry which contribute to water pollution and 

euthrophication problems. 

In this study, we examine the present water 

pollution problems of this area; we have studied the 

history of water pollution in the Seto Inland Sea, 

the influx of nutrients into all sections of-this area 

and the relationship between euthrophic levels and 

fishery productions in each section of the Seto 

Inland Sea. We have also analyzed the secondary 

organic pollution which is a result of the primary 

photosynthesis production and the comprehensive 

eutrhopic situation of the sea and each of its 

sections, in comparison with other coastal seas in 

Japan. 

From the results of this evaluation, we propose 

the following four strategies for eutrophication 
control in the Seto Inland Sea: (1) Elimination of 

nitrogen (N) and phosphorus (P) discharging loads 

by means of controlled chemical fertilizer 

consumption and recycling of human and livestock 

excreta as fertilizer (as a conclusion of the budget of 

N and P in Japan); (2) Consolidation of treatment 

facilities for the point sources of pollutants such as 

domestic sewage and industrial waste water; (3) 

Diversion of the discharging N and P loads from 

the Seto Inland Sea into the adjacent open sea aras 

of Japan w;hich have a larger carrying capacity 

through the exclusion system such as a sewer 

system (through comparison with each adjacent sea 

area of Japan); (4) Dredging of sediments over the 

polluted zones. 

Nalewajko, C. and M. A. O'Mahony (1989) 

Photosynthesis of algal cultures and 

phytoplankton following an acid pH 

shock. Journal of Phycology 25:3 19-325. 

{pH; acid; lake; phytoplankton; photosynthesis) 

In acidifying lakes, pH decreases abruptly in 

response to acid precipitation events. We tested the 

hypothesis that in comparison to a circumneutral 

lake, phytoplankton photosynthesis in an 

acidifying lake is less sensitive to a rapid decrease 

in pH (acid pH shock). 

Phytoplankton in Plastic Lake, which is 

undergoing acidification, was characterized by a 

predominance of Pyrrophyta, and phytoplankton 

photosynthesis decreased to a lesser extent in 

response to an acid pH shock than the 

photosynthesis of populations from St. Nora Lake, 

a circumneutral lake located nearby, in which 

Pyrrophyta were not abundant. Rates of 

phytoplankton photosynthesis in acid pH shock 

experiments were significantly correlated with 

hydrogen ion but not with dissolved inorganic 

carbon (DIC) concentrations. 

Depression of photosynthesis following an acid 

pH shock occurred in axenic cultures of Chlorella 

pyrenoidosa Chick but was not observed in axenic 

cultures of the acidophilic alga Chlorella 

saccharophila (Krug.) Nadson or in three species 

isolated from Plastic Lake. However, the three 

isolates were not acidophilic during growth. We 

conclude that phytoplankton in acidifying lakes 

consists predominantly of species which are 

tolerant to acid pH for short periods (hours) but 

cannot grow at these pHs. 

Nehring, D. (1981) Phosphorus in the Baltic 

Sea. Marine Pollution Bulletin 12: 194-198. 

{Baltic Sea; phosphorus; Gotland Sea) 

No abstract 

Nehring, D. (1984) The further development 

of the nutrient situation in the Baltic 

proper. Ophelia Supp1.3: 167- 179. 

{nitrogen; phophorus; nutrient; Baltic; 

hydrographic) 

Long-term nutrient trends were studied in the deep 

water (in general 1958-1982) and in the winter 

surface layer (in general 1958-1983) of the Baltic 

proper. Phosphate accumulated in the deep water, 

on average, over the whole period under 

investigation, whereas nitrate has decreased 

significantly since 1979. The concentrations of 

these nutrients also increased in the winter surface 

layer. The highly significant overall trends in this 

layer correlated closely with an increase in salinity. 

The reasons for the long-term variations in 

nutrients and the consequences of the eutrophication 

observed are briefly discussed. 

Nehring, D. (1992) Eutrophication in the 

Baltic Sea. In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{nitrogen, phosphorus, oxygen, Baltic Sea) 

Eutrophication is identified as one of the most 

serious problems in the Baltic Sea. Trend studies 

covering the period 1958-89 show increasing 

phosphate and nitrate concentrations in the surface 

layer in winter and in the deep water below the 

permanent halocline. Consequences of the 

eutrophication are discussed with respect to the 

biological productivity and the determination of the 

oxygen conditions in stagnant Baltic deep waters. 

Conditions for eutrophication in the Baltic Sea are 

compared with those in the Mediterranean Sea. 

Nehring, D., S. Schulz, and W. Kaiser (1984) 

Long-term phosphate and nitrate trends in 

the Baltic Proper and some biological 

consequences: A contribution to the 

discussion concerning the eutrophication 

of these waters. Rappots et Procb-verbaux des 

Re'unions 

Counseil international pour LIExploration de la 

Mer 183: 193-203. 

{Baltic; nutrients; nitrogen; phosphorus; fish; 

oxygen; primary production; phytoplankton; 

zooplankton) 

The phosphate and nitrate levels in the winter 

surface layers of the central Arkona, Bornholm, and 

southern Gotland Seas roughly tripled from 1969 to 

1978 (phosphate: 0.2 to 0.6 umol 1-l; nitrate: 1.5 

to 4.0 umol 1-I). In the deep water below the 

halocline, long-term accumulation of these 

nutrients was also found in the Gda'nsk, Gotland, 

FArS, and Landsort Deeps. This process was most 

significant at a depth of 100 m in the Gotland 

Deep, where on average phosphate increased from 

about 1 to 2.5 umol 1-I between 1958 and 1978, 

and nitrate from about 4 to 8 umol 1-I between 

1969 and 1978. 
The biological effects of the nutrient increase on 

the pelagic system of the Baltic are obviously low 

and masked by inadequate observation frequency or 

by annual fluctuations. On the other hand the 

increasing yields of the Baltic fisheries do not seem 

to be the result of the growing fishing effort alone 

but are the consequence of higher biological 

production. 

Neilson, B. J. and L. E. Cronin (ed.) (1981) 

Estuaries and Nutrients. Humana Press, 

Clifton, New Jersey. 

No abstract 

Nielsen, A. and G. Ertebjerg (1984) Plankton 

blooms in Danish waters. Op h e 1 ia, 

Supplementum 3:181-188. 

{phytoplankton; primary production; nitrogen; 

bloom) 

In the past fifteen years there seems to have been 

an increase in the occurrence of massive plankton 

blooms along the coast of the North Atlantic 

Ocean, including the North Sea, the Kattegat and 

the Belt Sea. Some of the plankton blooms have 

caused fish mortality, either directly by poisoning 

of the fish, or indirectly by suffocation due to 

oxygen deficiency. It is indicated in the paper that 

the increased pollution load on the coastal zone 

may play an important role in the development of 

massive plankton blooms in Danish waters. 

Niemi, A. (1972) Observations on 

phytoplankton in eutrophied and noneutrophied archipelago waters of the 

southern coast of Finland. Memoranda 

Societatis pro. Fauna et Flora Fennica 48:63-74. 

{phytoplankton; Finland; biomass; composition) 

A comparison is made of the summer course of 

the biomass and the composition of the 

phytoplankton populations of 3 archipelago zones 

in clean and polluted conditions off the southern 

coast of Finland. In summer low phytoplankton 

biomass follow a vigorous spring bloom in the 

oligotrophic areas. Strongly eutrophic waters are 

characterized by high biomasses (primarily due to 

the blue-green alga Oscillatoria agardhii) persisting 

throughout the summer. In addition to 

phytoplankton biomass, the composition of 

phytoplankton and the occurrence of certain 

indicator group may be a good criteria of the quality 

and level of eutrophication. 

Nienhuis, P. H. (1983) Temporal and spatial 

patterns of eelgrass (Zostera marina L.) 

in a former estuary in the Netherlands, 

dominated by human activities. Marine 

Technology Society Journal 17:69-77. 

{eelgrass; submerged vegetation; Netherlands) 

A review is given of the temporal and spatial 

patterns of the eelgrass population in Grevelingen 

estuary, SW Netherlands, after it has been 

transformed into a salt-water lake. In summer 1978, 

after a development of 8 years of opportunistic 

colonization by seeds and rhizomes, all potential 

habitats in the lake were occupied and covered 44 

km2 of sheltered sediment areas with a water depth 

of 0-2.5 m. After man-induced changes in 

environmental conditions the eelgrass population 

dropped down in 1980 to about half the 1978 

situation. The possible causes of this sudden 

decline have been discussed, viz seed production and 

germination, water temperature, wasting disease, 

transparency of the water and macro-algal 

competition. The most plausible explanation for 

mass decline of eelgrass in Lake Grevelingen in 

1980 is associated with an increase in organic 

matter deposition on the bottom, following a 

change in nutrient loading of the lake, causing rapid 

deoxygenation and toxification of the sediments, 

and consequent die back of rhizomes and roots. 

Water temperature and salinity may have played a 

minor role in the eelgrass wax and wane. 

Nilsson, P., B. Jonsson, I. L. Swanberg, and K. 

Sundback (1991) Response of marine 

shallow-water sediment system to an 

increased load of inorganic nutrients. 

Marine Ecology Progress Series 71:275-290. 

{sediment; nutrients; microflora; production; 

biomass; composition; productivity; meiofauna; 

bacteria; grazing; oxygen) 

An outdoor experimental system was used to 

investigate the effect of an~increased load of 

inorganic nitrogen and phosphorus on the lower 

trophic levels of the food web of a shallow-water 

sandy sediment. Estimates of structural changes 

were based on relationships between the biomass of 

autotrophs, heterotrophic bacteria and meiofauna 

and their qualitative composition. Effects on the 

functions of the sediment community were assessed 

by measuring primary and bacterial productivity and 

meiofaunal grazing rates using radiolabelling, as 

well as measuring changes in oxygen and nutrient 

fluxes and carbon pools. The sediment system 

responded within 2 to 3 wk to the nutrient 

enrichment. Meiofauna biomass increased, resulting 

in higher relative importance of oligochaetes and 

harpacticoid copepods. Primary productivity 

increased faster than meiofaunal grazing, resulting 

in an increase of microalgal biomass by a factor of 

4. Diatom and filamentous cyanobacteria were 

favoured by the increased nutrient levels. The 

stimulated photosynthetic activity has a negative 

feedback on the producing sediment layer, which 

was lifted off by oxygen bubbles entrapped in the 

mucus-rich top most layer. Stimulated growth of 

the filamenous green alga Enteromorpha clathrata 

resulted in a biomass of ca 2 g C m-2 after 4 wk, 

which was more than 2 times the biomass of 

rnicroautotrophs living in the sediment. Bacterial 
energy transfer because of a small Coriolis effect at 

low latitudes. 

Nixon, S. W. (1989) An extraordinary red 

tide and fish kill. In: E. M. Cosper, V. M. 

Bricelj and E. J. Carpenter (ed.) Nov e 1 

Phytoplankton Blooms Causes and Impacts of 

Recurrent Brown Tides and Other Unusual Blooms. 

Springer-Verlag, Berlin. 

{Narragansett Bay; red tide; sewage) 

No abstract 

Nixon, S. W. (1990) Quanifying the 

relationship between nitrogen input and 

the productivity of marine ecosystems. 

Eighth Marine Technology Conference and 

International Symposium for Ecology, Tokyo, 

August 1990, and Shimane, September 1990, 

Japan. 

{nutrients; productivity; relationship; estuary; 

nitrogen) 

The concept that the input of dissolved inorganic 

nutrients, especially nitrogen, regulated the basic 

level of primary production and, hence, the yield of 

fish from marine ecosystems developed in Western 

Europe and Scandinavia about a century ago. This 

"agricultural model" of marine production was 

conceived only after quantitative biological 

sampling methods and analytical techniques for 

measuring nutrients in the sea became available 

during the late 1800s and early 1900s. 

Verification of the model has been slow because 

of difficulties in measuring the rate of nutrient 

input to marine areas, problems in measuring 

primary production, lack of long-term 

comprehensive data, a system-specific view among 

researchers that discourages broad comparisons, and 

a reluctance to undertake ecosystem-level 

experiments in marine ecology. Widespread use of 

the 14c technique after about 1950, recent efforts 

to develop nutrient budgets, and the completion of 

a long-term nutrient addition experiment using 

marine mesocosms, have now provided data used in 

this report to quantify the broad, general 

relationship between primary production and 

fisheries yields from the sea have only been 

assembled in the past few years, but the evidence is 

now compelling. Despite the great complexity of 

marine ecosystems, the simple first-order 

relationship between nutrient inputs and 

productivity that was first conceived a century ago 

seems generally correct. 

Nixon, S. W. (1992) Quantifying the 

relationship between nitrogen input and 

the productivity of marine ecosystems. 

Advanced marine Technology Conference, NOS, 

Japan, The concept that the input of dissolved 

inorganic nutrients, especially nitrogen, regulated 

the basic level of primary production and, hence, 

the yield of fish form marine ecosystems developed 

in Western Europe and 

Nixon, S. W. and V. M. Berounsky (1984) The 

role of nitrification in contributing to 

low oxygen conditions in an urban 

waterway. Rhode Island Water Resources Center, 

FY-1983. 

{nitrification; oxygen; Providence River) 

No abstract 

Nixon, S. W., J. R. Kelly, B. N. Furnas, C. A. 

Oviatt, and S. S. Hale (1980) Phosphorus 

regeneration and the metabolism of 

coastal marine bottom communities. In: 

K. R. Tenore and B. C. Coull (ed.) Marine Benthic 

Dynamics. University of South Carolina Press, 

Columbia, South Carolina. 

{phosphate; Narraganett Bay; flux; sediment) 

In situ measurements of the exchange of 

phosphate between sediment and the overlying 

water at three stations in Narragansett Bay, Rhode 

Island, showed that there was almost always a net 

flux out of the sediments. The magnitude of the 

flux ranged from near zero in winter to almost 60 

umols m-2 hr-l in summer. The flux was strongly 

correlated with temperature during the spring 

warming and did not decrease with increasing 

phosphate concentrations in the overlying water. 

Calculations indicate that the phosphate cycle in 

coastal waters such as Narragansett Bay is 

dominated by sediment-water exchanges. Some 120 

mg-at of inorganic P m-2 yr-l were released 

annually to the overlying water, or enough 

phosphorus to support about 50% of the annual 

phytoplankton primary production. The flux of 

dissolved organic phosphorus was erratic and lower, 

and appreciable uptake as well as release was often 

observed. 

Laboratory experiments using replicate cores 

collected from the bay at different times of year 

showed that oxygen uptake, carbon dioxide release, 

and phosphorus exchange by the sediments were 

also influenced by the availability of fresh organic 

matter. It was clear from the laboratory and field 

measurements that the regeneration of organic 

matter by the benthos results in a return of 

inorganic nutrients to the water column that is 

anomalously low in fixed nitrogen relative to 

phosphorus. This remarkable feature of benthic 

regeneration, along with the fact that a large 

amount of organic matter is decomposed on the 

bottom in shallow ateas compared with the open 

sea, appears to be responsible for the 

characteristically low N/P ratio of coastal marine 

waters and for the importance of nitrogen rather 

than phosphorus as a majot limiting nutrient in 

these areas. 
Nixon, S. W., C. A. Oviatt, J. Frithsen, and B. 

Sullivan (1986) Nutrients and the 

productivity of estuarine and coastal 

marine ecosystems. Journal of Limnological 

Society of Southern Africa 12:43-7 1. 

{nutrients; primary productivity; estuary; coastal; 

microheterotrophs; recycling; MERL) 

Recent research on estuarine and coastal marine 

ecosystems has revealed two particularly interesting 

things about nutrient and productivity. First is the 

obgervation that these areas are among the most 

intensively fertilized environments on earth. Second 

is the common finding that much of the 

characteristically high primary productivity of these 

shallow waters is supported by nutrients released or 

recycled by pelagic and benthic microheterotrophs. 

Since nutrient inputs to coastal areas have probably 

been increasing and are likely to continue to do so, 

it is particularly important to understand the 

relationship between nutrient loading and nutrient 

cycling and the extent to which their interactions 

may set the levels of primary and secondary 

production in coastal systems. 

That some direct relationship exists between the 

input of nutrients and the productivity of higher 

trophic levels has been a principle of marine 

ecology since the turn of the century. It is 

surprisingly difficult, however, to find quantitative 

evidence showing that estuaries, lagoons, or other 

coastal waters respond to eutrophication by 

producing a larger biomass of animals. Part of this 

difficulty arises because the amount of nitrogen or 

phosphorus incorporated in animal tissue is a very 

small term in the total nutrient budget of an 

estuary, and the accuracy and precision of ecological 

field measurements may not be adequate to the task. 

In addition, the response of natural systems to 

nutrient enrichment is compounded by changes in 

climate, hydrography, harvesting effort and 

technology, and pollution. 

Attempts to avoid these problems by carrying out 

controlled nutrient addition experiments in the field 

or with mesocosms have been much rarer in marine 

ecology than limnology. The results that are 

available for such studies seem to suggest that there 

is a modest enhancement of primary production 

with nutrient addition, but that most of this extra 

organic matter is rapidly consumed, presumably by 

microheterotrophs. In other words, as nutrient 

inputs rise, so does the rate of nutrient recycling. 

Only a small fraction of the added nutrients appears 

as an increment in the production of higher trophic 

levels. 

Nixon, S. W., C. A. Oviatt, J. Garber, and V. Lee 

(1976) Diel metabolism and nutrient 

dynamics in a salt marsh embayment. 

Ecology 57:740-750. 

{diel; dissolved oxygen; metabolism; nitrogen; 

nutrients; phosphorus; salt marsh; silica; 

Narragansett Bay) 

Simultaneous measurements of the diel patterns 

of dissolved 02, pH-C02, dissolved organic 

nitrogen, ammonia, nitrite, nitrate, dissolved 

organic phosphorus, phosphate, and silicate were 

made on four occasions in a shallow salt marsh 

embayment (Bissel Cove, Narragansett Bay, Rhode 

Island, U.S.A.) that was closed to tidal water 

exchanges for 24-h periods. In spite of high rates of 

community photosynthesis and respiration, there 

appeared to be little diel exchange in dissolved 

nutrient concentrations, especially with respect to 

inorganic nitrogen. During summer and early fall, 

inorganic nutrients appeared to cycle within the 

sediment-detritus system of the embayment 

bottom, rather than being released to the overlying 

water. This suggest that there would be little export 

of inorganic nutrients from the marsh during tidal 

exchanges with the estuary. The behavior of this 

marsh embayment contrasts with reports from other 

marsh areas where large net imports or exports of 

nutrients have been observed. The concentrations of 

dissolved organic phosphorus and nitrogen were 

higher than inorganic forms, and showed 

substantial variation over each diel sampling 

period. The relationship between photosynthetic or 

respiratory gas exchange and nutrient cycling in 

waters influenced by benthic community 

metabolism is subtle and complex, and cannot be 

adequately approximated by simple models such as 

the Redfield ratio. 

Nixon, S. W. and M. E. Q. Pilson (1983) 

Nitrogen in estuarine and coastal marine 

ecosystems. In: E. J. Carpenter and D. G. 

Capone (ed.) Nitrogen in the Marine Environment. 

Academic Press, New York. 

{nitrogen; estuary; budget; distribution) 

No abstract 

Nixon, S. W. and M. E. Q. Pilson (1984) 

Estuarine total system metabolism and 

organic exchange calculated from nutrient 

ratios: An example from Narragansett 

Bay. In: V. S. Kennedy (ed.) The Estuary as a 

Filter. Academic Press, Orlando. 

{Narragansett Bay; estuary; nutrient; denitrification; 

net ecosystem metabolism; phosphorus; budget) 

The average ratio of the concentration of dissolved 

inorganic nitrogen (DIN) to phosphorus (DIP) 

found in many estuaries differs from that in the 

rivers, sewage, rain, and offshore waters that 

contribute nutrients to these systems. These 

differences are largely the result of various 

biological processes which take place in the 

estuary, including denitrification, the net production 

of organic matter, and the consumption and 

remineralization of organic matter carried into the 
estuary. Net production in this case represents the 

total amount exported, harvested, and removed 

through long-term burial that is in excess of the 

amount imported and consumed within the estuary. 

It is not the same as the commonly reported net 

production by the autotrophic components of the 

system. Together with physical mixing and 

chemical exchange processes which take place in 

the system, these biological transformations 

provide a dynamic "filter" which often makes the 

quantity and form of nutrients exported from an 

estuary quite different from those it imports. These 

changes can themselves provide insight into the 

internal processes of the estuary. 

For Narragansett Bay (RI, USA) we have 

combined measurements of the annual nutrient 

input with the ratio of the annual time and volumeweighted mean concentrations of DIN and DIP 

found in the Bay, measurements of the annual rate 

of denitrification, and data on the chemical 

composition of the plankton to calculate the total 

system metabolism of the Bay. The result suggests 

that the system is autotrophic, with net production 

exceeding consumption by about 80 g C m-2 y-l. 

Since only a small portion of this organic matter 

accumulates in the sediments (- 6 g C m-2 y-l) or 

is removed in the fishery (- 1 g C m-2 y-l), some 

70-75 g C m-2 y-l may be exported from the Bay, 

an amount equal to 22-24% of the reported 

production by the phytoplankton. 

Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P. 

Donaghay, B. Sullivan, S. Seitzinger, D. Rudnick, 

and J. Frithsen (1984) Eutrophication of a 

coastal marine ecosystem - An 

experimental study using the MERL 

microcosms. In: M. J. R. Fasham (ed.) Flows 

of Energy and Materials in Marine Ecosystems. 

Plenum Publishing Co., New York. 

{nutrients; phytoplankton assemblage; 

phytoplankton abundance; secondary production; 

MERL) 

No abstract 

NOAA (1990) Estuaries of the United 

States 

Vital statistics of a national resource 

base. NOAA, A special NOAA 20th anniversary 

report, {estuary; resource; review) 

No abstract 

Nowicki, B. L. and S. W. Nixon (1985) Benthic 

nutrient remineralization in a coastal 

lagoon ecosystem. Estuaries 8: 182- 190. 

{ammonia; nitrate; nitrite; phosphate; phosphorus; 

lagoon; Narragansett Bay; flux; N/P ratio) 

In situ measurements of the exchange of 

ammonia, nitrate plus nitrite, phosphate, and 

dissolved organic phosphorus between sediments 

and the overlying water column were made in a 

shallow coastal lagoon on the ocean coast of Rhode 

Island, U.S.A. The release of ammonia from mud 

sediments in the dark (20-440 umol per m2 per h) 

averaged ten times higher than from a sandy tidal 

flat (0-60 umol per m2 per h), and while mud 

sediments also released nitrate and phosphate, sandy 

sediments took up these nutrients. Fluxes of 

nutrients from mud sediments, but not from sandy 

areas, markedly increased with temperature. 

Ammonia release rates for mud sediments in the 

light (0-350 umol per m2 per h) were lower than 

those in the dark and it is estimated that some 25% 

of the ammonia released to the water column on an 

annual basis may be intercepted by the benthic 

microfloral community. Estimates of the annual 

net exchange of nutrients across the sediment-water 

interface, weighted by sediment type for the lagoon 

as a whole, showed a release of 450 mmol per m2 

of ammonia, 5 mmol per m2 of phosphate, 5 

mmol per m2 of dissolved organic phosphorus, and 

an uptake of 80 mmol per m2 of nitrate. Although 

rates of ammonia and nitrate exchange were 

comparable to those described for the deeper 

heterotrophic bottom communities of nearby 

Narragansett Bay, rates of benthic phosphate release 

were significantly lower. On an annual basis the 

Bay benthos released approximately 20 times more 

inorganic phosphate per unit area than did the 

lagoon benthos. As a result, the NIP ratio for the 

flux from the sediments was 74:l in the lagoon, 

compared with 16:l in "average" marine plankton 

and 8: 1 for the benthic flux from Narragansett Bay. 

The lack of remineralized phosphate in the lagoon 

is reflected in water column phosphate 

concentrations (always <lum) and water column 

NIP ratios (anual N/P = 27) and suggests that the 

lagoon may show phosphate limitation rather than 

the nitrogen limitation commonly associated with 

marine systems. 

O'Connor, D. J. (1981) Modeling of 

eutrophication in estuaries. In: B. J. 

Neilson and L. E. Cronin (ed.) Estuaries and 

Nutrients. Humana Press, Clifton, New Jersey. 

{nutrient; model; phytoplankton; estuary) 

No abstract 

O'Reilly and D. A. Busch (1984) 

Phytoplankton primary production on the 

northwestern Atlantic shelf. Rappots et 

Procb-verbaux des Riunions 

Counseil International pour LfExploration de la 

Mer 183:255-268. 

{Atlantic shelf; phytoplankton; primary 

production; sewage) 

Phytoplankton production of particulate and 

dissolved organic carbon was measured using 14c 

simulated in situ sunlight incubation method 
during 23 survey of the Mid-Atlantic Bight, 

Georges Bank, and the Gulf of Maine. Annual 

phytoplankton production (particulate plus 

dissolved) ranged between 260 and 470 g C m-2 yrin various regions, which places this continental 

shelf system among the most productive in the 

world. The highest rates of daily phytoplankton 

production were consistently found off the coast of 

New Jersey in the sewage-polluted apex of the New 

York Bight, followed by the shallow, well-mixed 

waters on Georges Bank. In general, high daily 

rates of primary production (1 g C m-2 yr-l) were 

observed during most of the months sampled and 

were not limited to the 'spring bloom' period. 

Available data indicate that microheterotrophs 

above the thermocline (not the seabed) supply most 

of the mineralized nutrients required by the 

productive summer phytoplankton communities. 

Size-fractionation of the 14c-labeled particulate 

organic carbon revealed that netplankton (> 20 um) 

are major primary producers during the spring and 

fall blooms in the Mid-Atlantic Bight, on Georges 

Bank, and in the Gulf of Maine; however, the 

nanoplankton (< 20 um) are responsible for most 

of the annual photosynthesis of organic carbon. On 

the annual basis, the percentage of particulate 

carbon productivity by nanoplankton increased from 

the shallow to the deep water. Euphotic per cent 

extracellular release (PER) of dissolved organic 

carbon by phytoplankton ranged between 0 and 55 

%, averaging 15 %, and increased from the shallow 

to the deep waters of the continental shelf. No welldefined seasonal cycle in euphotic PER was evident 

from our data. 

Ochi, T. (1987) The development of anoxic 

water and red tide associated with 

eutrophication in Hiuchi Nada Inland 

Sea, Japan. In: T. Okaichi, D. M. Anderson 

and T. Nemoto (ed.) Red Tides: Biology, 

Environmental Science, and Toxicology. Elsevier, 

New York. 

{red tides; oxygen) 

The mechanism of anoxic water mass formation 

was investigated in the eastern part of Hiuchi-Nada. 

It was found that the oxygen consumption by the 

sediment was unexpectedly small and that the 

oxygen consumption by organic matter freshly 

deposited on the bottom from phytoplankton 

blooms, followed by frequent breakdown of the 

thermocline were important contributing factors in 

the formation of anoxic water masses in HiuchiNada. 

Officer, C. B. and J. H. Ryther (1980) The 

possible importance of silicon in marine 

eutrophication. Marine Ecology Progress 

Series 3:83-91. 

{silicon; nitrogen; phosphorus; evidence; 

phytoplankton; diatoms; flagellates) 

Diatom phytoplankton populations are the usual 

food for zooplankton and filter feeding fishes and 

contribute in a direct way to the large fishable 

populations in coastal zones. Flagellates, on the 

other hand, are frequently poor foods for most 

grazers and can lead to undesirable eutrophication 

effects. Arguments are presented that silicon is 

often the controlling nutrient in altering a diatom 

to a flagellate community. The alteration is 

governed by the relative magnitudes of the natural 

fluxes of the nutrients nitrogen, phosphorus and 

silicon to the receiving water body and the recycled 

fluxes of nitrogen and phosphorus for oceanic, 

estuarine and inland water bodies. 

Okaichi, T. (1987) Red tide problems in the 

Seto Inland Sea, Japan. In: T. Okaichi, D. 

M. Anderson and T. Nemoto (ed.) Red Tides: 

Biology, Environmental Science, and Toxicology. 

Elsevier, New York. 

{red tides; blooms; oxygen; abundance) 

With the development of industries along the 

coast of Seto Inland Sea, Japan, aquaculture of 

yellowtail, oyster and nori (laver) have been 

suffering from outbreaks of red tides. During these 

two decades 20 billion yellowtail worth about 20 

billion yen were killed. Comprehensive surveys on 

the cause of outbreaks of red tides are carried out by 

workers at Universities and Prefectures. 

Countermeasures to avoid the fishkill are devised 

and the monitoring systems for red tides, PSP and 

DSP within Prefectures are organized by the Bureau 

of fisheries. 

Okaichi, T., D. M. Anderson, and T. Nemoto (ed.) 

(1988) Red Tides: Biology, Environmental 

Science, and Toxicology (Proceedings of 

the International Symposium on Red 

Tides, Takamastu Perfecture, Japan, 10- 

14 November, 1987). Elsevier, New York. 

No abstract 

Olenin, S. N. (1990) Benthic "desert" and 

transitional ecological zone at the 

bottom of the eastern part of the Baltic 

Sea. Oceanology of the Academy of Sciences of 

the USSR 29:751-753. 

{benthic; desert; Baltic; transition; macrofauna) 

As a result of studies performed during 198 1- 1987 

on the slope and floor of the Gotland basin in the 

eastern Baltic Sea, a transitional ecological zone 

was distinguished between the benthic "desert", 

which is devoid of macrofauna and situated on the 

floor and slope of the basin below 110-120 m, and 

the upper sections of the benthic zone above 80-90 

m, which has a comparatively rich and varied 

bottom populations. The zoobenthos of the 

transitional zone is extremely poor and is 
characterized by a predominance of mobile 

invertebrate forms. 

Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst 

(1984) Ganyaulax excavata on it or in^ in 

New Jersey. Coastal Ocean Pollution 

Assessment News 3:25-26. 

{nuisance bloom; Gonyaulax; red tide; nutrient; 

New York Bay; New Jersey) 

No abstract 

Olsson, G. E., B. Sundstrom, and L. Edler (1987) 

In situ studies of the effects of humic 

acids on dinoflagellates and diatoms. In: 

T. Okichi, D. M. Anderson and T. Nemoto (ed.) 

Red tides: Biology, Environmental Science, and 

Toxicology. Elsevier, New York. 

{blooms; humic) 

Experiments with 160 1 plastic bags were 

performed in September 1986 in situ on the west 

coast of Sweden. One series of bags were filled 

with water containing the natural phytoplankton 

community dominated by dinoflagellates (~80% of 

the total phytoplankton biomass). The other series 

was filled with 1 um filtered sea water inocculated 

with the diatom Phaeodactylum tricornutum. 

Phosphorus or nitrogen was added alone or together 

with commercially available humic acids to both 

series. Bags with no additions (C) or with only 

humic acids added (CH) were used as controls. P. 

tricornutum was stimulated by nitrogen addition, 

while no positive response was obtained by any 

nutrient addition to the dinoflagellate bags. 

Dinoflagellate biomass increased several fold when 

humic acids were added together with nitrogen. 

Also the alkaline biomass phosphatase activity 

(APA) increased drastically both in the CH and 

nitrogen + humic acid bags. Tentative conclusions 

are: 1) The dinoflagellates were inhibited by heavy 

metals and the humic acids alleviated this 

inhibition by working as chelating agents. 2) The 

diatom was not inhibited by heavy metals because 

it could still grow quite well without humic acids. 

3) As APA increased in the dinoflagellate control 

bag with only humic acids we assume that nitrogen 

in the humic fraction to some extent was available 

for the dinoflagellates. 

Oviatt, C. A., A. A. Keller, P. A. Sampou, and L. 

L. Beatty (1986) Patterns of productivity 

during eutrophication: A mesocosm 

experiment. Marine Ecology - Progress Series 

28:69-80. 

{oxygen; net ecosystem production; MERL) 

In a 28 months mesocosm experiment, levels and 

patterns of productivity and respiration were 

observed for a range of nutrient additions selected to 

provide a gradation from conditions in lower 

Narragansett Bay, Rhode Island, USA, to 

maximum impact for an urban estuary receiving 

present day levels of sewage effluents. For a 32-fold 

increase in nutrients, system apparent production 

increased by only a factor of 3.5. Seasonal patterns 

of autotrophy during the winter-spring diatom 

bloom and heterotrophy during summer and early 

fall occurred at all treatment levels. With the 

exception of the 8X treatment, all treatments above 

2X had a greater respiratory demand in the water 

column than the benthos. The highest treatment 

mesocosm (32X) went briefly anoxic during the 

second summer of the experiment during a period 

when little productivity was occurring in the water 

column. 

Oviatt, C. A., P. Lane, F. F. 111, and P. Donaghay 

(1988) Phytoplankton species and 

abundance in response to eutrophication 

in coastal marine mesocosms. Jaurnal of 

Plankton Research 11: 1223- 1244. 

{nutrient; species composition; phytoplankton; 

assemblage; abundance; MERL) 

In a mesocosm nutrient enrichment experiment 

the species (or categories) and abundances of 

diatoms, dinoflagellates, flagellates, monads and 

ciliates were identified and counted over a 16 month 

period. Diatoms and ciliates increased with 

increasing nutrient treatment while monads and 

flagellates, less than 10 urn, did not. In some 

instances grazing controlled abundances to low 

levels in nutrient enriched treatments. By contrast, 

in the field diatoms sometimes appeared to decrease 

while small phytoplankton less than 10 um 

appeared to increase under eutrophic conditions. 

While nuisance species were occasionally present in 

various nutrient treatments, the intensity and 

frequency of their presence did not tend to increase 

with nutrient treatment. Generally species (or 

categories) did not appear to change with nutrient 

treatment. 

Oviatt, C. A., J. G. Quinn, J. T. Maughan, J. T. 

Ellis, B. K. Sullivan, J. N. Gearing, P. I. Gearing, 

C. D. Hunt, P. A. Sampou, and J. S. Latimer 

(1987) Fate and effects of sewage sludge 

in the coastal marine environment: A 

mesocosm experiment. Marine Ecology 

Progress Series 41: 187-203. 

{production; respiration; fate; nutrients; sludge; 

oxygen; grazing; MERL) 

A mesocosm experiment to assess the fate and 

effects of sewage sludge in the coastal marine 

environment was conducted over 4 mo. during the 

summer of 1984 with 9 different treatments of 

sewage sludge and nutrient additions. Evidence from 

settling rate studies, accumulations of carbon and 

pollutant organics and ratios of 813c in the 

sediment indicated up to 83% of the sludge. 

particulate inputs settled to the bottom and up to 

51% accumulated there. While no toxicity due to 

organics or metals was apparent, prolonged hypoxia 
and even anoxia occurred in upper treatment levels. 

At summer temperatures, sludge particulate inputs 

in excess of 1 g C mm2 d-l caused these hypoxic 

conditions. In contrast to low primary production 

in low and medium sludge treatments during the 

last 2 mo. of the experiment, the highest sludge 

treatment had high production. We attribute these 

patterns to excessive grazing pressure from 

zooplankton and benthic fauna in the lower 2 

treatments and hypoxia-retarded grazing in the upper 

treatment. In the upper treatments, peaks of 

production stimulated peaks of excessive respiration 

suggesting that respiration of sludge particulates 

was enhanced or co-metabolized with the production 

of fresh organic material. 

Paerl, H. W. (1988) Nuisance phytoplankton 

blooms in coastal, estuarine, and inland 

waters. Limnology and Oceanography 33:823- 

847. 

{bloom; phytoplankton; marine; estuary; 

freshwater; cyanobacteria; dinoflagellates; review) 

Multiple interacting physical, chemical, and 

biotic factors, in proper combinations lead to the 

development and persistence of nuisance algal 

blooms. Upon examining combinations of 

environmental conditions most likely to elicit 

nuisance blooms, commonalities and analog 

situations become more apparent among coastal 

marine (dinoflagellate-dominated), 

estuarine(dinoflagel1ate- and cyanobacteriadominated), and freshwater (cyanobacteriadominated) ecosystem. A combination of the 

following hydrological, chemical, and biotic factors 

will most likely lead to bloom-sensitive waters: a 

horizontally distinct water mass; a vertically 

stratified water column; warm weather conditions, 

as typified by dry monsoon tropical climates and 

summer seasons in temperate zone; high incident 

photosynthetically active radiation (PAR); enhanced 

allochthonous organic matter loading (both as DOC 

and POC); enhanced allochthonous inorganic 

nutrient loading (nitrogen and/or phosphorus); 

adequate availability of essential metals, supplied 

by terrigenous inputs or upwelling; underlying 

sediments physically and nutritionally suitable as 

"seed beds" for resting cysts and alkinetes; algalmicrograzer (protists and rotifers) synergism, which 

also enhances nutrient cycling without 

consumption of filamentous and colonial nuisance 

taxa; and selective (for non-nuisance taxa) activities 

of macrograzers (crustacean zooplankton, larval 

fish). 

Nuisance bloom taxa share numerous additional 

physiological and ecological characteristics, 

including limited heterotrophic capabilities, high 

degrees of motility, and toxicity. Given such a set 

of commonalities, it would appear useful and 

timely to identify and address generally applicable 

criteria for deeming a water body "bloom sensitive" 

and to incorporate such criteria into the design of 

water quality management strategies applicable to 

both coastal marine and freshwater habitats. 

Pagou, K. and L. Ignatiades (1988) 

Phytoplankton seasonality patterns in 

eutrophic marine coastal waters. Biological 

Oceanography 5229-241. 

{phytoplankton; seasonality; Aegean Sea) 

Phytoplankton seasonality patterns were examined 

in an inshore eutrophic station fertilized 

continuously by domestic sewage. Data of certain 

physical (temperature), chemical (P-P04, N-N03, 

and Si-SI02 concentrations), and phytoplanktonic 

(chl a, total cell, diatom, and dinoflagellate 

concentrations) parameters were analyzed by time 

series analysis (autocorrelation, cross correlation). 

The phytoplanktonic and chemical parameters 

exhibited significant seasonal variability-in spite of 

the constant fertilization of the water-which was 

associated with the seasonal variability of 

temperature. 

Parker, C. A. (1983) Localized hypoxia 

recurs in the New York Bight. Coastal 

Ocean Pollution Assessment News 3: 1-3. 

{oxygen; New York Bight) 

No abstract 

Parker, C. A. and J. E. O'Reilly (1991) Oxygen 

depletion in Long Island Sound: A 

historical perspective. Estuaries 14:248-264. 

{oxygen; hypoxia; historical; Long Island Sound) 

A retrospective analysis of available data was 

conducted to characterize the spatial distribution and 

temporal trends in dissolved oxygen (DO) 

concentrations in Long Island Sound (LIS) over the 

past four decades. A general east-west gradient of 

decreasing bottom DO was evident in all historical 

data examined. In our review of data from the 1950s 

, collected by Gordon Riley and colleagues, and 

from contemporary surveys, we found no evidence 

of hypoxia (DO I 3.0 mg 1-l) in the Eastern Basin; 

however, in the deeper waters of the Central Basin, 

there is some evidence for recent (1986) emergence 

of moderate hypoxia. The Western Basin 

experienced episodes of hypoxia during the 1970s 

which became more recurrent and possibly more 

severe in the late 1980s. The most severe, 

persistent and chronically recurrent hypoxia 

occurred throughout the water column of the East 

River and in bottom water of the Western Narrows. 

An unprecedented episode of anoxia was observed in 

both the Western and Eastern Narrows regions of 

LIS in 1987. Previously, anoxia occurred rarely, 

was short-lived, and was confined to the East River. 

Statistical trend analyses revealed a significant 

increase in the summer minimum bottom DO in 

the lower and middle reaches of the East River over 

the past 20 years. Beginning in 1981, however, DO 
declined markedly in the adjacent Narrows bordering 

the Nassau County nearshore. The improvements 

in East River water quality over the previous 15-20 

years appear to have been gained at the expense of 

poorer water quality in the western sound. 

Mechanisms potentially responsible for the recent 

decline the bottom DO in western LIS are 

suggested. 

Pastorok, R. A. and G. R. Bilyard (1985) 

Effects of sewage pollution on coral-reef 

communities. Marine Ecology Progress Series 

21: 175- 189. 

{sewage; coral reef; lagoon; species composition; 

abundance; benthic algae; primary production; 

nutrient; oxygen; Kaneohe Bay; turbidity; 

sedimentation) 

Sewage pollution is an increasing problem in 

tropical marine environments. In this review we 

synthesize present knowledge of the effects of 

sewage pollution on coral-reef communities, and 

suggest directions for future research. A wide range 

of sewage impacts on coral-reef communities has 

been reported. Little or no impact has been 

observed on some reefs in well-flushed waters that 

receive small quantities of effluent, whereas large 

discharges of effluent into poorly-flushed lagoons 

and bays have caused major changes in species 

composition and abundance. The 3 components of 

sewage effluent most detrimental to coral 

communities are nutrients, sediments, and toxic 

substances. Nutrient enrichment by sewage effluent 

may enhance benthic algal biomass and primary 

production in the water column. Increased primary 

production in the water column favors benthic 

filter-feeding invertebrates which, with the benthic 

algae, may out-compete corals and other reefbuilding organisms. Anthropogenic inputs of 

dissolved nutrients and organic particulate matter 

may also depress oxygen levels. While heavy 

sediment loads on corals may be lethal, lesser 

quantities may inhibit growth, cause changes in the 

growth forms of colonies, decrease coral cover, alter 

species composition of reef-building organisms, 

and inhibit coral recruitment. Toxic substances may 

induce metabolic changes in corals, decrease rates of 

growth and reproduction, or reduce viability of 

corals. Although further research is needed on all 

three major components of sewage effluent, the 

most critical need is for comprehensive, long-term 

studies of sewage impacts. The combined effects of 

particulate and toxic substances are especially 

poorly documented at this time. 

Pearson, T. H. (1975) The benthic ecology 

of Loch Linnhe and Loch Eil, a sea-loch 

system on the west coast of Scotland. 

IV. Changes in the benthic fauna 

attributable to organic enrichment. 

Journal of Experimental Marine Biology and 

Ecology 20: 1-4 1. 

{sediments; benthic fauna; species; anoxic; 

effluent) 

Changes are described in the benthic fauna of 

Lochs Linnhe and Eil in response to organic 

enrichment of the loch sediments, brought about by 

the input of effluent material from a pulp and paper 

mill. Fluctuations in the amount of effluent 

discharges are related to subsequent successional 

changes in the fauna. As organic enrichment 

proceeds, there is progressive elimination of 

species, accompanied by an increase in the numbers 

of the surviving species. Excessive organic 

deposition leading to anoxic conditions at the 

sedimentlwater interface results in the elimination 

of all but a few annelid species. The observed 

successional changes are discussed in the light of 

recent hypotheses concerning ecological changes in 

relation to environmental stress. 

Pearson, T. H. (1980) Marine pollution 

effects of pulp and paper industry wastes. 

Helgolander Meerestuntersuchungen 33:340-365. 

{pulp; paper; oxygen demand; benthic; abundance; 

sediment; composition) 

The scale of waste discharges to the marine 

environment from the pulp and paper industry in 

various parts of the world is outlined and a brief 

description of the major characteristics of such 

wastes is given. The information available on the 

direct toxicity of these wastes to marine fauna and 

flora is assessed including both lethal and sublethal 

effects. The environmental impact of waste 

discharge and subsequent ecosystem modifications 

are considered in detail; the complex adjustments to 

the nutrient and carbon budgets of the water column 

and sediment involved in direct alterations to their 

oxygen balance and other indirect eutrophication 

effects are described. It is concluded that whereas the 

direct toxicity of these wastes has minimal effect in 

the marine environment the complex problems 

created by increasing the oxygen demand on the 

receiving waters can have considerable impact in 

inshore areas. Methods of minimizing the effect of 

such impacts and of forecasting their extent are 

outlined. 

Pearson, T. H. (1987) Benthic ecology in an 

accumulating sludge-disposal site. In: J. 

M. Capuzzo and D. R. Kester (ed.) Oceanic 

Processes in Marine Pollution. Vol. I. Biological 

Processes and Wastes in the Ocean. Robert E. 

Krieger Publishing Company, Malaba, Florida. 

{benthic; sludge; Eh; sediments; pH; abundance; 

biomass; species; annelids; nematods; depositfeeder) 

The distribution of benthic fauna in the area of the 

sludge-disposal site in the Firth of Clyde, Scotland, 

is related to the gradient of increasing organic 
enrichment created by dumping. Faunal succession 

in relation to enrichment is related to changing 

productivity along the gradient. Community 

changes are ascribed to progressive maximization of 

carbon utilization as input levels increase; this 

involves responses at many levels of organization. 

It is concluded that the benthic ecosystem has fully 

adapted to the continuously high organic carbon 

input. 

Pearson, T. H., A. B. Josefson, and R. Rosenberg 

(1985) Petersen's benthic stations 

revisited. I. Is the Kattegat becoming 

eutrophic? Journal of Experimental Marine 

Biology and Ecology 92: 157-206. 

{benthic; macrofauna; abundance; biomass; 

increase; decrease; composition; Baltic) 

The macrobenthic fauna present at the stations 

sampled by C.G.J. Petersen in 191 1-1912 in the 

Kattegat were re-assessed in 1984 using equipment 

and techniques similar to those used in the original 

surveys. The total biomass of organisms present in 

1984 was found to be significantly reduced in the 

north and west of the area and in areas shallower 

than 25 m. These changes were largely attributable 

to decreases in the populations of large echinoid, 

Echinocardium. When echinoids were excluded, 

significant increases in the biomass of other 

organisms present in 1984 were found to the east of 

the area and in areas deeper than 25 m. In general 

the majority of species were found to be smaller in 

individual size in 1984 when compared with the 

earlier surveys. Dominance comparisons based on 

biomass changes showed that deposit-feeders had 

decreased and suspension-feeders and carnivores had 

increased in dominance. Most notably 

Echinocardium had decreased and the ophiuroid 

Amphiura filiformis (Miiller) had increased in 

dominance at over 70% of the stations sampled. In 

general ophiuroids and annelids had increased and 

echinoids and molluscs had decreased in dominance. 

Comparisons of similarity values indicated that 

community composition has changed markedly 

over the area, with only =30% similarity of species 

between 191 1-1912 and 1984 at most stations. 

The possible causes underlying these general 

changes are considered, amongst which changes in 

fish predation pressures, the direct effects of 

trawling and long-term temperature fluctuations are 

suggested. Whilst such factors may be partially 

responsible, the most probable causes is considered 

to be a general eutrophication of the area. 

Pearson, T. H. and R. Rosenberg (1976) A 

comparative study of the effects on the 

marine environment of wastes from 

cellulose industries in Scotland and 

Sweden. Ambio 5:77-79. 

(benthos; abundance; diversity; indicator) 

The influence of effluents from cellulose 

industries on the benthic faunas of comparable 

fjordic systems on the west coasts of Sweden and 

Scotland is assessed. Generalizations are made 

about sequential changes in the interrelationship of 

benthos and sedimentary structure. Estimates are 

made of the rates of such changes along a gradient 

of environmetal stress brought about by the organic 

enrichment of marine sediments and these can be 

used to assess the probable effect of large inputs of 

organic material to marine sediment. The 

advantages and disadvantages of using indicators 

species, comparing faunal groups, and using 

diversity indices in the description of ecological 

changes are briefly considered. 

Pearson, T. H. and R. Rosenberg (1978) 

Macrobenthic succession in relation to 

organic enrichment and pollution of the 

marine environment. Oceanography and 

Marine Biology: Annual Review 16:229-311. 

{macrobenthos; benthic; organic enrichment; 

sediments; abundance; biomass; spatial; temporal; 

diversity; similarity; indicator; opportunistic) 

No anstract 

Pearson, T. H. and S. 0. Stanley (1979) 

Comparative measurement of the redox 

potential of marine sediments as a rapid 

means of assessing the effects of organic 

pollution. Marine Biology 53:371-379. 

{sediments; redox potential; macrofauna; benthos; 

abundance; species; biomass) 

A rapid method for measuring the redox potential 

(Eh) values of marine sediments is described. This 

method has been used to relate changes in the areas 

of highly reduced sediment in a sea loch system to 

the continuously fluctuation inputs of organic 

waste from a pulp and paper mill. Subsequent 

faunal changes in these areas can be related to 

changes in the measured sedimentary redox level. 

The method is suggested as a rapid means of 

assessing the potential impact of an additional 

organic input to a marine sediment. 

Pease, B. C. (1977) The effect of organic 

enrichment from a salmon mariculture 

facility on the water quality and benthic 

community of Henderson Inlet, 

Washington. Dissertation for Ph.D., University 

of Washington. 

{mariculture; benthos; infauna; epifauna; 

sediments) 

The effect of fish excretion and respiration at a 

salmon mariculture facility on the local water 

quality was evaluated. In the immediate vincinitiy 

of the facility, dissolved oxygen concentrations 

were consistently depressed while ammonia 

concentrations were increased, but neither effect was 

significant based on accepted criteria. The 
mariculture facility had no apparent effect on the 

local phytoplankton population. Major fluctuations 

in the local water quality were a result of natural 

hydrographic conditions and phytoplankton 

activity. 

The effect of fish feces and excess fish food from 

the mariculture facility on the benthic environment 

was also evaluated. There was a definite 

accumulation of organic matter under and extending 

at least 15 m from the salmon pens. This organic 

accumulation caused significant changes in the 

structure of the benthic epifaunal and infaunal 

communities. Epibenthic mats of the sulfideoxidizing bacterium, Beggiatoa, were observed 

under salmon pens along with an increased 

abundance of several large predatory species ( 

Cancer gracilis, Lepidopsetta bilineata, and 

Myoxocephalus polyacanthocephalus). The benthic 

infauna community under the salmon pens declined 

in species richness and became numerically 

dominated by the polychaete worm, Capitella 

capitata, a well-known ubiquitous indicator of 

organic pollution. Therefore, the mariculture 

facility would have little or no effect on the marine 

environment if the settleable solids were collected 

and removed, or depressed in such a way that there 

was no accumulation in the underlying sediments. 

Pekkari, S. (1973) Effects of sewage water 

on benthic vegetation. 0 I K 0 S 

Supplymentum 15: 185-1 88. 

{seaweeds; sewage; benthic; abundance; abnormal) 

The influence of sewage water on benthic 

vegetation was studied in the Stockholm 

Archipelago and in the Gulf of Bothnia. The most 

conspicuous effects of sewage effluents in the Gulf 

of Bothnia are the excalves of marine and freshwater 

vegetation distinctly differing from the vegetation 

in the vi cinity. Disappearance and abnormal shape 

of plants are conspicuous effects of sewage water in 

the inner part of the Stockholm Archipelago. 

Perciasepe, R. (1992) Progress Report of the 

Baywide Nutrient Reduction 

Reevaluation. Chesapeake Bay Program, 

{abundance; oxygen) 

In 1987, the parties to the original Chesapeake 

Bay Agreement signed a new Chesapeake Bay 

agreement. The 1987 Chesapeake Bay Agreement 

set a specific goal-to achieve at least a 40 percent 

reduction of nitrogen and phosphorus entering the 

mainstem Chesapeake Bay by the year 2000. 

Although this progress report's fingdings are 

preliminary, trends and generalizations of nutrient 

loads, water quality, and hibatat improvements are 

becoming evident. Most of the background studies 

for the Nutrient Reduction Reevaluation have been 

drafted. To date, seven model runs have been 

completed for use in this report. 

Perry, D. M., J. B. Hughes, and A. T. Herbert 

(1991) Sublethal abnormalities in 

embryos of winter flounder, 

Pseudopleuronectes americanus, from 

Long Island Sound. Estuaries 14:306-317. 

{abnormalities; winter flounder; disease; Long 

Island Sound; Boston Harbor) 

Sublethal abnormalities were examined in 

developing embryos of the winter flounder, 

Pseudopleuronectes arnericanus, as part of a study 

of its early reproductive success in Long Island 

Sound and two sites in Boston Harbor. These sites 

represented varying levels of anthropogenic 

contamination that were possibly affecting 

reproduction, as ascertained by chromosomal 

abnormalities and lowered developmental rates in 

the embryos. Effort was focused on examination of 

blastula and tail-bud embryos from field-captured 

females spawned at the laboratory. Abnormalities 

observed included evidence of cytotoxicity and 

chromosome damage. Embryos of fish from New 

Haven were usually the most aberrant, while 

embryos from other sites, notably Hempstead, 

Shoreham, and both Boston Harb or stations, 

showed subtle indications of abnormality. 

Persson, L.-E. (1987) Baltic eutrophication: 

A contribution to the discussion. Ophelia 

27:3 1-42. 

{Baltic; macrozoobenthos; abundance; diversity) 

Quantitative sampling of macrozoobenthos has 

been performed in the southern Baltic in the 1920s 

and the 1950s. Some of these stations (43) were 

revisited in 1982183. The total abundance and 

biomass was significantly increased in 1982183 

compared to the 1950s. It is suggested that the 

main part of the biomass increase might be due to 

reduced stocks of predatory flatfishes in the 1920s 

and 1930s. 

The most conspicuous difference in the 

comparisons is the widespread distribution of the 

polychaete Capitella capitata in 1982183. This is 

attributed to increased trawling activities in recent 

decades and not to organic enrichment. 

Phillips, D. J. H. and S. Tanabe (1989) Aquatic 

pollution in the Far East. Marine Pollution 

Bulletin 20:297-303. 

{Far East; algal blooms; oxygen; red tides) 

An overview of the problems experienced in the 

Far East relating to aquatic pollution is presented, 

with emphasis on the quality of coastal waters. The 

pollution problems faced by developing and 

developed nations in the region tend to vary in type 

according to socioeconomic factors. The appropriate 

treatment and disposal of sewage remains a 

particular problem in the developing nations. 

Industrial and agricultural effluents also exert 

measurable effects on marine resources in many of 

these regions. The environmental impacts of major 
engineering and development projects are also a 

cause for concern in certain areas. In the developed 

nations, the impacts of conservative contaminants 

(especially the highly persistent and toxic 

organochlorines) are considered to be the most 

significant problem at present. The region as a 

whole exhibited a great dependence on seafood as a 

primary or preferred source of protein, and the 

protection of coastal waters is.therefore of vital 

concern. In addition, tourism provides much-needed 

foreign exchange in many countries of the region, 

and the tourist facilities are mostly situated on the 

coast. Additional regulatory initiatives are required 

in many areas, coupled to strong legislative action 

and robust monitoring of coastal pollution, if the 

vital marine resources are not to be irreversibly 

damaged. 

Phoel, W. C., K. L. Webb, and C. F. D'Elia 

( 19 8 1) Inorganic nitrogen regeneration 

and total oxygen consumption by the 

sediments at the mouth of the York 

River, Virginia. In: B. 3. Neilson and L. E. 

Cronin (ed.) Estuaries and Nutrients. Humana 

Press, Clifton, New Jersey. 

{nitrogen; regeneration; oxygen} 

In situ measurements of inorganic nitrogen fluxes 

and riverbed oxygen consumption were made on 

sediments in 3, 9, and 16 m of water at the mouth 

of the York River during stratified and destratified 

water conditions. Ammonium was regenerated, the 

rate of which increased with depth and oxygen 

concentration in the overlying water. Nitrate and 

nitrite fluxes from the sediment were minimal or 

non-existent during stratification at the 16-m 

station but increased and the nutrients were taken 

up by the sediments under destratified conditions. 

At the 3-m station, which is above the halocline 

when developed, nitrate and nitrite appeared to be 

the major forms of nitrogen being released by the 

sediments. Oxygen consumption by the riverbed at 

the 16 and 9-m stations was higher during the 

increased oxygen tensions associated with vertical 

destratification. The 3-m stations maintained the 

higher rates of oxygen consumption throughout the 

sampling period. The in situ incubation of bottom 

water alone at all three stations indicated negligible 

rates of oxygen uptake. 

Polak, J. and G. D. Haffner (1978) Oxygen 

depletion of Hamilton Harbour. Water 

Research 12:205-215. 

{oxygen; eutrophic; demand; sediment} 

Processes involved in the oxygen cycles of the 

highly eutrophic Hamilton Harbour were studied. 

Sediment oxygen demand ks(02 rn-2 day-l) was 

measured by an in situ method and was determined 

to be dependent on oxygen concentration c (ml-l) 

in the water phase. This dependence was expressed 

by the equation k, = 0.72 + 0.26~ . The water 

column oxygen demand of the harbour was 

determined experimentally and revealed a significant 

dependence on oxygen concentration. It was 

estimated that over 80% of the oxygen supplied to 

the harbour was used within the water column. The 

sediment oxygen consumed about 18% of the 

oxygen entering the harbour and was relatively 

most important in the early stages of stratification 

when the hypolimnetic dissolved oxygen 

concentrations were high. The main sources of 

oxygen were atmospheric reaeration (80%), lakeharbour exchange (10%) and photosynthesis (10%). 

Portnoy, J. W. (1991) Summer oxygen 

depletion in a diked New England 

estuary. Estuaries 14: 122-129. 

{oxygen; estuary; organic; fish kill; marsh} 

The diked and freshened Herring River estuary 

(Wellfleet, Massachusetts) experiences regular 

summer hypoxia and one- to three-week period of 

main stream anoxia, often accompanied by fish 

kills. Stream hypoxia results from the temperaturedependent increase in oxygen demand of organic 

matter released by diked salt marsh deposits; periods 

of total anoxia are induced by heavy rains which 

increase the runoff of wetland organic matter. 

Historic reductions in tidal flushing have extended 

the low salinity region of the estuary normally 

characterized by high organic loads and minimal 

flushing. Recurrent main stream anoxia has 

depressed both migratory and resident aquatic fauna. 

Porumb, F. (1992) On the development of 

Noctiluca scintillans under 

eutrophication of Romanian Black Sea 

waters. In: R. A. Vollenweider, R. Marchetti 

and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{species, abundance, noctiluca scintillans, Black 

Sea} 

This paper analyses the development of Noctiluca 

scintillans during the 1980s and 1990s as a 

consequence of the intense eutrophication of the 

Romanian waters of the Black Sea. 

Price, K. S., D. A. Flemer, J. L. Taft, G. B. 

Mackiernan, W. Nehlsen, R. B. Biggs, N. H. 

Burger, and D. A. Blaylock (1985) Nutrient 

enrichment of Chesapeake Bay and its 

impact on the habitat of striped bass: A 

speculative hypothesis. Transactions of the 

American Fisheries Society 114:97-106. 

{Chesapeake Bay; fish; striped bass; oxygen; 

habitat; nitrogen; phosphorus; aquatic vegetation; 

chlorophyll) 

Stocks of striped bass Morone saxatilis have 

declined in the Chesapeake Bay system over the last 

decade. We present evidence for the working 

hypothesis that the decline has resulted, in part, 
from loss of deep-water habitat for adults, caused by 

limiting concentrations of dissolved oxygen that are 

related, in turn, to nutrient enrichment and greater 

planktonic production. A related hypothesis is that 

changes in the near-shore habitat for juvenile 

striped bass, involving severe declined in 

submerged aquatic vegetation due to nutrient-driven 

planktonic shading, also have contributed to the 

decline of striped bass. Nutrients (nitrogen and 

phosphorus) and in chlorophyll a, an indicator of 

phytoplankton biomass, have increased i many 

areas of the bay and tributaries over the past 20 to 

30 years. These trends are qualitatively correlated 

with greater deoxygenation of the deep channel in 

the mid and upper bay. During the late 1970s, 

summer oxygen concentrations as low as 2 mlniter 

approached to within 7-8 m of the surface, allowing 

water stressful to striped bass to intrude onto shoal 

areas of the bay. The volume of Chesapeake Bay 

bottom waters containing 0.5 mg 02/liter or less 

was about 15 times greater in July 1980 than in 

July 1950. The combination of the expanding 

hypoxic pool and summer temperature above 

preferred levels for adult striped bass may contribute 

to and "oxygen-temperature squeeze" that forces 

adults onto shoal areas of the bay or out of the 

upper bay. Many of these shoal areas now lack 

suitable cover for juvenile striped bass and their 

prey. Strong intraspecific competition among 

striped bass may be occurring there. 

Provini, A., G. Crosa, and R. Marchetti (1992) 

Nutrient export from the Po and Adige 

river basins over the last 20 years. In: 

R. A. Vollenweider, R. Marchetti and R. Viviani 

(ed.) Marine Coastal Eutrophication. Elsevier, 

Amsterdam. 

{phosphorus, nitrogen, Adriatic Sea) 

Available data on nutrient concentration and flow 

of the Po and Adige river waters were analyzed in 

order to evaluate the nitrogen and phosphorus load 

carried into the Adriatic Sea, as well as the changes 

that have occurred since 1968. Load data results 

were highly correlated with the flow, following 

relations of the linear type. The yearly mean loads 

amount to 13,000 t of total P and 110,000 t of 

total mineral N for the Po, and 1200 t and 13,000 t 

respectively for the Adige. These values are more 

than two times higher than those evaluated in 

1968. Nitrogen increased quite regularly, while the 

phosphorus load had already reached its maximum 

value at the end of the 1970s. Nutrient increase 

during 1968-88 can be explained by a higher 

production of detergents and fertilizers, and the 

conshctions of numerous sewer networks in the 

Po and Adige basins. 

Quinlan, A. V. (1982) An Ecodynamic 

Analysis of Algal Blooms Fouling 

Nahant Bay Beaches. MIT Sea Grant 

Program, MITSG 82-15. 

{seaweeds) 

Since the early 1900's the sandy beaches of 

Nahant Bay, Massachusetts have been chronically 

fouled by hundreds of tonnes of an abnormal form 

of the filamentous brown alga Pilayella littoralis. 

In warm weather, foul-smelling gaseous 

decomposition products sporadically cause great 

discomfort in the local residential and recreational 

populations. At the request of state and federal 

officials, a multidisciplinary research team was 

assembled to find out why this alga grows so 

abundantly in the surf zone and shallow sublittoral 

of the fouled beaches. The research team was 

specifically asked to evaluate the influence of 

effluents from a municipal sewage treatment plant 

outfall located on the bay's ocean boundary. The 

concentrations of algal nutrients were measured in 

the outfall boil and at foyrteen other locations 

throughout the bay at seventeen monthly intervals. 

In addition, dyes, drift-cards, and computer 

simulations were used to estimate the trajectory of 

effluents released from the treatment outfall. All 

the available information consistently indicated that 

the effluent plume is completly dispersed long 

before reaching the algal drift community, and at 

most, only 1-2% of the macronutrients available to 

the alga come from the outfall. Tidal exchange 

with Massachusetts Bay was found to contribute 

roughly 74 times as much nitrogen, the 

macronutrient apparently limiting the alga's 

growth, as the outfall does on each tide. As a 

consequence, the municipal sewage effluents 

released into Nahant Bay were exonerated as a cause 

of the algal fouling problem there. 

Quinn, H., J. P. Tolson, C. J. Klein, S. P. 

Orlando, and C. Alexander (1989) 

Susceptibility and Status of Gulf of 

Mexico Estuaries to Nutrient Discharges. 

NOAA National Ocean Service, {nutrients; 

nitrogen; phosphorus; estuary; susceptibility; N/P 

ratio; classification; review) 

No abstract 

Quinn, H., J. P. Tolson, C. J. Klein, S. P. 

Orlando, and C. Alexander (1989) 

Susceptibility of East Coast Estuaries to 

Nutrient Discharges: Albemarie/Pamlico 

Sound to Biscayne Bay. NOAA National 

Ocean Service, (nutrients; nitrogen; phosphorus; 

estuary; susceptibility; NIP ratio; classification; 

review) 

No abstract 

Quinn, H., J. P. Tolson, C. J. Klein, S. P. 

Orlando, and C. Alexander (1989) 

Susceptibility of East Coast Estuaries to 

Nutrient Discharges: Passamaquoddy Bay 
to Chesapeake Bay. NOAA National Ocean 

Service, {nutrients; nitrogen; phosphorus; estuary; 

susceptibility; N/P ratio; classification; review) 

No abstract 

Quinn, H., J. P. Tolson, C. J. Klein, S. P. 

Orlando, and C. Alexander (1991) 

Susceptibility of West Coast Estuaries to 

Nutrient Discharges: San Diego Bay to 

Puget Sound. NOAA National Ocean Service, 

{nutrients; nitrogen; phosphorus; estuary; 

susceptibility; NP ratio; classification; review) 

No abstract 

Rabalais, N. N. and D. F. Boesch (1986-1987) 

Extensive depletion of oxygen in bottom 

waters of the Louisiana shelf during 

1985. Coastal Ocean Pollution Assessment News 

3:45-47. 

{oxygen; Louisiana; Gulf of Mexico) 

No abstract 

Rabalais, N. N., R. E. Turner, J. W.J. Wiseman, 

and D. F. Boesch (1991) A brief summary of 

hypoxia on the northern Gulf of Mexico 

continental shelf: 1985-1988. In: R. V. 

Tyson and T. H. Pearson (ed.) Modern and Ancient 

Continental ShelfAnoxia. The Geological Society, 

London, England. 

{oxygen; hypoxia; Gulf of Mexico; Louisiana; 

phytoplankton) 

Oxygen-deficient conditions occur from April to 

October on the inner to middle continental shelf of 

the northern Gulf of Mexico and may cover up to 

9500 km2 during mid-summer off the Louisiana 

coast. Hypoxic bottom waters are found in 5-60 m 

water depth, 5-60 km offshore and extend up to 20 

m above the bottom. Salient forcing functions 

contributing to hypoxic water formation, 

maintenance and break-up are presented. 

Stratification is directly correlated with hypoxia in 

time and space, suggesting that reaeration of 

bottom waters is controlled by physical processes 

that are influenced by regional wind fields, river 

discharge and continental shelf scale currents. 

~h~to~lankton biomass reaches the bottom waters 

in the hypoxic zones in large amounts (>5 ugtl), 

fueling water column and benthic respiration rates, 

but to a currently unknown degree. Increased 

nutrient loadings in the two major rivers and 

changes in the proportion of those nutrient 

essential to phytoplankton growth have probably 

changed both the phytoplankton community 

species composition and community production. It 

is not yet clear whether the extent and severity of 

hypoxia on the Louisiana continental shelf have 

changed as a result of these riverine water quality 

changes. 

Radziejewska, T. and I. Drzycimski (1988) 

Meiobenthic communities of the Szczecin 

Lagoon. Kieler Meeresforschungen, Sonderheji 

6:162-172. 

{composition; distribution; meiobenthos; lagoon; 

Baltic; nematode; ostracod; copepods) 

Based on data collected at 7 stations 4 times a year 

in 1985 and 1986, composition and distribution of 

meiobenthic communities in the Polish part of the 

Szczecin Lagoon, a eutrophic and polluted water 

body connected with the Baltic Sea, is presented. 

The data show a tendency to reduced total 

meiobenthic densities and diversity from the lower 

to upper reaches of the lagoon. The meiobenthic 

communities studied were dominated by nematodes, 

ostracods ranking second in numerical importance. 

Harpacticoid copepods were most abundant at the 

outer stations which are influenced by Baltic 

inflows. Most of the 10 harpacticoid- species 

recorded in the lagoon were found at the outer 

stations (lower reached) as well. Similarity analysis 

allowed to separate three zones within the lagoon: 

(1) the outer zone, its stations showing most 

abundant and diverse meiobenthic communities; (2) 

the innermost zone (upper reaches) with the least 

abundant and qualitatively impoverished 

communities; and (3) the intermediate zone. 

Raffaelli, D., J. Limia, S. Hull, and S. Pont 

(1991) Interactions between the amphipod 

Corophium volutator and macroalgal 

mats on estuarine mudflats. Journal of the 

Marine Biological Association of the United 

Kingdom 71:899-908. 

{macroalgae; seaweeds; mudflat; amphipod; 

Corophium; Capitella) 

One of the most obvious effects of eutrophication 

in sheltered coastal areas and estuaries is enhanced 

growth of opportunistic macroalgae, which may 

form extensive mats over intertidal mudflats during 

the spring and summer. In the Ythan estuary, 

densities of the amphipod Corophium volutator 

(Pallas) in the sediment underlying weed mats were 

significantly lower than those in weed-free 

sediments, and are dominated by species 

characteristic of organically enriched, low oxygen 

environments such as Capitella capitata. Long-term 

data sets on Corophuim abundance in the Ythan 

suggest that this species has declined dramatically 

throughout those parts of the estuary affected by 

weed mats. 

Two fields experiments have been carried out to 

assess the effects of weed mats on assemblages. In 

the first experiment, sediments subjected to a range 

of weed densities displayed faunal responses 

consistent with those observed by survey of weedcovered and weed-free areas. Under a high biomass 

of weed, the amphipod Corophuim volutator 

disappeared almost completely from the mudflat. 

The second experiment indicated that the physical 
presence of weed filaments may be as important as 

the effects of weed on sediment chemistry in 

affecting the density of Corophium, probably 

through interference with the amphipod's normal 

feeding behavior. It is concluded that weed mats 

have a significant impact on Corophium, which is 

an important component of the diet of fish and 

shore-birds in the estuary. 

Raffaelli, D. G. and C. F. Mason (1981) 

Pollution monitoring with meiofauna, 

using the ratio of nematodes to copepods. 

Marine Pollution Bulletin 12: 158- 163. 

{meiofauna; nematode; copepod; ratio; sandy 

beaches) 

The ratio of nematodes to copepods was calculated 

for meiofauna samples from 17 beaches and 

additional values for intertidal and sublittoral sites 

were obtained from the literature. Intertidal ratios 

increased with decreasing particle size, but ratios 

from polluted sites were always extremely high. 

Sublittoral ratios increased with depth. It is 

suggested that this ratio has potential for 

monitoring organic pollution of sandy beaches. 

Raman, A: V. and K. P. Prakash (1989) 

Phytoplankton in relation to pollution in 

Visakhapatnam harbour, east coast of 

India. Indian Journal of Marine Sciences 18:33- 

36. 

{phytoplankton; nutrients; Visakhapatnarn harbour; 

sewage; abundance; diversity} 

Sporadic outbursts of phytoplankton, notably 

diatoms and phytoflagellates, are observed from 

Visakhapatnam harbour waters subjected to 

pollution. The harbour waters are characterized by 

high proportions of inorganic nutrients attributable 

to sewage and industrial waste discharges. High 

phytoplankton standing crop and chlorophyll 

concentrations accompanied by low species 

diversity at the harbour locations indicate severe 

eutrophidation of waters in this area as against near 

normal conditions in the open sea. 

Reguera, B. and Y. Oshima (1990) Response 

of Gymnodinium catenatum to increasing 

levels of nitrate: Growth patterns and 

toxicity. In: E. Graneli, B. Sundstrom, L. 

Edler and D. M. Anderson (ed.) Toxic Marine 

Phytoplankton. Elsevier, New York. 

{Gymnodinium; nitrate; phytoplankton; growth; 

red tides) 

Gymnodinium catenatum, responsible for PSP 

outbreaks in NW Spain, was cultured in K media 

with increasing levels of nitrate. Growth rates were 

calculated from measurements of in vivo 

fluorescence and cell counts. Toxicity was 

determined by HPLC. Toxin concentrations per cell 

reached a peak with nitrate concentrations of 110- 

220 uM. Much of the increase in toxin 

concentration was due to a higher production rate of 

GXT 6, showing that the toxin profile is not a 

conservative property in this species. 

Renk, H., J. Nakonieczny, and S. Ochocki (1988) 

Primary production in the southern Baltic 

in 1985 and 1986 compared with longterm mean seasonal variation. Kie 1 e r 

Meeresforschungen, Sonderheft 6:203-209. 

{primary production; Baltic; phytoplankton; 

Chlorophyll a) 

On the basis of long-term investigations, the 

course of seasonal variations of mean daily primary 

production and mean chlorophyll-a concentration in 

the Southern Baltic waters are presented. The mean 

annual primary production of the Gda'nsk Deep, 

Borholm Deep and the Gotland Deep amounted to 

12.4 gC m-2, 88.8 gC m-2 and 107.2 gC m-2, 

respectively. The primary production of the 

Southern Baltic in 1986 was higher than in 1985, 

and higher than the long-term mean value. The 

mean annual production for the Southern BalEic in 

1986 amounted to 130 gC m-2. On the basis of 

long-term observations of primary production and 

chlorophyll-a of the Southern Baltic, certain upward 

trends in phytoplankton production were observed. 

Revelante, N. and M. Gilmartin (1976) The 

effect of Po River discharge on 

phytoplankton dynamics in the Northern 

Adriatic Sea. Marine Biology 34:259-271. 

{phytoplankton; Adriatic Sea; community; 

nutrients; density; primary production; 

nanoplankton; microplankton; bloom) 

An oceanographic transect, extending from 

Yugoslavia across the Northern Adriatic Sea to the 

Po delta in Italy, was occupied during 1972 and 

1973 to establish the effect of Po river discharge on 

the phytoplankton communities of the region. 

Density distribution showed distinct seasonal 

features: a winter-spring period of low stability 

throughout the water column, and a summer period 

of stratification. The total water-column plant 

nutrients (nitrate, nitrite, phosphate, silicate) 

showed a 1.3 to 4.5-fold decrease eastward, with 

semi-eutrophic conditions restricted to an area off 

the Po delta. Nanoplankton usually dominated the 

phytoplankton community, in terms of cell 

density, surface chlorophyll a concentrations, and 

surface primary production rates. However, all 

significant maxima in these characteristics resulted 

from increases in the microplankton component. 

Changes in the frequency of major microplankton 

groups characterized three periods of the annual 

cycle: September-December, neritic, temperate 

diatom flora with some littoral elements (e.g. 

Nitschia seriata); January-May, neritic, temperate 

diatom flora of different composition (e.g. Lauderia 

borealis, Skeletonema costatum); May-August, 

dinoflagellates (e.g. Prorocentrum micans) at 
western stations and dinoflagellates plus neritic, 

warm-water diatoms at eastern stations. The 

seasonal cycle was characterized by spring and fall 

maxima tending to coincide with maximum Po 

river discharge andlor periods of low water-column 

stability and vertical mixing. The higher nutrient 

input at western stations was correlated with the codominance of only a few species of microplankton 

during bloom periods, suggesting that these species 

(S. costatum, N. seriata, and 5 others) can serve as 

indicators of eutrophic conditions in this region. 

Assimilation ratios of both the micro- and 

nanoplankton suggested borderline nutrient 

conditions. Phosphate was implicated as the 

limiting nutrient. 

Revelante, N. and M. Gilmartin (1977) The 

effects of Northern Italian rivers and 

Eastern Mediterranean ingressions on the 

phytoplankton of the Adriatic Sea. 

Hydrobiologia 56:229-240. 

{phytoplankton; Mediterranean; Adriatic Sea; 

standing crop; diversity; chl. a; density; 

microplankton; nanoplankton; Nitzschia seriata; 

indicator; dinoflagellate; coccolithophores) 

An analysis of the distribution of the 

phytoplankton standing crop during a period of 

high stability characterized by the reIative influence 

of northern Italian rivers and Mediterranean waters 

on the Adriatic Sea. The region influenced by 

northern Italian rivers was marked by low 

community diversities, relatively high chlorophyll 

a standing crops and cell densities, high 

microplankton to nanoplankton community 

volume ratios, and the dominance of Nitzschia 

seriata. To the contrary, the region influenced by 

Mediterranean waters exhibited high community 

diversities with prominent oceanic and warm water 

elements, low chlorophyll a standing crops and 

microplankton cell densities, low microplankton to 

nanoplankton ratios, and a relatively conspicuous 

coccolithophore flora. 

Nanoplankton populations remained relatively 

constant throughout the Adriatic. However 

eutrophication resulted in a marked increase in the 

microplankton component of the community, with 

the diatom Nitzschia seriata serving as a prime 

indicator of the process, even in regions several 

hundred kilometers downstream from nutrient 

sources. 

Even under relatively low rates of river discharge, 

the influence of northern Italian rivers could be 

traced along the entire western side of the sea to the 

Otranto Strait, with low north to south and high 

west to east gradients resulting from the effect of a 

cyclonic surface circulation pattern on inflowing 

Mediterranean waters in the south and river 

discharge in the north. 

Revelante, N. and M. Gilmartin (1980) 

Microplankton diversity indices as 

indicators of eutrophication in the 

Northern Adriatic Sea. Hydrobiologia 

70:277-286. 

{phytoplankton; diversity; Adriatic Sea; 

Mediterranean; indicator; chl. a) 

Phytoplankton community diversity indices are 

used to characterize effects of eutrophication in the 

Northern Adriatic Sea. A derived Shannon diversity 

frequency spectrum provided a single biological 

quantification which allowed an interpretation of 

temporal and regional differences and which can 

also be used to evaluate future changes in species 

diversity. The data base comprised a 4+ year time 

series involving 300 taxa. 

Reys, E. and M. Merino (1991) Die1 dissolved 

oxygen dynamics and eutrophication in a 

shallow, well-mixed tropical lagoon. 

Estuaries 14:372-38 1. 

{oxygen; dynamics; eutrophication; lagoon; 

Mexico; primary productivity) 

Bojorquez Lagoon (BL), located on the Mexican 

Caribbean, has received sewage and dredging 

impacts as a result of tourism development. The 

lagoon supports a high diversity of primary 

producers compared to sheltered adjacent lagoons 

dominated by Thalassia testudinum communities. 

The Diurnal Curve Method (Odum and Hoskin, 

1958) was used to measure community metabolism 

and asses eutrophication in BL by comparing it to 

the nonimpacted lagoons and to other systems 

studied with this method. Dissolved oxygen 

community input to the water column in BL ranged 

between 8.3 g 02 m-2 d-l and 41.5 g 02 m-2 d-l 

during 1985 and 1986, and averaged 17.1, whereas 

dissolved oxygen community consumption ranged 

from 6.4 g 02 m-2 d-l to 37.6 g 02 m-2 d- l and 

averaged 15.2. These values are higher than those 

found for the adjacent lagoons and similar coastal 

lagoons, and are similar to results from other 

lagoons with sewage or seafood waste discharge. 

Net flux of oxygen from the community to the 

water column averaged 1.9 g 02 m-2 d-l and ranged 

from -9.8 g 02 m-2 d-l to 8.1 g 02 m-2 d-l. 

These values are low compared to the adjacent 

lagoons, and close to zero, as in dystrophic 

environments. Primary productivity, as estimated 

by oxygen input, increased in BL during the period 

of study, indicating that eutrophication is 

proceeding, but the lagoon has not reached yet a 

level of "critical eutrophication" as defined by Mee 

(1988). 

Riegman, R., A. Rowe, A. A. M. Noordeloos, and 

G. C. CadCe (1993) Evidence for 

eutrophication induced Phaeocystis sp. 

blooms in the Marsdiep area 
< ether lands). In: T. J. Smayda and Y. 

Shimizu (ed.) Toxic Phytoplankton Blooms in the 

Sea. Elsevier, Amsterdam. 

{Phaeocystis; ammoniurn/nitrate ratio} 

Recently, a global epidemic of novel and nuisance 

phytoplankton blooms has been suggested. 

Antluopogenic linkage is still questionable because 

the involved mechanisms are unclear. In all coastal 

areas of the European continent, anthropogenic 

enrichment of N and P (but not Si) has favoured 

non-diatom blooms during the nutrient controlled 

period (late spring-autumn). The colonial 

flagellate, Phaeocystis sp., regarded harmful1 

because of its massive foam production during the 

wane of the bloom, has increased in Dutch coastal 

waters. This increase coincides with a decrease of 

the average N/P ratio from 38 down to 13 dutring 

the nutrient controlled period (summer) as a 

consequence of altered nutrient discharges from 

Lake IJssel and the river Rhine. Competition 

experiments in continuous cultures showed that the 

Phaeocystis could become dominant at N/P ratios 

below 16. At higher N/P ratios Phaeocystis did 

not become dominant. In laboratory cultures, 

colony formation was absent under P-limitation. 

Under N-limitation, Phaeocystis formed only 

colonies when nitrate was the nitrogen source. 

Dutring growth on ammonium, no colonies were 

formed under limiting conditions. The importance 

of the ammoniumlnitrate ratio for Phaeocystis 

colony formation under nutrient limiting conditions 

was illustrated in 1991 by a summer bloom which 

occurred in the Marsdiep area after the 

ammoniumlnitrate ratio had dropped remarkably. 

Riisgird, H. U. and E. Poulsen (1981) Growth 

of Mytilus eudulis in net bags transferred 

to different localities in a eutrophicated 

Danish Fjord. Marine Pollution Bulletin 

12:272-276. 

{growth; bivalves; Mytilus; fjord; H2S; oxygen} 

The growth rate has been measured in Mytilus 

edulis transferred in net bags to seven localities in 

the brackish Danish fjord, Limfjorden, in which 

certain areas are heavily eutrophicated. The increase 

in shell length, shell weight and flesh body weight 

was measured after growth periods of 14-18 days. 

The increase in, e.g., flesh dry weight ranged from 

twice the start weight to a four-fold increase of the 

start weight. The net growth efficiencies were 

estimated to be between 54 and 73%. Algal 

concentration was in no case the limiting growth 

factor, but low growth rates were observed in areas 

with seasonal oxygen depletion and release of toxic 

H2S from the sediments. It is suggested that 

measurements of actual growth in Mytilus edulis 

can be a useful technique in the study of biological 

effects in marine recipients. 

Ritz, D. A., M. E. Lewis, and M. Shen (1989) 

Response to organic enrichment of 

infaunal macrobenthic communities under 

salmonid seacages. Marine Biology lO3:2 1 1 - 214. 

{macrobenthos; benthic; infauna; organic 

enrichment; abundance; biomass; diversity} 

The response of infaunal macrobenthic 

communities beneath salmonid seacages to solid 

organic wastes (food pellets and fish excreta) was 

tested by means of the ABC-method (abundance, 

biomass comparison methods: Warwick 1986) on 

samples collected from the SAFCOL seafarm at 

Badger Cove, southeast Tasmania, from February 

to July 1988. The intermittent nature of the 

addition of organic waste allowed us to monitor 

decline and recovery of the macrofauna, i.e., 

harvesting of a cage permitted a period of recovery 

whilst restockmg precipitated a decline. The ABCmethod proved to be a sensitive indicator of 

community health. Under a normal feeding regime 

the macrofaunal community structure indicated a 

moderately disturbed condition. Only 7 wk after the 

cage was harvested, species richness had increased 

markedly and the community adopted an 

undisturbed condition. Further improvement was 

apparent 14 wk post-harvest. Similarly, a decline to 

a moderately disturbed condition was apparent 7 wk 

after restocking and species richness had declined. 

No such changes occurred under a cage which 

contained fish continuously over the same period. 

Rosenberg, R. (1985) Eutrophication-the 

future marine coastal nuisance? Marine 

Pollution Bulletin 16:227-23 1. 

{oxygen; benthic fauna; fish; Sweden; bloom; 

review) 

Increased inputs of nutrients to marine coastal 

areas over the last decades have created a basis for 

eutrophication of the waters surrounding Sweden. 

In combination with relatively low water exchange 

in these vertically stratified and almost non-tidal 

waters, local and regional effects of increased 

macro-algal biomass, and decreased oxygen 

concentrations in bottom water leading to 

mortalities of benthic animals and decreased fish 

catches have at times been observed. The effects 

were first noted in the Baltic, but are now obvious 

also in Swedish and Danish coastal areas in the 

Kattegat and the Belt Sea. Similar symptoms have 

recently also been recorded off the Danish North 

Sea coast. Other shallow coastal and shelf areas, 

where stratification occurs, can be regarded as 

potentially eutrophic risk areas. 

Rosenberg, R. (1992) Eutrophication-related 

marine ecosystem studies in western 

Sweden. In: G. Colombo, I. Ferrari, V. U. 

Ceccherelli and R. Rossi (ed.) Marine 
Eutrophication and Population Dynamics. Olsen & 

Olsen, Fredensborg. 

{ecosystem; sediment transport; Kattegat) 

A six year multidisciplinary programme in the 

Kattegat on the Swedish west coast is summarized. 

There are indications that primary production and 

sedimentation of organic material have increased 

lately due to eutrophication. Although primary 

production is not exceptional (annual mean 144 g 

C per square meter) the special geomorphological 

and hydrographical featurew of the southern 

Kattegat (shallow, semi-enclosed, no tide, strong 

halocline) promote development of periodic 

bottom-oxygen deficiency. The ecological 

importance of vertical sedimentation and of lateral 

transport of ephemeral blankets into areas of 

deposition is di-scussed. Several examples of 

rapidly occurring negative ecological effects caused 

by eutrophication are given. 

Rosenberg, R., R. Elmgren, S. Fleischer, P. 

Jonsson, G. Persson, and H. Dahlin (1990) 

Marine eutrophication case studies in 

Sweden. Ambio 19: 102-108. 

{review; Sweden; nutrient; effects) 

This article summarizes the eutrophication of two 

sea areas: 1) the Baltic Sea and in particular a 

coastal area which is affected primarily by effluents 

from a tertiary sewage plant; and 2) the Kattegat 

and in particular the Laholm Bay, which is affected 

primarily by nutrient inputs from agriculture and 

forest land. During this decade, inputs of N and P 

to these large sea areas have increased by factors of 

approximately 4 to 6 and >8, respectively. Internal 

nutrient transport and atmospheric deposition is 

discussed. It is concluded that N availability 

generally limits primary production, except in 

Bothnian Bay, where P has this role. P is, 

however, the limiting factor at times in some 

coastal areas, and in the Baltic for N-fixing bluegreen-algae, which bloom in warm summers when 

low NIP ratios prevail. Noted eutrophication effects 

are localized elevated primary production, decreased 

depth distribution and species richness of 

macroalgae. Eutrophication is most likely a 

stimulus to high benthic infaunal biomass and fish 

biomass in well ventilated sea areas. Other areas 

suffer today from almost permanant (the Baltic 

proper) or seasonal (the Kattegat) hypoxia with 

devastating effects on benthos and demersal fish. 

The N cycle and future possible consequences in 

relation to nutrient input are discussed. It is 

concluded that a reduction by at least half of 

external N and P to the Baltic sea, and N inputs to 

the Kattegat is needed to mitigate the negative 

effects of eutrophication. 

Rosenberg, R., J. S. Gray, A. B. Josefson, and T. 

H. Pearson (1987) Peterson's benthic 

stations revisited. 11. Is the Oslofjord and 

eastern Skagerrak enriched? Journal of 

Experimental Marine Biology and Ecology 

105:219-251. 

{benthic; fauna; abundance; biomass; composition; 

organic enrichment) 

In summer 1985,23 of the stations established in 

1914 by C.G.J. Petersen in the northern Skagerrak 

and the Oslofjord, were revisited to compare the 

present state of the macrobenthic assemblages with 

the previous condition. The methods used were 

similar to those of the original survey. 

The total benthic biomass was significantly 

higher in 1985 by, on average, a factor of 1.8 over 

the whole area. Most of this change was due to a 

highly significant increase of worms, notably 

polychaetes, and a significant increase of ophiuroid 

and echinoid echinoderms. Similarity values in 

terms of biomass suggested that community 

composition had changed-considerably over the 70- 

yr period, in particular in the inner and central 

Oslofjor (11% similarity) but also in the outer 

Oslofjord and the Skagerrak (33% similarity). There 

was no difference in the comparative size of 

individual organisms over the period. 

The observed increase in biomass is greater than 

the increase along the Swedish Skagerrak coast 

observed in the late 1970s, and well beyond the 

observed limits of annual variability in such areas. 

Different causes for the observed changes are 

discussed and it is suggested that a general organic 

enrichment has taken place in the Oslofjord. 

Rosenberg, R., B. Hellman, and B. Johansson 

(1991) Hypoxic tolerance of marine 

benthic fauna. Marine Ecology Progress Series 

79:127-131. 

{hypoxia; oxygen; benthic; fauna; tolerance) 

In stratified coastal marine waters hypoxia is a 

growing problem affecting bottom-dwelling 

animals. Earlier studies suggest oxygen 

concentrations of about 2 ml l-I (Rosenberg, 1980) 

as the lower tolerance limit for many benthic 

species in coastal areas. We exposed several 

conspicuous infaunal species on the NE Atlantic 

continental shelf, contained within their sediment 

habitat, to gradually reduced oxygen concentrations. 

Tolerance to hypoxia for the 8 species examined 

was in the range of 0.5 to 1.0 ml 1-I (8 to 15% 

saturation), which they could tolerate for several 

days to weeks. The ophiuroid Amphiura filiformis 

left its protected position in the sediment at an 

oxygen concentration of 0.85 ml 1-I (13% sat.), 

whereas A. chiajei emerged from the sediment at 

0.54 ml 1-I (8% sat.). 

Rosenberg, R. and L.-0. Loo (1988) Marine 

eutrophication induced oxygen deficiency: 

Effects on soft bottom fauna, western 

Sweden. Ophelia 29:213-225. 
{oxygen; benthic; macrofauna; increase; mortality) 

Marine eutrophication is studied in a 

multidisciplinary project in the Kattegat, western 

Sweden. Input of nutrients has increased severalfold during this century; nitrogen = 6 times, 

phosphorus = 10 times. This has caused elevated 

concentrations of nutrients in the water and 

increased primary production. Some adverse effects 

at the bottom have been noted annually from about 

August through October in the 19801s, e.g. fish 

catches have dropped and mortalities have been 

reported for fish and benthic animals including 

Nephrops norvegicus. 

In this paper we evaluate the effects on benthic 

macrofauna which are suggested to be caused 

mainly by oxygen deficiency occurring especially 

around the depth of a strong halocline ( = 15 m), 

but also along a bottom transect down to 57 m. At 

some sites the benthic communities were 

impoverished, and at others only some species 

seemed to be especially sensitive, e.g. the bivalve 

Abra alba and the brittle star Amphiurafiliformis. 

In shallow ( < 10m ) exposed waters an annually 

occurring mortality of molluscs was observed over 

large areas from 1980 onwards. Affected species 

were mainly the bivalves Cardium edule and Mya 

arenaria. The causes behind this mortality may also 

be oxygen deficiency occurring during calm weather 

periods. 

Rydberg, L., L. Edler, S. Floderus, and W. GranBli 

(1990) Interaction between supply of 

nutrients, primary production, 

sedimentation and oxygen consumption 

in SE Kattegat. Ambio 19: 134- 14 1. 

{nutrients; nitrogen; phosphorus; oxygen; Baltic) 

Intensive measurements of nutrient fluxes, carbon 

and nitrogen assimilation, sedimentation and 

oxygen consumption within the SE Kattegat are 

used in an effort to follow the coupling between 

increasing nutrient supply and decreasing deep-water 

oxygen concentration. The investigated area is a 

part of the strongly stratified Baltic estuary, with a 

hampered deep-water exchange and a large supply of 

nitrogen from nearby anthropogenic sources. 

Measurements were undertaken during 198 1 - 1988. 

The results are discussed, mainly in terms of annual 

or seasonal mean values. The average primary 

production, 11.4 mol C m-2 (corresponding to 45 

mmol C m-2 d-l during the "productive season"; 

from March to October), was surprisingly well 

correlated with the uptake of inorganic nitrogen, 

which averaged 1.68 mol N m-2 (1 1.1 mol C m-2, 

using the Redfield ratio). The oxygen consumption 

(deep water and benthic) which averaged 20 mol02 

m-2 d-l, was well correlated with the measured 

nitrate uptake and with the external supply of 

nitrate to the surface water. The consumption was 

twice as high as that in the open Kattegat. 

Ryther, J. H. and W. M. Dunstan (1971) 

Nitrogen, phosphorus, and eutrophication 

in the coastal marine environment. 

Science 171:1008-1013. 

{nutrients; nitrogen; phosphorus; phytoplankton) 

The distribution of inorganic nitrogen and 

phosphorus and bioassay experiments both show 

that nitrogen is the critical limiting factor to algal 

growth and eutrophication in coastal marine waters. 

About twice the amount of phosphate as can be 

used by the algae is normally present. This surplus 

results from the low nitrogen to phosphorus ratio 

in terrigenous contributions, including human 

waste, and from the fact that phosphorus 

regenerates more quickly than ammonia from 

decomposing organic matter. Removal of 

phosphate from detergents is therefore not likely to 

slow the eutrophication of coastal marine waters, 

and its replacement with nitrogen-containing 

nitrilotriacetic acid may worsen the situations. 

Saad, M. A. H. and E. I. M. Hemeda (1992) 

Effect of pollution on the Western 

Harbour of Alexandria. I. Environmental 

characteristics. In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{PHI 

The Western Harbour of Alexandria has an area of 

about 1962 acres and a water depth ranging from 6- 

14 m. This harbour is protected by two water 

breaks and is divided into two areas; the inner and 

outer ports. The water quality of the harbour has 

deterioated due to the effects of various pollutants 

discharged from ships at anchor and sewage waste 

outfalls. Variations of some of the environmental 

conditions in the harbour were studied. 

Temperature measurements generally decreased with 

depth. The highest average water temperatures were 

recorded in July and August, whereas the lowest 

were in January and February. The Secchi disc 

readins showed remarkable variations: the 

minimum and maximum regional average Secchi 

values were accompanied by the highest and lowest 

regional averages of suspended matter (SM). The 

irregularity in the vertical distribution of SM 

possibly coincided with turbulence of the wsater 

column from navigation and the effect of pollution. 

Salinity values generally increased with depth. The 

discharge of sewage wastes from numerous ships 

and land-based sources decreased the surface salinity. 

The highest regional average salinity values were 

found at locations far from the direct effect of 

sewage pollution. The pH values generally 

decreased with depth: low pH values were found at 

locations directly affected by sewage wastes. 

Sales, D., A. Gbmez, and D. Cantero (1983) 

Incidence of urban sewage disposal in the 
salt-ponds areas of the south of the Bay 

of Cadiz. Marine Pollution Bulletin 14:447- 

452. 

(Cadiz Bay; oxygen; salt pond} 

The highest pollution indices in the marsh and 

tide-lands situated to the south of the Bay of Cadiz 

have been evaluated. At the present, the salt 

industry of this area is being transformed into 

mariculture installations. This study consisted of 

the systematic evaluation of different parameters at 

26 sampling stations during the period 1976- 198 1. 

The results suggest that extensive zones are too 

seriously affected to make use of them as marine 

piscifactories. 

Samanidou, V., K. Fytianos, and G. Vasilikiotis 

(1989) Distribution of nutrients in the 

Thermaikos Gulf, Greece. Toxicological and 

Environmental Chemistry 20-21:29-37. 

(nutrients; Greece; methods; nitrogen; phosphorus; 

distribution; sewage} 

The distribution of nutrients (phosphate, nitrate, 

nitrite and ammonium) in water samples from 

eleven sampling stations in the Thermaikos Gulf 

was studied during a period of one year (1985- 

1986). Significant seasonal fluctuations were 

observed mainly for nitrate and ammonium, with a 

maximum during winter and a minimum in the 

summer. The obtained data showed that the 

nutrients are present at relatively high 

concentrations near the agricultural areas, in the 

harbour of Thessalonki and in the area where 

municipal sewages are discharged. 

Sampou, P. and C. A. Oviatt (1991) Seasonal 

patterns of sedimentary carbon and 

anaerobic respiration along a simulated 

eutrophication gradient. Marine Ecology 

Progress Series 72:27 1-282. 

{organic carbon; oxygen uptake; sulfate reduction} 

Concentrations of organic carbon and rates of 

dissimilative sulfate reduction in surface sediments 

of marine mesocosms were examined along an 

experimental eutrophication gradient. 

Phytoplankton biomass increased due to addition of 

inorganic nutrients (N, P, Si). This increase was 

especially pronounced during the winter-spring 

diatom blooms, which increased in magnitude and 

duration along the nutrient gradient. Net system 

production in winter and spring resulted in carbon 

deposition and accumulation in surface sediments 

(maximum net accumulation 17 mol C m-2). 

Benthic remineralization of carbon exceeded 

depositional supply during summer and fall. 

Sediment carbon concentration approached 

background levels in December and February, 

suggesting very little annual accumulation of 

sediment carbon. Sediment oxygen consumption 

and sulfate reduction rates in organic enriched 

sediments were an order of magnitude higher than 

control and were correlated with temperature and 

carbon concentrations (r2=0. 85). Anaerobic 

metabolism was the dominant metabolic pathway 

in control and treated sediments, with 50 to 70% of 

annual carbon remineralization due to sulfate 

reduction. 

Sampou, P. A. (1989) Effects of 

eutrophication on the biogeochemical 

cycling of carbon, oxygen, sulfur and 

energy in coastal marine ecosystems. 

Dissertation for Ph.D., University of Rhode Island. 

{biogeochemical cycling; energy flow; MERL; 

sewage; nutrients; production; ecosystem; oxygen} 

The effects of eutrophication on the 

biogeochemical cycling of the major elemental 

cycles, carbon, oxygen, and sulfur and on energy 

flow, were investigated in experimental marine 

ecosystems (mesocosms). Eutrophic environmental 

conditions were created withln mesocosms by daily 

addition of varying amounts and sources of 

nutrients and sewage sludge. Inorganic nutrient 

loading resulted in an increase in the magnitude and 

duration of winter-spring phytoplankton 

populations and production with a correlated 

increase in sedimentation and accumulation of 

carbon in surface sediments. Net ecosystem 

production during the winter and spring was 

followed by net system heterotrophy during the 

summer and fall with a concomitant decrease in 

sedimentary carbon concentrations. Carbon 

sedimentation and accumulation (up to 19 mol C 

M-~) insurface sediments stimulated sediment 

oxygen consumption and sulfate reduction 

metabolism. Sulfate reduction increased by an order 

of magnitude in carbon enriched sediments and was 

a dominant pathway for benthic carbon 

remineralization in all the mesocosms. However, 

the fraction of benthic metabolism using sulfate for 

a terminal electron acceptor did not increase along 

the nutrient addition gradient when annual 

integrated estimates of metabolism were compared. 

Whole system carbon and oxygen budget showed 

the sediments to be an important and influential 

component in eutrophic marine ecosystems. A 

significant fraction of allochthonous sewage carbon 

was remineralized via sulfate reduction in the 

sediments. This led to the dissociation of carbon 

cycling from oxygen and energy flow in the system 

due to an increased role of sulfur biogeochemistry. 

Accumulation of solid sulfides in the sediments 

from sulfate reduction during the experiment 

explained the discrepancy between carbon dioxide 

and oxygen consumption at the sediment water 

interface (respiratory quotients > 2) and represented 

a large storage of energy originally contained in 

organic matter. 

Oxygen dynamics in eutrophic marine systems 

were regulated by phytoplankton cycles and sulfur 

cycling in the sediments. Initiation of hypoxia 
(oxygen less than 2 mg 1-l) was caused by rapidly 

declining phytoplankton populations and a large 

imbalance between day oxygen production and 

night oxygen consumption in the water column. 

However, high sulfate reduction rates and the 

accumulation of hydrogen sulfide in surface 

sediments was found to be influential in the demise 

of the macrofauna and the precipitation and 

maintenance of oxygen concentrations less than 1 

mg 1-1. 

Sand-Jensen, K. and J. Borum (1983) 

Regulation of growth of eelgrass (Zostera 

marina L.) in Danish coastal waters. 

Marine Technology Society Journal 17:15-21. 

{submerged vegetation; eelgrass; growth; shading) 

Eelgrass is the dominant macrophyte in coastal 

Danish waters. The productivity of eelgrass is at 

the same level as fertile terrestrial crops and it can 

be substantially higher than the phytoplankton 

production in open waters when compared per unit 

surface area. Eelgrass may contribute significantly 

to the total production of all autotrophic 

components in shallow coastal areas. 

The above-ground biomass of eelgrass is more 

variable than the below-ground biomass due to a 

higher turnover rate and exposure to physical 

factors of the former. In shallow water, biomass of 

eelgrass increases rapidly from spring to early 

summer and stabilizes at a high level during late 

summer. Growth is continuous throughout the year 

although it occurs at a higher rate during spring and 

summer. Individual plants produce about 16 leaves 

and rhizome segments per year. During spring and 

summer each plant produces a new leaf every 9-15 

days and their life span is about 2 months. The 

rapid turnover is important in keeping down the 

epiphytic load by producing new tissue. It also 

produces a continuous supply of senescing leaves 

which presumably leads to stabilized energy 

transformations within the detritus food web. Since 

the leaf biomass increases from spring to summer, 

the ratio of daily productivity to biomass (PIB) 

falls because of increased self-shading. Water 

temperature and availability of nutrients do not 

affect seasonal variation in leaf productivity. 

Nutrient availability may, however, affect the 

biomass as a yield limiting factor. 

Eutrophication of coastal waters changes in the 

balance among different autotrophic components. 

Growth of phytoplankton, epiphytes and freefloating algae, which take up inorganic nutrient 

from the free water phase, is greatly stimulated. 

This will result in decreased growth rates, depth 

penetration and, eventually, survival of eelgrass 

though shading. 

Biomass and growth measurements of epiphytic 

algae provide a convenient method for assessing the 

effect of inorganic nutrient loading in different parts 

of a receiving area. Eelgrass is suitable as an 

indicator of trace-metal pollution as it occurs over a 

widespread area and chemical analysis for trace 

metals is a relatively simple procedure. Due to the 

high growth potential of eelgrass leaves (about 5-6 

cm leaf turion-l day-l), growth rates of leaves can 

be used as a very sensitive method of assessing the 

effects of industrial waste products both in the field 

and in laboratory tests. 

Sandulli, R. and M. d. N. Giudici (1989) Effects 

of organic enrichment on meiofauna: A 

laboratory study. Marine Pollution Bulletin 

20:223-227. 

{meiofauna; sewage; nematode; harpacticoid; 

copepod; ratio) 

A 84 day laboratory experiment indicates the 

decline of intertidal meiofauna abundance may be 

directly related to treatment level and time of 

exposure to sewage sludge. The observed changes 

revealed a marked decrease of nematodes and total 

disappearance of mesobenthic harpacticoids while 

the non-interstitial copepod species were little 

affected. The possibility of a new interpretation of 

the NematodeICopepod ratio as an index of organic 

pollution in sediments is discussed. 

Schiewer, U., R. Borner, and N. Wasmund (1988) 

Deterministic and stochastic influence of 

nutrients on phytoplankton function and 

structure in coastal waters. Ki e 1 e r 

Meeresforschungen, Sonderheft 6: 173- 183. 

{phytoplankton; nutrients; composition; 

abundance; chlorophyll a) 

Knowledge of how phytoplankton responds to 

nutrient inputs is essential for water management 

and for minimizing eutrophication. Only processes 

that are deterministic, i.e. that can respond as 

algorithms, are controllable. The study area is the 

chain of inshore waters (so-called Bodden) south of 

the Darss-Zingst peninsula- shallow eutrophic 

waters of estuarine character in the Southern Baltic. 

Monitoring programmes and laboratory 

experiments have revealed an annual periodicity of 

the phytoplankton and of the physico-chemical 

factors influencing it. On the basis of these results, 

experiments were carried out in enclosures to study 

the effects of nutrient loading on phytoplankton. 

The purpose was to test the feasibility of 

influencing phytoplankton development under field 

conditions during the transition period from late 

spring to mid-summer. 

This contribution presents results from the 1985 

shallow water enclosure experiments (FLAK 85) 

which demonstrate that 

- the scale of phytoplankton reactions and the 

species involved are stochastic in character and are 

governed by stochastic interactions between 

meteorological events and water exchange processes 

in the chain of Bodden; 
- all processes affecting phytoplankton growth are 

deterministic in character, conforming to simple 

batch theories: simultaneous addition of nitrogen 

and phosphorus favours green algae, and in 

exceptional cases one algal species became 

dominant 

- nutrient loadings do not affect the time of 

transition to the mid-summer phytoplankton 

population, the most important regulating factor 

obviously being temperature. 

Schindler, D. W. (1981) Studies of 

eutrophication in lakes and their 

relevance to the estuarine environment. 

In: B. J. Neilson and L. E. Cronin (ed.) Estuaries 

and Nutrients. Humana Press, Clifton, New Jersey. 

{lake; estuary; comparison; Vollenweider; nutrient; 

composition) 

No abstract 

Schlungbaum, G. and G. Nausch (1988) 

Nutrient turnover at the sedimentlwater 

interface in shallow eutrophic coastal 

waters. Kieler Meeresforschungen, Sonderheft 

6:184-190. 

{sediment; interface; nutrient; phosphorus; 

phosphate; phytoplankton; primary production) 

The nutrient situation in eutrophic shallow 

coastal water systems is characterized by particular 

features. In addition to water exchange with the 

open sea, interactions at the sedimentlwater 

interface play a significant role. With the help of 

investigation of phosphate sorptionldesorption 

balances and on the denitrification capacity of 

shallow coastal waters, the following general 

conclusions can be made: The phosphate 

concentrations in the water are primarily determined 

by physico-chemical reactions with the sediment, 

and less by the rhythm of the phytoplankton 

primary production. Through intensive interactions 

between sediment and water, a phosphate 

"equilibrium" concentration which fluctuates very 

little is maintained over the whole year. High 

primary production rates are possible at these low 

equilibrium concentrations of phosphate sorption 

capacities. With the development of nearly 

anaerobic conditions, nitrate reduction can occur at 

a rate, when the appropriate amount of nitrate is 

made available to the reaction. Under optimal 

conditions for nitrate reduction, nitrate 

concentration is also not a parameter characterizing 

the nature of the water body. 

Schramm, W., D. Abele, and G. Breuer (1988) 

Nitrogen and phosphorus nutrition and 

productivity of two community forming 

seaweeds (Fucus vesiculosus, Phycodrys 

rubens) from the Western Baltic (Kiel 

Bight) in the light of eutrophication 

processes. Kieler Meeresforschungen, Sonderheft 

6:221-240. 

{nitrogen; phosphorus; seaweeds; productivity; 

Fucus; Phycodrys; Baltic) 

Considerable amounts of nutrients enter, as a 

result of human activity, the Western Baltic Sea in 

the form of discharge of seepage from land or as 

atmospheric input. Benthic macrophytes which 

preferably inhabit the nutrient recipient shallow 

near-shore areas, and where they often constitute the 

major primary producers may thus play an 

important role in eutrophication processes by 

uptake and accumulation on nutrient elements, 

enhanced primary production and following 

degradation of the produced organic material. 

Nutrient uptake, binding capacity and illumination 

as well as nutrient dependent growth of the two 

community forming seaweeds Phycodrys rubens 

and Fucus vesiculosus have been investigated and 

related to the seasonal patterns of the nutrient 

conditions in their respective habitats. Nutrient 

concentrations, both phosphorus and nitrogen, in 

the algal beds are markedly higher than in the 

surface water of the open Kiel Bight. In general, the 

seaweeds seem not to be nutrient limited under 

natural conditions except for nitrogen in Phycodrys 

during summer. Nutrient tissue contents are 

saturated only for nitrogen during winter. The 

significance of these findings is discussed in view 

of the observed increasing nutrient levels and 

changes of the vegetation in the Kiel Bight. 

Schultz, S., G. Wrtebjerg, G. Behrends, G. Breuel, 

P. Ciszewski, U. Horstmann, K. Kononen, E. 

Kostrichkina, J. Leppanen, F. Mohlenberg, 0. 

Sandstrom, M. Viitasalo, and T. WillCn (1992) 

The present state of the Baltic Sea 

pelagic ecosystem - an assessment. In: 

G. Colombo, I. Ferrari, V. U. Ceccherelli and R. 

Rossi (ed.) Marine Eutrophication and Population 

Dynamics. Olsen & Olsen, Fredensborg. 

{Baltic Sea; assessment; pelagic biology; 

phytoplankton assemblage) 

In recent years, the decrease of salinity as well as 

of the temperature and oxygen in the deep water has 

continued in the Baltic Sea. In addition, the 

content of phosphate and nitrate has declined in the 

winter surface water at least during the last two 

years. However, both the primary production 

capacity and the assimilation numbers have been 

increasing for most sub-areas of the Baltic Sea, 

indicating a higher phytoplankton activity. The 

stock of zooplankton increased in tha Arkona, 

Bornholm and Eastern Gotland Seas. Changes in 

the species composition or in the structure of the 

phytoplankton and zooplankton community have 

not been observed. 

The present status of the Baltic Sea pelagic 

system reflects clear signs of eutrophication. The 

reasons for this are seen in the anthropogenic 
impact in conjunction with the features of the 

Balitc Sea and a stagnation period lasting thirteen 

years. 

Schulz, S. and G. Breuel (1981) Response of 

plastic-bag enclosed summer 

phytoplankton to a nutrient input under 

in situ conditions in the Arkona Sea 

(Baltic Sea). Kieler Meeresforschungen, 

Sonderheft 5:9 1- 100. 

{phytoplankton; nutrients; Baltic; primary 

production; Arkona; biomass; composition) 

In July 1878 several experiments with enclosed 

summer phytoplankton communities were carried 

out in the Arkona Sea using 25 1 cylindrical bags 

made of 0.2 mm polyethylene film. These 

experiments concerned the response of 

phytoplankton to nutrient input. Phytoplankton 

biomass, primary production and chlorophyll 

content increased rapidly in enriched water, in 

contrast to the controls where changes were 

negligible. Nutrients were depleted to the same 

levels as in the controls after a few days. The palgae were the first to respond to nutrient input, 

followed later by diatoms and dinoflagellates. In the 

course of the experiments, however, the energy was 

stored to an increasing extent in the cyanophytes at 

the expense of other groups where energy content 

remained constant or decreased slightly. 

Schulz, S., G. Breuel, and A. Irmisch (1988) 

Simulated eutrophication in enclosure 

experiments in the Arkona Sea. Kiele r 

Meeresforschungeb, Sonderheji 6:73-84. 

{Arkona Sea; Baltic; plankton; nutrients; primary 

production; biomass; productivity) 

In spring, summer and autumn enclosure 

experiments were performed in the central part of 

the Arkona Sea. The natural water with the 

plankton community was enriched by nutrients to 

about winter levels, and to the double of these 

concentrations. 

In spring and summer, the phytoplankton 

responded with rapid uptake of nutrients and an 

increase in primary production and biomass. In 

autumn, the uptake of nutrients was also fast, 

whereas productivity did not increase, and biomass 

only in diatoms. Not only did the production 

increase with higher nutrient supply, but also the 

productive season was prolonged. This fact is 

important for the function of the pelagic system in 

the Baltic, because the biomass and nutrients 

remaining in the spring bloom determine to a great 

extent the productivity of the whole year. 

Seiki, T., E. Date, and H. Izawa (1991) 

Eutrophication in Hiroshima Bay. Marine 

Pollution Bulletin 23:95-99. 

{nutrients; primary production; budget) 

Several researches for eutrophication in the 

northern Hiroshima Bay were carried out to 

investigate nutrient budgets in aquatic circulation 

processes, i.e. primary production, settling flux of 

particulate organic matter (POM) and the benthic 

remineralization. 

It became apparent from these investigations that 

primary production contributed more to the organic 

pollution in the northern bay than land based 

organic loadings by a factor of 10, that approx. 70- 

80% of the POM originating in phytoplankton 

were easily remineralized into inorganic nutrients 

during the settling process in water, and that 

approx. 60% of nitrogen and 70% of phosphorus in 

POM settled on the sediments might be returned to 

the water column by the release from benthic 

sediment. 

Seitzinger, S. P. and S. W. Nixon (1985) 

Eutrophication and the rate of 

denitrification and N20 production in 

coastal marine sediments. Limnology and 

Oceanography 30: 1332-1339. 

{denitrification; N20 production; nutrient; MERL) 

Large (13 m3, 5m deep) microcosms with coupled 

pelagic and benthic components were used to 

measure the effect of nutrient loading and 

eutrophication in coastal marine ecosystems on the 

rates of benthic denitrification (N2) and N20 

production. After 3 months of daily nutrient 

additions, average denitrification rates ranged from 

about 300 umol N m-2 h-l in the sediments of the 

control microcosm to 880 in the most enriched 

microcosm, which received 65 times the nutrient 

input of the control. Increases in the production of 

N20 were more dramatic and increased by a factor - 

of about 100, from 0.56 N m-2 h-l in the control 

to 51 in the most enriched microcosm. Although 

there was a clear increase in the denitrification rate 

in the more eutrophic systems, the amount of fixed 

nitrogen removed was a constant or progressively 

smaller fraction of the nitrogen input. Even in the 

most enriched microcosm, at least 16% of the N 

input was removed by denitrification. 

Seliger, H. H. (1989) Mechanisms for red 

tides of Pyrodinium bahamense var. 

compressum in Papua New Guinea, Sabah 

and Brunei Darussalam. In: G. M. 

Hallegraeff and J. L. Maclean (ed.) Biology, 

Epidemology and Management of Pyrodinium red 

Tides. Fisheries Department, Ministry of 

Development, Manila. 

{red tides; blooms; abundance) 

Mechanisms are proposed to explain the 

development of red tides of the toxic dinoflagellate 

Pyrodinium bahamense var. compressum along the 

northern and southern coasts of Papua New Guinea 

and along the northwest coast of Borneo, from 
Sabah to Burnei Darussalam. The "spread of red 

tides to areas not previously affected and the 

increased frequency of annual occurrences of 

paralytic shellfish poisoning in already colonized 

areas are suggested to be the result of coastal 

hypertrophication. Increased nutrients along 

coastlines permit sufficient reproduction of 

dinoflagellates in entrained surfaces patches to 

compensate dilution losses, enhancing emigration 

over longer distances. 

Sellner, K. G. and M. M. Olson (1985) 

Copepod grazing in red tide of 

Chesapeake Bay. In: D. M. Anderson, A. W. 

White and D. G. Baden (ed.) Toxic Dinoflagellates. 

Elsevier, New York. 

{secondary production; zooplankton; bloom; 

Chesapeake Bay; red tide; grazing; dinoflagellates) 

Copepod grazing of dinoflagellate blooms in 

Chesapeake Bay and its tributaries was investigated 

using 14c-labelling procedures. Grazing rates for 

Eurytemora affinis, the late winter - early spring 

dominant herbivore, on Heterocapsa triquetra were 

low with a mean rate of 0.011 +- 0.002 ml 

c~~e~od-~h-~. However, at a cell density of 2.43 X 

lo5 m1-l, the ingestion rate corresponds to 1.88 

ugC copepod-l h-l, or approximately one-third of 

the copepod's daily ration. Fecal pellet production 

every 6-9 min. Low grazing rates for the spring-fall 

dominant herbivore, Acartia tonsa, were also 

recorded where the copepod was feeding on blooms 

of Gymnodinium sp. and Katodinium rotundatum. 

Mean rates ranged from 0.005 - 0.048 ml copepodlh-l, corresponding to estimated ingestion rates of 

0.038 - 0.102 ugC copepod-lh-l, well below the 

rate recorded for Eurytemora. These data indicated 

that A. tonsa may not remove significant quantities 

of dinoflagellate carbon in Chesapeake Bay and its 

sub-estuaries. 

Silva, E. S. (1985) Ecological factors 

related to Prorocentrum minimum blooms 

in Obidos Lagoon (Portugal). In: D. M. 

Anderson, A. W. White and D. G. Baden (ed.) 

Toxic Dinoflagellates. Elsevier, New York. 

{red tide; bloom; environmental condition) 

Prorocentrum minimum Schiller red waters have 

often been observed in Obidos Lagoon and have 

caused toxicity of bivalve there. Particular attention 

is given to two of those blooms, separated by 

about 10 years, in May-June 1973 and in JanuaryFebruary 1983. A comparative study of 

environmental conditions during the two red waters 

of P. minimum is presented. 

Skjoldal, H. (1992) 

Personal Communication. 

Skjoldal, H. R. and I. Dundas (ed.) (1991) The 

Chrysochromulina polylepis bloom in 

the Skagerrak and the Kattegat in MayJune 1988: Environmental conditions, 

possible causes, and effects. ICES, 

Copenhagen. 

{bloom; Chrysochromulina; causes) 

No abstract 

Smayda, T. J. (1989) Primary production 

and the global epidemic of phytoplankton 

blooms in the sea: A linkage? In: E. M. 

Cosper, N. M. Bricelj and E. J. Carpenter (ed.) 

Novel Phytoplankton Blooms; Causes and Impacts 

of Recurrent Brown Tides and Other Unusual 

Blooms. Springer-Verlag, Berlin. 

{primary production; phytoplankton bloom) 

No abstract 

Smayda, T. J. (1992) A phantom of the 

ocean. Nature 358:374-375. 

{dinoflagellate; red tides; foodweb) 

No abstract 

Smayda, T. J. and Y. Shimizu (ed.) (1993) 

Toxic Phytoplankton Blooms in the Sea 

(Proceedings of the Fifth International 

Conference on Toxic Marine 

Phytoplankton, Newport, Rhode Island, 

U.S.A., 28 October -1 November 1991). 

Elsevier, Amsterdam. 

No abstract 

Smayda, T. J. and A. W. White (1990) Has 

there been a global expansion of algal 

bloom? If so, is there a connection with 

human activities? In: E. GranBli, B. 

Sundstrom, L. Edler and D. M. Anderson (ed.) 

Toxic Marine Phytoplankton. Elsevier, New York. 

{phytoplankton; algae; bloom; nutrient 

enrichment) 

No abstract 

Smith, D. W. (1984) Responses of 

aufwuchs, phytoplankton and 

macrophytes to municipal and industrial 

wastes in San Francisco Bay and bay 

analog systems. Dissertation for Ph.D., 

University of California, Berkeley. 

{auhwuchs; phytoplankton; seaweeds; San 

Francisco Bay; method; mesocosm) 

The use of aquatic organisms which colonize 

artificial substrates (aufwuchs) is currently being 

developed as a method of assessing localized effects 

of municipal and industrial discharges into San 

Francisco Bay, California by state regulatory 

agencies. The method is intended to provide an 

inexpensive and robust bioassay which will serve 

as an index of wastewater effects on the 

phytoplankton community, since the effects on 
phytoplankton cannot be directly observed. In this 

study functional or physiological responses 

(photosynthesis, respiration, chlorophyll a, and 

biomass) were measured in the laboratory, in 

mesocosms and in the field to characterize the 

response of the aufwuchs community to wastewater 

and some of its individual stimulatory and toxic 

components. The mesocosm consisted of 12 

outdoor 3000L tanks with a constant supply of 

baywater. The response of aufwuchs under 

experimental conditions confirmed and elucidated 

the causes of the stimulatory response of aufwuchs 

grown on substrates moored in dilution-fields in 

San Francisco Bay. 

The effects of colloidal and microbial components 

of wastewater on earlier colonizing and later rapid 

growth phase of the auchwuchs community were 

determined to be unimportant with respect to the 

measurable differences caused by exposure to 

wastewater. Wastewater was found to stimulate 

growth of aufwuchs. Inorganic nitrogen was 

determined experimentally and deductively to be the 

causative stimulatory component of wastewater. 

Aufwuchs were also shown to be sensitive to toxic 

compounds found in the wastewater. 

The stimulatory and toxic response of aufwuchs 

amplified the response of phytoplankton. Various 

aspects of the Bay's physical and chemical 

environment such as light turbulence and ambient 

nutrient concentration may result in this 

amplification by moderating factors. Aufwuchs, 

then, may serve as an early warning of potential 

effects of wastewater on phytoplankton. 

Strategies of substrate deployment and harvest 

were also assessed. Increased biomass, facilitated by 

relatively long incubation intervals (7-14 days), 

was shown experimentally to maximize the 

precision of the technique. The precision of the 

method is adequate from a regulatory point of view 

to delineate the region where biota are influenced by 

wastewater. 

Smith, S. V. (1981) Responses of Kaneohe 

Bay, Hawaii, to relaxation of sewage 

stress. In: B. J. Neilson and L. E. Eugene (ed.) 

Estuaries and Nutrients. Humana Press, Clifton, 

New Jersey. 

{nitrogen; sewage; nutrient; cycling; Hawaii} 

Kaneohe Bay is a subtropical coral reeflestuary 

complex which was subjected to increasing sewage 

loading; that sewage was diverted in 1977 and 

1978. We have treated the loading and diversion as 

a controlled , total-ecosystem experiment to 

evaluate the chemical and biological responses to 

external subsidy of nutrients. We here consider the 

Bay's response to sewage loading and to its 

diversion largely in the context of a nitrogen 

budget. 

Even the most heavily impacted portion of the 

Bay showed only modest increases in dissolved 

nitrogen levels within the water column. 

Particulate materials including plankton biomass, 

as well as dissolved inorganic phosphorus, were 

elevated substantially. The benthos showed 

increased biomass and metabolic rates, especially of 

heterotrophs responding to fallout of organic 

particulate materials. Nutrient recycling within the 

Bay was the major immediate source for the 

nutrients for the observed rapid metabolic activity. 

Sewage accounted for approximately 80 percent of 

the -inorganic nitrogen and 90 percent of the 

inorganic phosphorus delivery to Kaneohe Bay. 

Diversion lowered the land-derived inorganic 

nitrogen and phosphorus input to the Bay by 70-80 

percent. Virtually all components of the system 

have responded to this diminished nutrient subsidy, 

but the water column nutrient washout and 

biological recovery are predictably occurring more 

rapidly than the benthos responses. 

Snelgrove, P. V. R. and J. B. Lewis (1989) 

Response of a coral-associated crustacean 

community to eutrophication. Marine 

Biology 101:249-257. 

{crustacea; coral; Madracis; Barbados; composition; 

density) 

Seventy-five samples of the crustacean fauna 

associated with the coral Madracis mirabilis 

(Duchassaing and Michelotii) were obtained at five 

sites along the west coast of Barbados in 1985. 

Samples were analyzed to determine whether any 

community response to eutrophication was evident. 

Ordination (multidimensional scaling) and 

classification (group average clustering) based on 

Bray-Curtis dissimilarity measures separated the 

fauna of highly eutrophied sites from those that 

were less eutrophied. Dominance plots based on 

relative abundance and log-normal plotting methods 

were ineffective in distinguishing the fauna of 

eutrophied sites. The crustacean fauna responded to 

eutrophication mainly by reduction of density of 

animals. No differences in species richness, number 

of site endemic species or rare species were 

observed between highly eutrophied and less 

eutrophied sites. Nine "indicator" species of 

eutrophy were identified on the basis of strong 

density differences between sites. 

Soulsby, P. G., D. Lowthion, and M. Houston 

(1982) Effects of macroalgal mats on the 

ecology of intertidal mudflats. Marine 

Pollution Bulletin 13: 162- 166. 

{macroalgal mats; mudflat; benthos; abundance; 

biomass} 

The standing biomass and distribution of the 

invertebrate fauna of mudflats covered by 

macroalgal mats has been determined and compared 

with that of mudflats unaffected by algae. The 

faunal biomass of algal-covered mud was 

consistently higher than uncovered mud, mainly 
due to the presence of large populations of the snail 

Hydrobia ulvae. The structure of the faunal 

composition of algal-covered mud was altered and 

the numbers of at least one species of polychaete 

worm markedly reduced. The effects of these 

changes on the bird population exploiting the 

mudflats is discussed. 

Soulsby, P. G., D. Lowthion, and H. A. C. 

Montgomery (1985) The role of sewage 

effluent in the accumulation of 

macroalgal mats on intertidal mudflats in 

two basins in southern England. 

Netherlands Journal of Sea Research 19:257-263. 

{macroalgae; nutrients) 

Two intertidal basins in southern England, 

Portsmouth Harbour and Langstone Harbour, both 

support large crops of the macroalgae Ulva and 

Enteromorpha each summer. Both Harbours have 

received discharges of sewage but the discharge in 

Portsmouth has been removed. Data from surveys 

of algal distribution, standing biomass, tissue 

nutrient concentration and water column nutrient 

concentration have been evaluated in order to 

deternine the relationship between effluent discharge 

and macroalgal growth. No direct relationship 

between water column nutrient concentration and 

macroalgae biomass or area cover was found in the 

field. Concentrations of Nitrogen and 

Nitrogen:Phosphorus ratio in the tissues suggest 

that Nitrogen is stored in excess during the growing 

season. It is concluded that the evidence strongly 

indicates that further increases in sewage derived 

nutrient will not result in higher standing crops of 

algae than currently observed. 

Spencer, C. P. (1985) The use of plant 

micro-nutrient and chlorophyll records as 

indices of eutrophication in inshore 

waters. Netherlands Journal of Sea Research 

19:269 -275. 

{indices; chlorophyll; nutrients) 

The records for 14 years of plant micro-nutrient 

concentrations and the quantities of chlorophyll in 

Liverpool Bay are reviewed. The complicated and 

varied hydrographical regimes which characterize 

the area preclude the use of the former variables as 

precise indicators of the extent of enrichment of the 

water. There is nonetheless evidence that the area 

can sometimes support high standing crops of 

phytoplankton. Advective dispersal processes and 

variations in their intensity are probably 

particularly important in controling the magnitude 

of these crops of phytoplankton. It is suggested 

that adequate long term records of the frequency and 

durations of such blooms are likely to be a more 

useful indication of the extent or eutrophication of 

the area and of any long term trends which might 

occur than are the hydrochemical data. 

Stachowitsch, M. (1991) Anoxia in the 

Northern Adriatic Sea: rapid death, slow 

recovery. In: R. V. Tyson and T. H. Pearson 

(ed.) Modern and Ancient Continental Shelf 

Anoxia. The Geological Society, London, England. 

{Adriatic; oxygen; benthos; anoxia; mortality) 

The Northern Adriatic Sea is characterized by the 

full range of features associated with sensitive 

marine ecosystems. In the Gulf of Trieste, 

mortalities due to anoxia have been reported in 

1974, 1983 and 1988. The anoxic event in 

September 1983 led to mass mortality in a highly 

developed and wide-ranging community in the Gulf. 

This destroyed over 50% of the epifaunal biomass 

in only two days, and over 90% within four days. 

The increased frequency, severity, and range of 

recent oxygen deficiencies suggest that, in addition 

to meteorological and hydrographic factors, 

anthropogenic influences are a contributing or 

enhancing factor in the onset of anoxic conditions 

and associated mass mortalities. These 

anthropogenic influences may be subsumed under 

the heading eutrophication. If eutrophication only 

slightly increases the rate of oxygen decrease below 

the thermocline, it may advance the point at which 

critical oxygen levels are attained by several days. 

Since only a few days of anoxic conditions are 

required to destroy this benthic community (with an 

equally short time required for reoxygenation) the 

degree of eutrophication may determine whether an 

ecological catastrophe occurs or not. 

The recolonization process between 1984 and 

1987 has been slow: the total biomass and the 

relative contribution of the designating species have 

remained distinctly below previously recorded 

values. Additional anthropogenic influences 

(intensive dredging) may further impede 

recolonization. A further, small-scale mortality was 

registered in 1988. 

Stambler, N., N. Popper, Z. Dubinsky, and J. 

Stimson (1991) Effects of nutrient 

enrichment and water motion on the coral 

Pocillopora damicornis. Pacific Science 

45:299-307. 

{nutrient; coral; phosphorus; nitrogen; density; 

growth) 

Exposure of the hermatypic coral Pocillopora 

damicornis (Linnaeus) to elevate levels of dissolved 

inorganic phosphorus did not affect the colony or 

the zooxanthellae. Exposure to elevated levels of 

dissolved inorganic nitrogen and inorganic nitrogen 

+ phosphorus led to an increase in algal density, 

and as a result, to an increase in the chlorophyll 

concentration. These latter two experimental 

enrichments slowed skeletal growth rate of the 

corals, probably because of a decrease in the 

photosynthetic rate of the algae and perhaps a 

decrease in the translocation of photosynthetic 

products from the algae to the coral. The algae 
probably used the photosynthetic energy for their 

own increased growth. Experimental manipulation 

of water motion used in these experiments did not 

affect the coral or the symbiotic algae. 

Stockner, J. G. and D. D. Cliff (1979) 

Phytoplankton ecology of Vancouver 

Harbor. Journal of the Fisheries Research Board 

of Canada 36: 1-10. 

{phytoplankton; Vancouver Harbor; primary 

production; sewage) 

Phytoplankton production and distribution were 

examined over a 2-yr period in the Burrard Inlet 

system, which includes a true fjord (Indian Arm), a 

shallow blind inlet (Port Moody arm), and a 

turbulent narrows region that is contiguous to the 

Port of Vancouver. Greatest annual production 

occurred in Port Moody Arm with a mean of 532 g 

C m-2 yr-l while the lowest values were in Indian 

Arm and the Narrows region, averaging about 260 

g C m-2 yr-l. Nitrate and zooplankton grazing were 

the main factors limiting phytoplankton production 

in Indian Arm, while flushing and poor light 

conditions influenced phytoplankton growth in the 

Narrows and outer Burrard Inlet. Most of the 

discharges of domestic and industrial wastes have 

been diverted to the Fraser River, and Vancouver 

Harbor can be considered relatively clean and 

pollution-free because of strong tidal mixing and 

seaward flushing. The only sign of eutrophication 

in the inlet is in Port Moody Arm where sufficient 

nutrients from sewage discharges and a relatively 

stable mixed-layer depth create near optimal 

conditions for phytoplankton growth. Daily 

production here is among the highest recorded in 

the literature for Pacific coastal marine waters. 

Stockner, J. G., D. D. Cliff, and K. R. S. 

Shortreed (1979) Phytoplankton ecology of 

the Strait of Georgia, British Columbia. 

Journal of the Fisheries Research Board of Canada 

36:657-666. 

{phytoplankton; primary production; chlorophyll a; 

Strait of Georgia) 

Observations of phytoplankton production, 

abundance, and distribution were made at 16 

stations in the Strait of Georgia from 1975 to 

1977. The discharge of turbid Fraser River water 

exerts a strong influence on phytoplankton 

production and distribution in surface waters by 

rapid light attenuation and horizontal advection. At 

plume boundaries and back eddied where light 

conditions improve, very high production occurs 

(>4-5 g C m-2 d-I), because of rapidly replenished 

nutrients supplied by the Fraser River. Advection, 

turbulence, zooplankton grazing, and summer 

nitrate depletion collectively impart a 

heterogeneous distribution pattern to phytoplankton 

in the surface waters of the Strait of Georgia. Mean 

annual production varies from lows of 150 g C nr2 

in Fraser River plumes to highs over 500 g C m-2 

in sheltered boundary waters of inlets. Recent 

increases in ammonia and nitrate from land drainage 

and domestic sewage, mainly through the Fraser 

River, are related to increases in phytoplankton 

standing stocks in the Strait. 

Stockner, J. G. and K. S. Shortreed (1988) 

Response of Anabaena and Synechococcus 

to manipulation of nitrogen: Phosphorus 

ratios in a lake fertilization experiment. 

Limnology and Oceanography 33: 1348- 136 1. 

{lake; nutrient addition; picoplankton; chlorophyll; 

NIP ratio; bacteria) 

Additions of inorganic nitrogen and phosphorus to 

selected areas of oligotrophic Kennedy Lake began 

in 1978 and resulted in development of late-summer 

blooms of nitrogen-fixing cyanobacteria Anabaena 

circinalis in 1981 and 1982. In subsequent years the 

blooms were successfully eliminated by increasing 

the molar N:P ratio of added nutrients to 35: 1. The 

nutrient additions also resulted in dramatic increases 

in both autotrophic (predominantly Synechococcus 

sp.) and heterotrophic (bacteria) picoplankton 

numbers, with respective maxima of 4.0 x lo5 mland >4.0 x lo6 ml-l. Chlorophyll concentrations 

were generally <2 ug liter1 during untreated 

conditions and increased to as much as 50 ug liter1 

during blooms. In years following the Anabaena 

blooms, hypolimnetic nitrate concentrations were 

significantly higher in the treated basin of the lake 

than they were in an untreated basin or than they 

were before the blooms. Development of nitrogenfixing cyanobacteria blooms is dependent on both a 

low N:P supply ratio and a sufficient phosphorus 

supply 

Sukhanova, I. N., M. V. Flint, G. Hilbaum, V. 

Karamfilov, A. I. Kopylov, E. Matveea, T. N. 

Rat'kova, and A. F. Sazhin (1988) Exuviaella 

cordata red tide in Bulgarian coastal 

waters (May to June 1986). Marine Biology 

99: 1-8. 

{Exuviaella; red tide; bloom; Burgas Bay; 

phytoplankton; plankton; species composition) 

The structure and some functional characteristics 

of the plankton community at the times of a 

Exuviaella cordata red tide were investigated in 

Burgas Bay, Bulgaria, in May and June 1986. 

Characteristics of main plankton components 

(phyto-, bacterio-, nanoheterotrophic plankton, 

ciliates and mesoplankton) in the bloom area are 

presented. Development of the E.cordata bloom was 

determined by abiotic conditions among which 

eutrophication and salinity decrease caused the patch 

character of its rapid development. Attainment of 

maximum red tide (ca. 1 x lo9 cells 1-l; 1 x lo3 g 

m-3) from background (500 to 800 x lo3 cells 1-l; 
600 g m-3) took 3 to 7d. Growth rate (u) during 

that period was 1.2 to 2.2 doublings per day. A 

sharp bloom decline (3 to 4d) was caused by 

parasitic flagellates destroying the alga's 

chloroplast. Die1 biomass losses due to grazing 

remained below 5%. Metabolites and degradation 

products of Exordata revealed no pronounced toxic 

effects on the other components of the planktonic 

community. The rapid bloom degradation due to 

effects of parasitic flagellates indicates the high 

potential ecosystem "self-regulation". 

Sullivan, B. K. and P. V. Banzon (1990) Food 

limitation and benthic regulation of 

population of the copepod Acartia 

hudsonica Pinhey in nutrient-limited and 

nutrient-enriched systems. Limnology and 

Oceanography 35: 16 18- 163 1. 

{secondary production; copepods) 

Evidence that population size of copepods in 

coastal marine ecosystems can be both resource and 

predator limited was obtained from experimental 

enclosures with and without the sediment 

community in place. The novel finding was that 

meroplankton and attached benthos were apparently 

effective competitors and predators of planktonic 

copepods in both nutrient-limited and nutrientenriched systems. During the spring period of 

increase Acartia hudsonica, as well as other 

abundant species of copepods, reached considerably 

higher numbers in four mesocosms without 

sediments than in four other mesocosms with the 

normal sediment community in place regardless of 

rate of nutrient input to the mesocosms. 

Measurements of population parameters of A. 

hudsonica indicated equivalent rates of reproduction, 

growth, and recruitment in nutrient enriched 

treatments, with and without sediments. 

Suppression of population growth in the presence 

of sediments was therefore not due to food 

limitation but, by inference, to higher rates of 

mortality associated with high abundances of 

potential predators of benthic origin. In unenriched 

control mesocosms, resource competition with the 

benthos may also have contributed to low 

population densities in tanks with sediments 

(reproductive rates were more food limited in 

enclosures with sediment). These results 

demonstrate important interactions between benthic 

and pelagic fauna that have often been overlooked 

but nevertheless may be important in regulating 

dynamics of pelagic copepod populations in 

shallow, well-mixed waters. 

Sullivan, B. K., P. H. Doering, C. A. Oviatt, A. 

A. Keller, and J. B. Frithsen (1991) Interactions 

with the benthos alter pelagic food web 

structure in coastal waters. Canadian 

Journal of Fisheries and Aquatic Science 48:2276- 

2284. 

{benthos; food web; pelagic ecosystem) 

Results from studies in experimental enclosures 

containing both water column and benthic 

communities show that the benthos has an 

important effect on the structure and productivity of 

pelagic food webs whether that change is nutrient 

enriched or nutrient limited. Research over a 10-yr 

period in 13m-3 mesocosms showed that changes 

in the pelagic food web were correlated with 

different sediment communities and with the 

presence or absence of a benthos. The abundance of 

copepods was inversely correlated with numbers of 

macrofauna. At both low and high nutrient levels, 

systems without benthos had greatly enhanced 

numbers of carnivorous holozooplankton including 

ctenophore, medusans, chaetognaths, and fish. Our 

observations indicate that the presence of the 

benthos shortens the pelagic food web and inhibits 

the response of pelagic fauna to nutrient enrichment 

in well-mixed coastal waters. The strength of 

benthic-pelagic coupling, which is controlled by 

the amount of turbulence in the water column, may 

be more important to food web structure than the 

rate of nutrient supply and could determine which 

subsystem responds to eutrophication. 

Sullivan, B. K. and P. J. Ritacco (1985) 

Ammonia toxicity to larval copepods in 

eutrophic marine ecosystems: A 

comparison of results from bioassays and 

enclosed experimental ecosystems. Aquatic 

Toxicology 7:2O5-217. 

{ammonia toxicity; copepods; eutrophic marine 

ecosystems; morta1ity;MERL) 

In an experiment designed to simulate 

eutrophication of a shallow coastal ecosystem, 

nutrients were added to experimental ecosystems 

(MERL mesocosms) in six different treatment 

levels. We observed large reductions in the numbers 

of normally dominant copepods of the species 

Acartic tonsa and A. hudsonica associated with a 

high concentrations of unionized ammonia (NH3) 

in the two most nutrient enriched treatments. 

Comparison of 48h LC50 values of 10-15 uM1- 

INH~ obtained from laboratory bioassays with 

concentrations of NH3 associated with increased 

mortality in the MERL tanks indicated that 

bioassay data correctly predicted trends of high and 

low mortality as well as fluctuations in the 

numbers of copepods in MERL tanks. Actual 

mortality rates of the mesocosm copepods was 

sometimes higher than predicted, however. 

Sullivan, B. K. and P. J. Ritacco (1988) Effects 

of nutrients and copper on copepod 

population dynamics: A mesocosm 

study. In: M. S. Evans (ed.) Toxic 

Contaminants and Ecosystem Health; A Great 

Lakes Focus. John Wiley & Sons, Inc., New York. 
{copepod; biomass; copper; survival; egg 

production; MERL) 

No abstract 

Swartz, R. C., F. A. Cole, D. W. Schults, and W. 

A. DeBen (1986) Ecological changes in the 

Southern California Bight near a large 

sewage outfall: benthic conditions in 

1980 and 1983. Marine Ecology Progress 

Series 31:l-13. 

{sewage; species richness; biomass; density; 

macrobenthos; California) 

The structure of the macrobenthic community, 

sediment toxicity, and sediment contamination 

changed greatly between 1980 and 1983 along a 

pollution gradient from the Los Angeles County 

Sanitation Districts (LACSD) sewage outfalls on 

the Palos Verdes Shelf, California, USA to a 

reference site in the northern Santa Monica Bay. 

Stimulation of the species richness, biomass, and ' 

density of the benthos along the 60 m contour 5 to 

11 km from the outfalls was significantly reduced 

in 1983 when compared to 1980. The clam 

Parvilucina tenuisculpta and the polychaete Tharyx 

sp. A were the numerically dominant species in the 

area of benthic stimulation in both years. Stations 

1 to 3 krn from the outfalls in 1980 were occupied 

by a depauperate fauna strongly dominated by the 

opportunistic polychaete Capitella spp. In 1983 

species richness and biomass significantly increased 

close to the outfalls, Capitella spp. was much less 

abundant, and the assemblage was dominated by 

ostracods of the genus Euphilomenedes that are 

characteristic of background conditions or areas 

only slightly affected by wastewater discharges. At 

the reference site in Santa Monica Bay there were 

no significant changes in species richness, 

biomass, or density between 1980 and 1983, and 

the brittlestar Amphiodia urtica remained the 

dominant species. Sediment contamination by most 

measured chemicals and parameters of organic 

enrichment decreased on the Palos Verdes Shelf, but 

not at the reference station, between 1980 and 

1983. Sediments collected within 3 km of the 

outfalls in 1980 were acutely toxic to the 

phoxocephalid amphipod Rhepooxynnius abronius. 

There was no significant toxicity associated with 

any sediment from the shelf in 1983. These 

ecological changes correspond with the reduction in 

the mass emission of BOD and chemical 

contaminants from the LACSD outfalls between 

1980 and 1983. Severe storms in the winter of 

1982-1983 may also have contributed to 

improvements in sediment quality and the partial 

recovery of the macrobenthos. 

Takahashi, M. and N. Fukazawa (1982) A 

mechanism of "Red-Tide" formation. 11. 

Effect of selective nutrient stimulation 

on the growth of different phytoplankton 

species in natural water. Marine Biology 

70:267-273. 

{red tide; nutrient; phytoplankton; growth; 

abundance; macronutrient; micronutrient; 

chlorophyll a) 

Three semi-continuous culture experiments were 

conducted by using natural algal populations 

[dominants: Skeletonema costatum, Thalassiosira 

sp., Gymnodinium sp., Heterosigma sp. (formerly 

called Olisthodiscus luteus) and Eutreptilla sp.] 

collected from Tanigawa Harbor, Japan, where redtides occurred occasionally. The effects of macroand micro-nutrients on the growth of different 

species of algae were specially evaluated by a 

dialysis bag culture technique. Two types of 

responses for nutrients were clearly observed: 

macro- and micro-nutrient dependency. The former 

group includes S. costatum, Thalassiosira sp. and 

Eutreptilla sp. which mainly stimulated their 

growth by macro-nutrients. Heterosigma sp. was 

the latter group which was stimulated by micronutrients. Gymnodinium sp. showed an 

intermediate type. A red-tide of Heterosigma sp. 

formed during the experiment in the harbor and is 

discussed with the results of the present culture 

experiments and some field observations. 

Takano, H. (1987) Red-tides at the mouth 

of Sumida River, Tokyo, during the last 

eleven years, 1976-1986. In: T. Okaichi, 

D. M. Anderson and T. Nemoto (ed.) Red Tides: 

Biology, Environmental Science, and Toxicology. 

Elsevier, New York. 

{phytoplankton; abundance) 

The quantities of chlorophyll a, including 

pheopigments, in the brackish water of Sumida 

River were measured at a fixed station near the 

laboratory at least once in ten days from January 

1976 to December 1986. Values higher than 30 

ugll were obtained mainly from the middle of April 

to the end of September, and in February and 

October in some years. The highest value was 356 

ugll at the end of July 1977. The main growth 

areas of plankton were in Tokyo Harbour. 

Populations were swept to the sampling station by 

tidal currents. 

Tamminen, T. (1982) Effects of ammonium 

effluents on planktonic primary 

production and decomposition in a 

coastal brackish water environment. I. 

Nutrient balance of the water body and 

effluent tests. Netherlands Journal of Sea 

Research 15:455-464. 

{ammonia; primary production; decomposition; 

nutrients; archipelago; Baltic; effluent) 

No abstract 

Tamminen, T. (1982) Effects of ammonium 

effluents on planktonic . primary 
production and decomposition in a 

coastal brackish water environment. 11. 

Interrelations between abiotic and biotic 

components of the planktonic ecosystem. 

Netherlands Journal of Sea Research 15:349-361. 

{ammonia; phosphate; effluent; phytoplankton; 

primary production; decomposition; nutrients; 

archipelago; Baltic; abiotic) 

No abstract 

Tamminen, T. (1984) Ammonium effluent 

tests with phytoplankton and 

bacterioplankton communities and with a 

test alga. Ecological Bulletins 36: 120- 124. 

{ammonia; phytoplankton; bacterioplankton; 

primary production; effluent) 

Stimulative and toxic effects of industrial 

effluents containing high concentrations of 

ammonium were assayed in natural brackish water 

communities of phytoplankton and 

bacterioplankton and with laboratory cultures of a 

test alga (Chlorella sp.). Acute effluent effects on 

planktonic communities were measured with shortterm primary productivity and heterotrophic activity 

assays. Effects on Chlorella were determined as 

biomass increase during 14 d. 

Activity of bacterioplankton was inhibited by the 

lowest effluent additions (corresponding 

approximately to 100 ug NH4-N 1- I). 

Phytoplankton was stimulated over a wide range of 

effluent additions (5 to 1000 ug NH4-N 1-I), and 

toxicity was observed with effluent additions over 

1000 ug NH4-N 1-l. Chlorella showed the largest 

stimulation (up to 1000% 'of the control), and 

inhibition occurred only with the highest effluent 

additions (over 6000 ug NH4-N 1-I). The different 

responses of these organisms to effluent additions 

are discussed with regard to applying effluent tests 

to environmental monitoring and research. A 

coupling of test results and field data is considered 

to be a special advantage if the pollutant under 

study is significant in the cycles of matter in the 

environment. 

Tamminen, T., S. Kaitala, K. Kivi, and J. 

Kuparinen (1985) Response of a planktonic 

brackish water community to single and 

combined additions of ammonium and 

phosphate in a factorial mesocosm 

experiment. In: J. S. Gray and M. E. 

Christiansen (ed.) Marine Biology of Polar Regions 

and Effects of Stress on Marine Organisms. John 

Weily & Son Ltd., New York. 

{ammonia; phosphate; mesocosm; Finland; 

phytoplankton; primary productivity; model) 

Effects of single and combined addition of 

ammonium (38 and 75 mg N m-3) and phosphate 

(10 and 20 mg P -3) on the structure and function 

of enclosed (220 1) planktonic communities were 

examined during a 12-day 32 factorial experiment 

on the southern coast of Finland, in early August, 

at the commencement of the annual blue-green 

algal bloom in the area. Particulate and excreted 

primary productivity, heterotrophic activity (3~- 

glucose turnover rate), 33~04 assimilation, 14cmethylamine (ammonium analogue) assimilation 

and chlorophyll a were measured on four dates 

(activity parameters in >3 um and <3 um size 

fractions). Nitrogen fixation (acetylene reduction 

method) was measured at the end of the experiment. 

Phytoplankton responded rapidly to the nutrient 

addition, with up to a 3-fold increase of chlorophyll 

a on the first date, when the single effect of 

ammonium was the most significant. Ammonium 

was the main limiting nutrient for phytoplankton 

production during the experiment, but also 

significant phosphate effects and combined effects 

of ammonium and phosphate were observed. 

Nitrogen-fixing blue-green algae were only 

phosphate-limited. The interaction of nitrogen and 

phosphorus cycles in the regulation of primary 

productivity demonstrates that the concept of the 

limiting nutrient cannot be rigorously applied at 

the community level. Hetertrophic activity was 

stimulated by both nutrients, and the results 

emphasized the functional role of <3 um 

picoplankton in both nutrient and carbon cycles. 

Mean percentages of the total particulate 

assimilation were 44% (C02), 69% (PO4) and 68% 

(methylamine) in the < 3um fraction during the 

experiment (n=45-54). The result suggest that 

bacterioplankton would directly affect primary 

production through competition for inorganic 

nutrients. 

Tamminen, T. (1990) Eutrophication and the 

Baltic Sea: Studies on phytoplankton, 

bacterioplankton, and pelagic nutrient 

cycles. Dissertation for Ph.D., University of 

Helsinki. 

{Baltic; phytoplankton; bacterioplankton; 

nutrients; cycles; bloom; nitrogen; phosphorus; 

Finland) 

Planktonic processes were studied at the SW coast 

of Finland, the Baltic Sea, in order to examine 

causal connections relevant to eutrophication in the 

ecosystem. The annual pattern of nutrient 

limitation was investigated by means of 

experiments on several system levels, and by 

analyzing in situ nutrient ratios. Nitrogen was 

shown to be the basic limiting nutrient throughout 

the growth seasons, and colimitation by 

phosphorus occurred in early summer. 

The shift from new production of the spring 

bloom to regenerated production of the summer 

season was studied in terms of nitrogenous 

nutrition. Clear preference succession from nitrate 
to ammonium was demonstrated, but nutrient 

availability overruled inherent preferences in 

nutrient assimilation. During regenerated 

production, utilization of organic nutrients was 

(organic N) uptake, while bacterial regeneration of 

organic P provided an important P sources also for 

phytoplankton. 

Research strategies for the dynamically fluctuating 

pelagic environment were outlined with special 

emphasis on the interrelation between fluctuations 

in the physical environment and within the 

planktonic food web at different time scales. 

Continuous interaction between experimental food 

web fluctuations, and system modelling appears 

inevitable in order to understand nutrient cycle 

phenomena in the special conditions of the Baltic 

Sea. 

Nitrogen discharge was shown to promote 

eutrophication in the Baltic Sea. Effective reduction 

of nitrogen loading is therefore of vital importance. 

Taslakian, M. J. and J. T. Hardy (1976) Sewage 

nutrient enrichment and phytoplankton 

ecology along the central coast of 

Lebanon. Marine Biology 38:3 15-325. 

{sewage; phytoplankton; Lebanon; nutrients; 

abundance; diversity; dominance; blue-green algae; 

dinoflagellates; diatom; indicator) 

The abundance and taxonomic diversity of 

phytoplankton has been studied in relation to 

sewage pollution (proximity to outfalls) south of 

Beirut, Lebanon. Surface-water samples were 

collected from a series of beach stations extending 

from the American University of Beirut to 20 km 

south from June, 1973 to July, 1974. Samples 

were preserved, concentrated by settling, and the 

concentration of each taxon of phytoplankton 

enumerated in an inverted microscope. Water 

samples from the vicinity of two major sewer 

outfalls (Carton and Khalde sewers) showed very 

high concentrations of NH~+, NO2-, NO3- and 

PO 4-3, a greater total concentration of 

phytoplankton, and a lower taxonomic diversity 

than samples remote from outfalls. A considerable 

variation in the occurrence of species and 

dominance occurred along the pollution gradient. 

Blue-green algae and dinoflagellates were dominant 

in polluted waters, while diatoms dominated in 

cleaner water away from major sewage outflow. 

From the dominance and relative distribution of the 

taxa along the pollution gradient, certain taxa 

(Oscillatoria spp., Spirulina spp., Phormidium 

spp., Synechococcus custos and S. elongatus, 

Gymnodinium spp., and Prorocentrum spp.) 

emerge as indicator species of pollution. These 

changes correspond to a typical degradation of a 

complex community to a less mature state by the 

inflow of nutrient-rich sewage (eutrophication) 

along a coastal region about 10 km long. 

Thompson, G. B. and J. Ho (1981) Some 

effects of sewage discharge upon 

phytoplankton in Hong Kong. Marine 

Pollution Bulletin 12: 168- 173. 

{sewage; phytoplankton; abundance; assemblage; 

diversity; Hong Kong) 

The coastal waters of Hong Kong constitute a 

transition from estuarine conditions in the west to 

more oceanic conditions in the east, with a major 

discharge of untreated sewage located at the midpoint. Chlorophyll a was determined and net 

phytoplankton was sampled at 45 stations 

throughout this transition. Over a period of 20 

months, Chlorophyll a values rarely exceeded 2 ug 

1-I in unpolluted coastal waters. Estuarine waters 

generally contained 2-6 ug 1-I and, in waters 

influenced by sewage discharge, values sometimes 

exceeded 20 ug 1-l. There was no evidence of a 

reduction in taxonomic diversity in polluted areas 

except in summer, when the net phytoplankton was 

dominated by Chaetoceros spp. In the autumn and 

early winter, Skeletonema costatum was abundant 

in the central polluted areas. 

Tolmazin, D. (1985) Changing coastal 

oceanography of the Baltic Sea. I. 

Northwestern Shelf. Progress in 

Oceanography l5:2 17-276. 

{Black Sea; environment; nutrient; nitrogen; 

oxygen; hypoxia; anoxia; mortality; benthos) 

This article describes the hydrography of the 

Northwestern Shelf (NWS), of the Black Sea 

emphasizing the changes induced by water 

management in the Dnieper and Dniester river 

basins. The existing literature and previously 

unpublished data have been reviewed and 

synthesized to describe water property fields and 

transport mechanisms of NWS and the Dnieper and 

Dniester estuaries before the early 1960s, or the socalled precontrol period, when the effect of artificial 

river flow control upon the coastal waters was 

insignificant. 

After the hydroenergy complexes and water 

withdrawal and disposal systems on the river 

became fully operational in the early 1970s (the socalled postcontrol period), the annual river discharge 

from the Dnieper and Dniester rivers had noticeably 

decreased and seasonal river flow patterns had been 

artificially modified. Instead of a powerful and short 

early spring flood, typical for the natural conditions 

in the Dnieper river, the hydrographs in the post 

control period exhibit two smaller peaks of river 

discharge of much longer period. One of them 

(winter-early spring) is caused by intense 

hydroenergy generation and weir discharge through 

the cascade of storage reservoirs. Another is 

associated with the spring flood, modified by 

intense water consumption and storage in this 

period. High average river discharge in late May- 
early June strengthened the summer pycnocline 

which inhibits vertical mixing in the estuaries and 

coastal waters. Owing to a slow summer 

circulation, the rate of natural purification of the 

entire coastal system has been reduced. This 

coupled with the increased nutrient, organic and 

pollutant transports, decreased the dissolved-oxygen 

concentration and led to anoxic events and mass 

mortalities of marine organisms in the previously 

productive regions. These effects have primarily 

plagued the benthic communities along the entire 

western coast of the NWS since 1973. 

Winter convective overturn in the Black Sea 

reaches its maximum depth at the southern 

boundary of the NWS. Thus, the NWS waters 

descend beneath the seasonal and main thermoclines 

in the open sea and are spread by the prevailing 

currents across the entire sea in the cold 

intermediate layer (CIL). By the dynamic 

mechanism the projected man-made modifications 

in the riverine-estuarine systems of the NWS will 

affect and change the large-scale thermocline 

structure and marine life of the Black Sea. 

Tomascik, T. (1990) Growth rates of two 

morphotypes of Montastrea annularis 

along a eutrophication gradient, 

Barbados, W.I. Marine Pollution Bulletin 

21:376-381. 

{coral; Barbados; growth rate; reef; growth form; 

morphotype) 

Growth rates (skeletal linear extension) of 

columnar and lobate morphotypes of the reefbuilding coral Montastrea annularis were compared 

within and among seven fringing reefs along a 

previously described eutrophication gradient on the 

west coast of Barbados using X-ray radiography. 

Both morphotypes of M. annularis exhibit similar 

patterns of increasing average growth rates with 

improving environmental conditions. However, the 

average growth rates of the columnar morphotype 

are statistically higher (P<0.001) on all reefs in the 

study, when compared to the lobate morphotype. It 

is suggested that while environmental conditions 

exert a measurable effect on the growth rates of 

both morphotypes, the differences in the average 

growth rates between the two growth forms, under 

similar environmental conditions, may be related 

to genetic conditions. 

Index master chronologies computed for a 

columnar morphotype of M. annularis indicate a 

general pattern of decreasing growth rates within 

the past 30 yr at each of the seven fringing reefs. It 

is postulated that this general pattern of decreasing 

growth rates may be directly related to the 

deterioration of water quality along the west coast 

of the island. 

Tomascik, T. (1991) Settlement patterns of 

Caribbean scleractinian corals on 

artificial substrate along a eutrophication 

gradient, Barbados, West Indies. Marine 

Ecology Progress Series 77:26 1-269. 

{corals; Barbados; settlement; reef) 

Artificial substrate settlement plates (terracotta 

tiles) were set out on 3 fringing reefs for a period of 

12 mo to study settlement patterns of juvenile 

scleractinian corals along a eutrophication gradient 

on the west coast of Barbados, W.I. A total of 716 

coral planulae settled on 120 experimental plates 

after 12 mo of exposure. Statistically higher 

(p<0.05) average number of juvenile corals per 

plate (X = 9.2+- 3.3; N = 40) was recorded on a 

less eutrophic reef compared to 2 more eutrophic 

reefs (X = 6.9 +- 3.1; N = 40 and X = 1.9 +- 1.3; 

N = 40). Differences in juvenile coral settlement 

between reef zones, within each reef, were 

dependent on the reefs position along the 

eutrophication gradient. Statistically higher number 

of coral planulae (X = 7.2 +- 4.5; N = 60) 

successfully settled on vertical plates compared to 

horizontal plates (X = 4.8 +- 3.3; N = 60). Coral 

planulae did not settle on upper surfaces of 

horizontal plates. In terms of relative abundance, 

the most common coral species in the juvenile 

population on the experimental plates were Poritea 

asteroides Lamarck which accounted for 42% of the 

settled planulae, followed by Agaricia spp. (23%); 

and P. asteroides (Pallas) (19%). Juveniles of 

Montastrea annularis (Ellis & Solander), 

Siderastrea spp. and Diploria spp., while present at 

2 northern reefs, were absent from the most 

eutrophic reef. 

Tomascik, T. and F. Sander (1985) Effects of 

eutrophication on reef-building corals. I. 

Growth rate of the reef-building coral 

Montastrea annularis. Marine Biology 

87: 143-155. 

{coral; reef; growth; zooxanthellae; SPM) 

Fourteen environmental variables were monitored 

at seven locations along the west coast of Barbados 

on a weekly basis over a one-year period, 1981 to 

1982. The physicochemical and biological data 

indicate that an environmental gradient exists as a 

result of increased eutrophication of coastal waters. 

Growth rates (linear extension) of Montastrea 

annularis (Ellis and Solander), measured along the 

environmental gradient, exhibit high correlation 

with a number of water quality variables. 

Concentration of suspended particulate matter is the 

best univariate estimator of M. annularis skeletal 

extension rates (r2 = 0.79, P<0.0001). The results 

suggest that suspended particulate matter may be an 

energy source for reef corals, increasing growth up 

to a certain maximum concentration. After this, 

reduction of growth occurs due to smothering, 

reduced light levels and reduced zooxanthellae 

photosynthesis. 
Tomascik, T. and F. Sander (1987) Effects of 

eutrophication on reef-building corals. 11. 

Structure of scleractinian coral 

communities on fringing reefs, Barbados, 

West Indies. Marine Biology 94:53-75. 

{corals; community; reef; species composition; 

zonation; diversity; sedimentation) 

Seven fringing reef complexes were chosen along 

the leeward coast (west) of Barbados to study the 

effects of eutrophication processes upon the 

scleractinian coral assemblages. The structure of 

scleractinian coral communities was studied along a 

eutrophication gradient with a quantitative 

sampling method (line transect) in terms of species 

composition, zonation and diversity patterns. On 

the basis of these data the fringing reefs were 

divided into three ecological zones: back reef, reef 

flat, and spur and groove. Statistically discernible 

and biologically significant differences in 

scleractinian coral community structure, benthic 

algal cover and Diadema antillarum Philippi 

densities were recorded among the seven fringing 

reefs. High correlations between environmental 

variables and biotic patterns indicate that the effects 

of eutrophication processes (nutrient enrichment, 

sedimentation, turbidity, toxicity and bacterial 

activity) were directly and/or indirectly affecting the 

community structure of scleractinian coral 

assemblages. In general, species diversity was most 

sensitive in delineating among-reef, and amongzone, differences, which were attributed to 

intensification of eutrophication processes. Porites 

asteroides Lamarck, P. porites (Pallas), Siderastrea 

radians (Pallas), and Agaricia agaricites (Linnaeus) 

were the most abundant coral species in the 

polluted southern reefs. The absence and/or low 

abundance of coral species previously characterized 

as well adapted to high turbidity and sedimentation 

[i.e. Montastrea cavernosa Linnaeus, Meandrina 

meandirtes (Linnaeus)] indicate that eutrophication 

processes may adversely affect these species. It is 

suggested that sediment rejection abilities, 

combined with feeding and reproductive strategies, 

are the primary biological processes of scleractinian 

corals through which eutrophication processes 

directly and/or indirectly affect the structure of coral 

communities. 

Tomascik, T. and F. Sander (1987) Effects of 

eutrophication on reef-building' corals. 

111. Reproduction of the reef-building 

coral Porites porites. Marine Biology 94:77- 

94. 

{reef; corals; reproduction; larval development; 

maturation; zooxanthellae; gonad index) 

The sexual reproduction of Porites porites 

(Pallas), a shallow water hermatype coral, was 

studied over a one-year period (June, 1982 to June 

1983) on three fringing reef complexes lying along 

an eutrophication gradient on the west coast of 

Barbados, West Indies. The data suggest that P. 

porites is a gonochoric species with a brooding 

mode of reproduction, but a low incidence (2.7%) 

of hermaphroditism was detected in a population 

sampled from a reef subjected to urban and 

industrial pollution. Gonadal development occurs 

within the mesenteries between the retractor 

muscles and the mesenterial filaments. 

Gametogenesis occurs during nine to ten months of 

the year, with the peak reproductive activity 

occurring predominantly in the fall and winter 

(November to January). Gametogenesis was 

therefore loosely synchronized between colonies; 

however, gonad in all stages of development were 

present within colonies throughout the reproductive 

season. The reproductive season of two P. porites 

populations sampled from two polluted reefs began 

one to two months earlier than that of a P. porites 

population sampled from a less polluted reef. The 

simultaneous presence of ova and larvae within a 

colony between November and April suggests that 

larvae may be released repeatedly during an extended 

breeding season. No correlation was found between 

the average number of gonads and polyp size. 

However, the gonad index (average number of 

gonads based on the sum of male and female 

gonads) showed an inverse relationship with a 

number of environmental variables. It is suggested 

that zooxanthellae photosynthesis through reduced 

light levels may significantly lower the energy 

available from photosynthates to the maturing ova 

and/or embryos, thus depressing larval development 

and maturation. Coral colonies samples from two 

polluted reefs contained lower numbers of larvae 

than colonies sampled from a less polluted reef. 

The 2:l sex ratio observed in a P. porites 

population sampled from a polluted reef may result 

from rapid asexual reproduction (fragmentation), 

indicating that the mode of reproduction may be 

influenced by environmental conditions. 

Topping, G. (1976) Sewage and the sea. 

In: R. Johnston (ed.) Marine Pollution. Academic 

Press, London, England. 

{sewage; nutrients; benthos; review; oxygen; 

phytoplankton} 

No abstract 

Tracey, G. A. (1988) Effects of 

eutrophication on growth and 

bioenergetics of the blue mussel, 

Mytilus edulis. Dissertation for Ph.D., 

University of Rhode Island. 

{mussel; growth; energetics; physiology; toxicity; 

Narragansett Bay; MERL; sewage; phytoplankton; 

food) 

Eutrophication may be an important environmental 

problems in marine water receiving excess 

inorganic or organic nutrients from urban or 

agricultural activity. Nutrient-induced changes may 
include both altered nutritional and toxicological 

environments. Shifts in food quality may occur by 

the mixture of sewage sludge with natural 

particulates of shifts in phytoplankton species 

composition to food organisms of lower nutritional 

value. In addition, the toxicity of environmental 

conditionsmay be adjusted by environmental 

conditions created by eutrophication. These aspects 

of eutrophication were examined in this research. 

Growth and bioenergetics of the blue mussel, 

Mytilus edulis, was used to indicate ecosystem 

alteration caused by eutrophication. Bioenergetics 

measurements included the physiological responses 

of clearance rate, assimilation efficiency, respiration 

rate and ammonia excretion. Apparatus used in the 

development of the research included the use of 

experimental niarine ecosystems (mesocosms). 

Mesocosms were amended daily with varying 

amounts and sources of nutrients, and mussels were 

exposed to the mesocosm waters. Additional 

research involved examination of effects on mussels 

caused by a dense algal bloom in Narragansett Bay. 

A portion of one study involved documentation of 

mortality in mussels along a transect in 

Narragansett Bay. 

Mussels are sensitive to copper and toxicity is 

strongly dependent upon environmental conditions 

characteristic of a particular eutrophic state. Factors 

which influence the eutrophication-toxicity 

relationship include the rate of biological removal 

of Cu from the water column and the partitioning 

of Cu between dissolved and particulate phases. 

Sewage sludge does not serve as a potential food 

resource for mussels, and acts to dilute out 

available food. Food availability appears largely 

dependent upon plant biomass as indicated by 

chlorophyll a concentration. Mussels may be 

extremely sensitive to shifts in phytoplankton 

species composition, especially during the 

reproductive phase of the species. A picoplanktonic 

algal bloom caused reduced feeding in mussels 

when present in sufficient densities. 

The results of the research indicate that 

eutrophication may profoundly effect the growth 

and physiological responses of mussels. 

Mechanisms of effects include alteration of food 

quality and modification of pollutant availability. 

Of all physiological measurements, clearance rates 

were generally most sensitive. 

Tsutsumi, H., S. Fukunaga, N. Fujita, and M. 

Sumida (1990) Relationship between 

growth of Capitella sp. and organic 

enrichment of the sediment. Ma r i n e 

Ecology Progress Series 63: 157- 162. 

{Capitella; sediments; organic matter; growth; 

algae; opportunistic) 

Laboratory colonies of Capitella sp., which is 

found densely distributed in organically enriched or 

polluted areas, were cultured in sediments with 

various levels of organic matter to examine the 

relationship between level of sediment organic 

matter and growth of individuals. Growth was 

independent of absolute levels of sediment organic 

matter, but was significantly correlated with 

increases in the levels of sediment organic matter 

on addition of algal powder to the sediment. These 

results indicate that Capitalla species predominating 

in organically enriched areas have a physiological 

requirement for organic materials discharged from 

sources of organic enrichment, e.g. abundant 

growth of algae, fish farming, industrial effluent, 

sewage, if they are to grow normally. Although 

association of some Capitella species with 

sediment organic enrichment has previously been 

recognized as a reflection of their opportunistic 

characteristics, a physiological requirement for 

organic materials discharged from sources of 

organic enrichment may be the single factor most 

responsible for the concentration of Capitalla 

species in organically enriched areas. However, it is 

unlikely that they can directly ingest and assimilate 

such organic materials. Since the additional organic 

materials are easily decomposed in the sediment, 

Capitella species may require either (1) some 

specific microorganisms, the levels of which 

increase with levels of the organic materials, or (2) 

certain substance(s) produced during decomposition 

of the organic materials. 

Tsutsumi, H., T. Kikuchi, M. Tanaka, K. Imasaka, 

and M. Miyazaki (1991) Benthic faunal 

succession in a cove organically polluted 

by fish farming. Marine Pollution Bulletin 

23:233-238. 

{benthic; fauna; succession; abundance; 

community) 

In the past two decades, fish farming using net 

cages has developed in the coastal water throughout 

Japan. Such fish farming has allowed the 

production of large amounts of valuable fish and 

their supply to the markets in major cities on a 

regular basis. However, fish farming is often 

followed by serious organic pollution of the water 

and bottom sediments in the vicinity of the cages 

since approximately 90% of the food for the fishes 

results in organic discharge to the environment 

around the fish farm. Organic pollution of soft 

bottom sediment is apt to be accompanied by the 

development of reducing conditions in the sediment 

and deoxidization of the bottom water, as a results 

of the decomposition of abundant organic matter. 

The benthic- communities in the organically 

polluted areas are subjected to catastrophic 

environmental disturbances. We have assessed the 

environmental conditions and abundance of benthic 

communities in a specific cove since 1966. The 

results of the present study clearly show the 

dramatic changes in fauna and the reduction in the 

abundance of members of the benthic communities 
in the cove that have accompanied the progress of 

the organic pollution associated with fish farming. 

Tubbs, C. R. and J. M. Tubbs (1983) 

Macroalgal mats in Langstone Harbour, 

Hampshire, England. Marine Pollution 

Bulletin 14: 148-149. 

{green algae; seaweeds; sewage; bird) 

The spread of green algae over mudflats in 

Langstone Harbour has been caused by increased 

discharges of both treated and untreated sewage 

effluent. The decline in some species of estuarine 

birds may be a secondary effect of the blanketing of 

the muds by algae, but Soulsby et al., 1982 

concluded that the evidence for a causal link was 

inconclusive. This note examines the points they 

put forward in support of this. 

Turner, R. E. and N. N. Rabalais (1991) 

Changes in Mississippi River water 

quality in this century. BioScience 41:140- 

147. 

{nutrients, nutrient ratios) 

No abstract 

Turner, R. E. and N. N. Rabalais (1994) 

Coastal eutrophication near the 

Mississippi River delta. Nature 368:6 19- 

621. 

{nutrients) 

Changes in delivery of river-borne nutrients such 

as dissolved phosphate, nitrate and silicate, owing 

to land-use changes and anthropogenic emissions, 

are known to result in eutrophication-enhanced 

phytoplankton blooms-and more severe hypoxic 

events in many enclosed bays and seas. Although 

similar ecological effects might be expected on 

continental shelves, the occurrence of such 

eutrophication has been unresolved. Here we 

present evidence of eutrophication of the 

continental shelf near the outflow of the 

Mississippi River, obtained by quantifying 

biologically bound silica (BSi) in diatom remnants 

within dated sediment cores. BSi accumulation rates 

are greatest in water depths of 20 to 50 m within 

100 km of the river mouth, and have increased by 

as much as 100% this century. The increases were 

substantial by 1980, by which time riverine 

nitrogen loading had doubled relative to the 

beginning of the century, even though silica 

loading had declined by 50% over the same period. 

Thus changes in river-borne nutrient loadings can 

modify coastal coastal food webs and affect the 

amount and distribution of oxygen in bottom water 

on the scale of continental shelves. 

Twilley, R. R., W. M. Kemp, K. W. Staver, J. C. 

Stevenson, and W. R. Boynton (1985) Nutrient 

enrichment of estuarine submerged 

vascular plant communities. 1. Algal 

growth and effects on production of 

plants and associated communities. 

Marine Ecology Progress Series 23: 179- 19 1. 

{submerged plants; community; growth; estuary; 

nutrients; estuarine ponds; epiphytes; biomass; 

light; mechanism) 

Eight experimental ponds containing submerged 

vascular plants (predominantly Potamogeton 

perfoliatus and Ruppia maritima) were subjected in 

duplicate to 4 levels (including controls) of 

fertilization from June to August 1981. Seston and 

phytoplankton chlorophyll a increased with 

fertilization, and pronounced algal blooms were 

evident under high dosage. Of the total seston, 

phytoplankton exerted the greatest influence on 

attenuation of photosynthetically active radiation 

(PAR), such that there was insufficient light for 

submerged vasculat plant growth at the sediment 

surface during blooms. An extensive epiphytic 

community developed on plants in all nutrienttreated vonds at densities similar to those observed 

in nature on senescent plants. At high nutrient 

treatments the accumulation of epiphytic material 

resulted in >80% attenuation of the incident 

radiation at the leaf surface. Biomass of submerged 

macrophytes decreased significantly under high and 

medium nutrient treatments compared to control 

and low treatments within 60d following initial 

fertilization. Apparent production of vascular plants 

(based on oxygen production and 14c-bicarbonate 

uptake) was reduced at the higher nutrient 

treatments for both R. maritima and P. pelfoliatus. 

Most of this reduction in macrophyte 

photosynthesis could be explained by attenuation of 

PAR associated with epiphytic material. However, 

without PAR attenuance in the overlying water, 

observed levels of epiphytic growth would be 

insufficient to reduce light below compensation 

levels needed to sustain vascular plant growth. At 

the high fertilization rates, integrated primary 

production of pond communities was significantly 

reduced with the loss of vascular plants, even 

though phytoplankton and epiphytic growth were 

enhanced. 

Tyson, R. V. and T. H. Pearson (ed.) (1991) 

Modern and Ancient Continental Shelf 

Anoxia. Special Publication No. 58 of 

The Geological Society. The Geolocical 

Society, London. 

No abstract 

Ursin, E. and K. P. Andersen (1978) A model 

of the biological effects of 

eutrophication in the North Sea. Rappots - - 

et Proc2s-verbaux des Rkunions 

Counseil International pour L'Exploration de la 

Mer 172:366-377. 

{model; North Sea; nutrients; yield; fish) 
In a model of the pathways of phosphorus 

through the North Sea ecosystem, first, a steady 

supply of nutrients from rivers etc. was simulated, 

and next, an increasing supply. With apparently 

realistic data for water exchange and nutrient supply 

the model predicted a slight increase in the yield of 

fish, far from sufficient to describe the observed 

yield increase over the period 1960-1970. 

Valente, R. M., D. C. Rhoads, J. D. Germano, and 

V. J. Cabelli (1992) Mapping of benthic 

enrichment patterns in Narragansett Bay, 

Rhode Island. Estuaries 15:l-17. 

{sediments; redox potential; remote sensing; 

benthos; methods; stage; infauna; assemblage; 

bacteria; habitat; review; Narragansett Bay) 

A synoptic reconnaissance survey was performed 

over a five-day period in August 1988 to assess 

benthic habitat quality throughout Narragansett 

Bay, Rhode Island, using REMOTS@ analysis 

sediment-profile photography and analysis in 

combination with measurements of the levels of 

Clostridiurn pelfringens spores (a fecal indicator) in 

sediments. Three main areas of degraded benthic 

habitat quality related to either excessive organic 

enrichment or physical disturbance were identified 

based solely on the REMOTS@ analysis: the 

Providence River Reach, Greenwich Bay and its 

associated coves and harbors, and an area located 

along the southwest side of Prudence Island. 

Sediments at many stations in these areas exhibited 

shallow apparent redox-potential discontinuity 

(RPD) depths, high apparent oxygen demand, and 

low-order benthic successional stages. Elevated 

Clostridiurn perfringens spore counts in surface 

sediments were attributed to inputs from wastewater 

treatment facilities. The highest spore counts 

occurred at the head of the bay, where wastewater 

treatment discharges and associated combined sewer 

overflows are numerous. Using data from the 

REMOTS@ analysis and the sediment inventory of 

C. perfringens spores a distinction was made 

between organic enrichment of the bottom from 

sewage, versus nonsewage enrichment or physical 

disturbance. The combination of techniques 

employed in this investigation could be used to 

design more efficient monitoring to assess 

eutrophication effects in estuaries and determine the 

effectiveness of regulatory or management 

initiatives to reduce organic overenrichment of 

benthic habitats. 

Van Es, F. B., V. A. M.A., L. A. Bouwman, and 

H. G. J. Schroder (1980) Influence of organic 

pollution on bacterial, macrobenthic and 

meiobenthic populations in intertidal 

flats of the Dollard. Netherlands Journal of 

Sea Research 14:288-304. 

{oxygen; bacteria; benthos; biomass; diversity; 

Ems estuary; tidal flat; macrofauna; meiofauna} 

Large amounts of organic waste-water are 

discharged into the Ems-Dollard estuary, mainly in 

autumn and early winter. In November 1975 and 

June 1975 a number of samples were taken from 

the sediment of the tidal flats and analyzed for 

bacterial numbers, organic carbon and mud content. 

Cores were taken to quantify macro- and meiofauna. 

Oxygen saturation was measured in the water 

column over the sediment. 

In November the oxygen saturation was less than 

20% in a considerable part of the Dollard. It is not 

clear to whatextent the inward increase in organic 

matter content of the sediment was influenced by 

the waste-water discharge. 

The correlation between the viable counts of 

aerobic heterotrophic bacteria and the organic matter 

content, as observed in the other parts of the 

estuary, was absent in the most polluted part of the 

Dollard. However, the differences between the 

numbers counted in November and June were not 

significant in that area. 

Numbers of sulphate-reducing bacteria decreased 

sharply between November and June, indicating a 

dependency on the waste-water discharges. Numbers 

of sulphide-oxidizing bacteria decreased in summer 

concomitantly with the sulphate-reducing bacteria. 

These data suggest that, in contrast to the other 

parts of the estuary, in the Dollard obligate 

autotrophs play a predominant part in the 

populations of sulphide-oxidizing bacteria during 

the time of high waste discharges. 

Population diversities of meio- and macrofauna of 

the tidal flats were strongly decreased in the vicinity 

of the outfall. Total biomass of macrofauna 

decreased to zero, whereas meiofauna biomass 

significantly increased. Some possible explanations 

for the effects on the benthic fauna are given. 

Veer, H. W. v. d. (1989) Eutrophication and 

mussel culture in the western Dutch 

Wadden Sea: Impact on the benthic 

ecosystem; A hypothesis. Helgolander 

Meeresuntersuchungen 435 17-527. 

{mussel; Dutch Wadden Sea; benthos; nutrient; 

macrofauna; biomass) 

Since 1950, two large-scale changed have taken 

place in the western Dutch Wadden Sea, namely the 

eutrophication of the area and introduction of an 

extensive mussel culture. Although eutrophication 

in the fresh waters started already around 1950, 

nutrient concentrations in the western Wadden Sea 

remained fairly constant until about 1970, due to 

the retention of nutrients in Lake IJssel, the main 

source. From 1970-1980 concentrations increased 

strongly, and during the last years the situation has 

stabilized. Mussel culture was introduced in 1950 

and expanded during the next decade to an area of 70 

km2, all situated in the sublittoral area. From 1960 

the area of mussel culture remained about constant 

with fluctuating yields of between 3.5 and 120 
millions of kg fresh weight. Due to a lack of data 

for the period until 1970 the impact of 

eutrophication and mussel culture on the 

ecosystem, cannot be assessed. From 1970 onwards 

an increased biomass and production of the 

macrofauna in the intertidal zone has been observed, 

which is attributed to eutrophication. The 

hypothesis is postulated that the introduction of 

mussel culture between 1950 and 1960 has resulted 

in an increased food competition in the area, leading 

to a decreased stock of the macrofauna in the 

intertidal. Eutrophication from about 1970 onwards 

has improved the food conditions and as a result 

both in-macrofauna in the intertidal and the mussel 

in the sublittoral area would have increased in 

biomass, allowing higher maximum yields of the 

mussel culture. The importance of monitoring 

programs is stressed to follow these trends in the 

near future and to check the above hypothesis in 

areas where it is decided to introduce or intensify 

mussel culture. 

Veer, H. W. v. d., W. v. Raaphorst, and M. J. N. 

Bergman (1989) Eutrophication of the 

Dutch Wadden Sea: External nutrient 

loadings of the Marsdiep and Vliestroom 

basin. Helgolander Meeresuntersuchungen 

43:SOl-515. 

{Dutch Wadden Sea; nutrients; nitrogen; 

phosphorus) 

The increasing P and N content in the two main 

tidal basins in the western Dutch Wadden Sea, the 

Marsdiep and the Vliestroom basin, has been 

reconstructed from the 50s onwards. The area is 

enriched with nutrients by two sources both 

originating from the river Rhine, one being the 

discharge from Lake IJssel and the other the 

exchange with the coastal zone of the North Sea. 

Due to a buffering by Lake IJssel for about 15-20 

years, the eutrophication of the western Wadden Sea 

showed a time lag compared with the continuously 

increasing nutrient concentrations in the river 

Rhine and the coastal zone of the North Sea. At 

present, the primary production in part of the area 

still seems to be nutrient limited in summer, while 

loadings have already been decreasing in recent 

years. So far, no severe, negative effects on the 

ecosystem have been reported. Some remarks are 

made on the eutrophication in other parts of the 

Dutch Wadden Sea in relation to the hydrographic 

characteristics of these areas. 

Vidakovi'c, J. (1983) The influence of raw 

domestic sewage on density and 

distribution of meiofauna. Marine Pollution 

Bulletin 14:84-88. 

{sewage; meiofauna; abundance; density; Adriatic 

Sea) 

The influence of raw domestic sewage on density 

and distribution of meiofauna in the sea bottom 

sediments in the Northern Adriatic Sea, in the area 

of Rovinj (Yugoslavia), was investigated, 

comparing meiofaunal density at the stations which 

are under direct influence of sewage and the stations 

which are far from the source of pollution. It is 

concluded that raw domestic sewage does not have a 

negative influence on the density and distribution of 

meiofauna, but Parker's index of pollution indicates 

that meiofauna at these stations were under stress. 

Vogt, H. and W. Schramm (1991) 

Conspicuous decline of Fucus in Kiel 

Bay (Western Baltic): What are the 

causes? Marine Ecology Progress Series 69: 189- 

194. 

{seaweeds; Baltic; Fucus; model; biomass) 

In the framework of investigations on possible 

effects of eutrophication on the macrophytobenthos 

in Kiel Bay (Western Baltic), a large-scale survey of 

the distribution and occurrence of the genus Fucus 

was carried out in 1987188. For large-scale 

quantitative mapping, underwater television was 

employed or direct observations from the water 

surface were made. Species composition and 

quantitative biomass data were obtained by dredging 

and SCUBA-diving. Comparison with the results 

of earlier investigations revealed a drastic decline in 

Fucus biomass from between 40 000 and 45 000 t 

wet wt down to only 2400 t wet wt in 1987188, 

which means a decrease by 94 to 95 %. Whereas 

Fucus vesiculosus as well as F. serriatus were still 

frequent at depths below 2 m down to 13 m in the 

seventies, during our investigation Fucus spp. was 

not found in water depths greater than 2 m. 

Possible causes for the observed changes are 

discussed, and it is concluded that decreased light 

levels or increased epiphyte growth as a result of 

eutrophication, and the reduction of substrate for 

algal growth due to stone fishing and sand 

deposition, are the main causes for the decline of 

Fucus spp. 

Vollenweider, R. A., A. Rinaldi, and G. Montanari 

( 1992) Eutrophication, structure and 

dynamics of a marine coastal system: 

results of ten-year monitoring along the 

Emilia-Romagna coast (Northwest 

Adriatic Sea). In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{oxygen, phosphorus, Adriatic Sea) 

The process of eutrophication, which has been in 

progress for some two decades in the coastal area of 

the Northwest Adriatic Sea, shows mainly cyclicseasonal features. In winter and spring, large 

blooms of diatoms usually occur detenning the socalled "dirty waters", while in summer blooms of 

dinoflagellates cause "red tides". Low salinity and 

high nutrient loads from the River Po and most 

minor rivers favour the production of these blooms. 
Recurrent episodes of anoxia in bottom waters 

cause fish kills and other nuisances that impair 

fisheries. Algal blooms over the years, and two 

episodes of mucilage formation during the summers 

of 1988 and 1989 have become a serious menace to 

the regional tourist industry. 

Vukadin, H. (1991) Impacts of nutrient 

enrichment and their relation to the algal 

bloom in the Adriatic Sea. Marine Pollution 

Bulletin 23: 145-148. 

{nutrients; algal bloom; sewage; stratification) 

Extraordinary manifestations of eutrophication in 

the Adriatic Sea during the last few years have been 

due to the combined effects of different physicochemical and meteorological factors. Permanent 

inputs of nutrients, particularly in the northern 

Adriatic via river runoffs and municipal sewage 

during calm summers, cause marked stratification 

of the water column and reduction of the horizontal 

advection. These two effects provide the ideal 

conditions for single species bloom. An attempt 

has been made to calculate the nutrient balance 

which allows a better interpretation of algal blooms 

in the Adriatic. 

Vukadin, I. and V. Hulji'c (1981) Nutrient 

salts in the surface microlayer and 

subsurface layer of Kastela Bay waters. 

Thalassia Jugoslavica 17: 89093. 

{nutrients; nitrogen; phosphorus; microlayer; 

subsurface; Kastela Bay) 

This paper gives values of the inorganic form of 

the main nutrient salts in the surface and subsurface 

layers of Kastela Bay waters (the Middle Adriatic). 

Analyses of all nutrients dissolved show an 

enrichment in nutrients in the surface layer if 

compared with the subsurface water. Only silicon 

in the surface layer showed no significant 

enhancement with respect to silicon in the 

subsruface water. 

Vuorinen, I. and E. Ranta (1988) Can signs of 

eutrophication be found in the 

mesozooplankton of Seili Archipelago 

Sea? Kieler Meeresforschungen, Sonderheft 

6: 126-140. 

{mesozooplankton; Archipelago Sea; hydrography; 

composition) 

In this study, we evaluate whether it is possible 

to distinguish the effects of eutrophication from 

other, i.e. hydrographical, factors affecting 

zooplankton. We illustrate our arguments with 

examples from an old set of zooplankton data 

which includes 9-year records of mesozooplankton 

and hydrography collected from Seili, off the south 

coast of Finland. The present study shows that 

hydrographic changes override or mask, at least the 

time period studied, possible eutrophication effects. 

Vuorinene, I. and E. Ranta (1987) Dynamics of 

marine meso-zooplankton at Seili, 

Northern Baltic Sea, in 1967-1975. 

Ophelia 28:3 1-48. 

{zooplankton; Baltic; Kelatella) 

Long-term (1967-1975) dynamics of mesozooplankton population densities at Seili (Northern 

Baltic Sea) are described. The data were obtained 

from quantitative net hauls taken at 10 day internals 

during the open water season and once a month in 

winter. Water salinity and temperature were recorded 

concurrently. Altogether 34 taxa (species, species 

groups, instars) were frequent enough for time 

series analysis. 

The most abundant rotifers were Synchaeta 

species (S. baltica and S. monopus for the most 

part). Among cladocerans Podon polyphemoides 

and Eubosmia longispina maritima were 

numerically dominant, while Acartia spp. and 

Eurytemora affinis hirundoides were the most 

abundant copepods. The nauplii of Balanus 

improvisus were the most frequent and abundant 

meroplankton. 

In winter meso-zooplankton biomass is dominated 

by copepods (nauplii), in spring rotifers increase, 

followed by cladocerans and then by copepod 

copepodids and adults in autumn. From 20 to 100% 

of the total meso-zooplankton biomass is 

attributable to copepods. There is considerable yearto-year variation in meso-zooplankton 

composition. For example, biomass in the 

summers of 1968 and 1974 was dominated (40- 

60%) by rotifers, while in 1967, 1970 and 1973 

rotifers were less important (10-15%) in the 

biomass. 

The salinity at Seili rose substantially during 

1970- 197 1, followed by another inflow of saline 

water in 1973. These salt water intrusions kept the 

salinity at an elevated level to the end of the 

sampling period. The meso-zooplankton taxa 

responded to the ~alinity change: 11 out of 34 taxa 

increased in numbers (seasonal effect removed), 

while most (23) of the taxa decreased in numbers. 

However, in most cases these changes were not 

distinguishable from random fluctuations. Presently 

it is difficult to assess the significance of other 

factors (human-caused eutrophication, or biotic 

interactions) to the documented abundance changes. 

Waern, M. and S. Pekkari (1973) Outflow 

studies. Nutrients and their influence on 

the algae in the Stockholm Archipelago 

during 1970. No. 1. Oikos Supplymentum 

15:155-163. 

{nutrient; nitrogen; phosphorus; Baltic) 

When comparing 1970 with other years in order 

to study the effects of different treatments of sewage 

water in the Stockholm Archipelago some strong 

influences depending upon climatic causes have to 

be taken into consideration such as, 
1) the accumulation of nutrients inherited from the 

years before, 

2) the discharge below the lasting ice cover, 

3) the vigorous spring flood, and 

4) the extremely small discharge from Lake 

Malaren during the summer. 

The years 1968-1970 had 3) and 4) in common. 

In the Baltic proper an upwelling ~f phosphaterich water appeared in 1970. In the Aland Sea the 

N:P ratio varied between 11 : 1 and 19: 1 when total 

N and total P are compared. When the mineralized 

fractions were compared (below the photosynthetic 

layer) the N:P ratio varied between 2.1: 1 and 2.6: 1. 

This is far from an N:P ratio by assimilation of 7: 1 

(all ratios on a ug basis). 

Walker, D. I. and R. F. G. Ormond (1982) Coral 

death from sewage and phosphate 

pollution at Aqaba, Red Sea. Marine 

Pollution Bulletin 13:21-25. 

{sewage; coral; Red Sea; algal growth; sediment} 

Localized pollution of coral reef areas is occurring 

at Aqaba, Red Sea, as a result of sewage discharge, 

and as a result of spillage of phosphate dust during 

loading of phosphate mineral onto ships. The rate 

of death of colonies of the coral Stylophora 

pistillata was found to be 4-5 times as great in the 

polluted area as in a control area. Coral damage in 

the control area is generally caused by grazing or by 

extreme low tide, but the cause of coral death in the 

polluted area was not readily apparent. The growth 

of algae, both on damaged corals , and on glass 

slides placed out in the reef, was greatly stimulated 

in the polluted area, but it appeared that such algal 

growth was not the direct cause of coral death. 

Corals in the polluted area may be under stress 

because of reduced light intensity, inhibition of 

calcification by excess phosphate, and increased 

sediment load. It was found that in the polluted area 

there was a greater weight of sediment settling on 

the glass slides for a given weight of algae. But in 

addition, since algal growth was faster in the 

polluted area, the sediment load was increased by 

the sediment trapping capacity of the enhanced algal 

growth. Thus, it is suggested that increased algal 

growth stimulated by increased nutrient 

concentrations may be important in greatly 

increasing the sediment load experienced by corals. 

Wallstrom, K. (1988) The occurrence of 

Aphanizomenon flos-aquae 

(Cyanpphyceae) in a nutrient gradient in 

the Baltic. Kieler Meeresforschungen, 

Sonderhej? 6:2 10-220. 

{blooms; blue-green algae; NIP ratio; nitrogen; 

phosphorus} 

The investigation area, Himmerfjardan Bay 

situated at the Swedish coast of the northern Baltic 

Proper, receives waste water from a sewage 

treatment plant. Phosphorus is efficiently reduced 

in the sewage treatment resulting in a high N : P 

ratio (= 65 : 1) in the discharge. In the near future 

also the nitrogen discharge will be reduced, and it is 

feared that a lowered N : P ratio in the receiving 

waters may favour an undesired increase of nitrogen 

fixing blue-green algae. This study is focused on 

the development of a common nitrogen fixing 

species in the area, Aphanizomenon flos-aquae, at 

varied nutrient loadings in 1983-85. The biomass 

as well as the number of heterocysts was low near 

the treatment plant and increased with increasing 

distance from the sewage discharge. The low 

biomass and number of heterocysts in 

Himmerfjardan Bay is interpreted as an effect of 

competition with other algae and of low level of 

phosphorus relative to nitrogen, i.e. unfavourable 

conditions for nitrogen fixation. The abundance of 

Aphanizomenon was highest in 1984 when the 

phosphorus load was higher than the other years of 

investigation. As a tentative conclusion it is 

suggested that nitrogen fixing algae may increase as 

a result of reduction in the nitrogen discharge to 

Himmerfjikdan Bay in the future. At the reference 

station outside the Bay, nutrient conditions 

favoured nitrogen fixing algae, as interpreted from 

inorganic N:P ratio less than 10:l during the 

summers of the investigation period. Here the 

variation in biomass between the years was mainly 

due to the variation in water temperature; 

Aphanizomenon was most abundant during the 

warm summer of 1984 and less abundant during the 

cold summer of 1985. A significant correlation was 

obtained between the heterocyst frequency and 

concentration of inorganic phosphorus in the 

trophogenic layer at the reference station. 

Walsh, J. J., G. T. Rowe, R. L. Iverson, and C. P. 

McRoy (1981) Biological export of shelf 

carbon is a sink of the global C02 cycle. 

Nature 291: 196-201. 

{carbon; cycle; nutrients; shelf; C02} 

Measurements of carbon metabolism, production 

and exchange along food webs suggest that large 

fractions of the organic matter produced on 

continental shelves must be exported to continental 

slopes. The annual loss of organic matter from 

continental shelf ecosystems is far greater than in 

the open ocean. If part of the loss of nearshore 

primary production has increased in those coastal 

zones where anthropogenic inorganic nutrient 

supplies have been consistently increasing since the 

industrial revolution, then burial and diagenesis of 

this material in slope depocentres could represent 

the 'missing BMTs of the carbon' in global C02 

budgets. 

Wang, Z. (1987) Evaluation of water 

quality in the Zhujiang Estuary, China. 

In: T. Okaichi, D. M. Anderson and T. Nemoto 
(ed.) Red Tides: Biology, Environmental Sciences, 

and Toxicology. Elsevier, New York. 

{phytoplankton; abundance; red tide) 

This paper presents preliminary results of a 

monthly comprehensive investigation of 

eutrophication and red tides in the Zhujiang (Pearl 

River) estuarine area, beginning in February, 1987. 

The purpose of this paper is to discuss the trophic 

level of water quality in the Zhujiang estuary 

during Spring and Summer. A single parameter as 

well as a multiparameter Trophic State Index was 

adopted to assess the trophic status in the estuarine 

area. Using nutrients (DIN and DIP), COD loading, 

and changes in phytoplankton (cell counts and 

chlorophyll a) as the major parameters, a synoptic 

assessment was made. 

Weaver, A. M. (1978) Aspects of the effects 

of particulate matter on the ecology of 

kelp forest (Macrocystis pyrifera) (L.) 

C.A. Agardh) near a small domestic 

sewer outjall. Dissertation for Ph.D., Stanford 

University. 

{particulate; sewage; Ulva ; sedimentation; kelp} 

An investigation of the relation of particulate 

matter to the ecology of a kelp forest was 

undertaken near a small sewer outfall located on a 

flat shale reef in 11 m of water which supported a 

canopy of the kelp Macrocystis pyrijera. The 

sewage effluent had an unusually high particulate 

content which varied several fold through time. The 

area immediately in front of the outfall terminus, as 

compared to adjacent stations, was conspicuously 

more turbid, had higher sedimentation and many 

patches of exposed shale devoid of organisms, 

including Macrocystis. 

The purpose of the study was fourfold: 1) to 

observe how the quality and quantity of particulate 

matter fluctuated through time and space; 2) to 

determine some of the major reasons for these 

fluctuations; 3) to determine the influence of the 

quantity and quality of particulate matter on the 

distribution of benthic organisms; and 4) to 

determine if and how the sewer outfall influenced 

particulate matter in the study area. 

Particulates were trapped over time in plastic 

tubes mounted vertically on the bottom at stations 

located within and outside of the sewage field as 

defined by coliform levels. The contents were 

subjected to various analyses. 

Total sedimentation (grams of particulate matter 

trapped per unit time per unit area) fluctuated 

through time in a seasonal cyclic manner, increased 

in the fall, was at its maximum in the winter and 

decreased again in the spring. This pattern was 

observed at 11 different stations separated by a 

maximum distance of 1400 meters. At any one 

point in time, however, large differences (up to 20 

fold) in sedimentation were observed between 

individual stations. 

Water movement generated by onshore swells 

appeared to be a major influence on total 

sedimentation both in time and space. The amount 

and proximity of bottom sediments also played a 

major role in determining spatial differences. 

Total sedimentation in the area of the outfall was 

disrupted. Pronounced gradient in total 

sedimentation were demonstrated both laterally 

(parallel to shore) and in a seaward direction. 

However, these differences appeared to derive from 

the outfall structure and the local topography rather 

than the direct sedimentation and accumulation of 

sewage particles. 

The lack of organisms at the near sewer station 

was attributed to (1) the scouring of organisms 

from the substrate when total sedimentation was at 

a high level, (2) burial of available substrate during 

periods of high sediment accumulation. 

Like total sedimentation, macrodetritis 

(fragmented plants and animals) content of the 

trapped sediments displayed a seasonal behaviour, 

generally increasing and decreasing with total 

sedimentation. 

The fluctuations in macrodetritis levels appeared 

to be a major influence upon the levels of organic 

carbon in the particulates within the study area. 

The outfall station had consistently higher 

macrodetritis levels than adjacent stations and the 

fall macrodetritis at all station within the sewage 

filed contained large quantities of detached Ulva not 

found elsewhere. 

The occurrence of the Ulva bloom (which took 

place annually for three consecutive years) appeared 

to be based on: 1) the high water clarity during 

periods of low sedimentation which allowed 

sufficient light for Ulva growth at 11 m depth; 2) 

the presence of sewage effluent which is known to 

stimulate the growth of Ulva. 

Distinct concentration gradients of copper were 

often, but not always, demonstrable in the 

sediments trapped around the outfall. The highest 

concentrations were at the outfall station. 

During the period of high sediment accumulation 

(summer), the direct sedimentation of copper with 

particles from the sewage effluent appeared to be 

responsible for the gradient. However, during the 

fall, copper accumulating in the detached Ulva 

appeared to be responsible. 

The importance of these findings to future 

environmental surveys and the location of sewer 

outfalls is discussed. The significance of the Ulva 

phenomena as a potential pathway for toxins to get 

into the food chain is also considered. 

Weigelt, M. and H. Rumohr (1986) Effects of 

wide-range oxygen depletion on benthic 

fauna and demersal fish in Kiel Bay 

1981-1983. Meeresforschung 31:124-136. 

{abundance; biomass; benthic; macrofauna; 

oxygen; anoxia; hydrogen sulfide) 
In late summer 1981 an extraordinary and wide 

ranging oxygen depletion killed most of the benthic 

fauna below the halocline (>20m) in Kiel Bay. 

This catastrophic period lasted for several weeks, 

and even areas were affected where such events had 

never been observed before. Fishermen reported 

mass catches of moribund polychaetes and 

migrations of fishes into nearshore waters. Only a 

few macrofauna species survived (Arctica, Astarte, 

Corbula, Halicryptus). 

About 30,000 t of macrofauna died in Kiel Bay, 

but recolonization in the following months was 

rapid. Stomach analyses of demersal fish revealed 

significant changes in food composition of cod and 

dab. 

In late summer of 1983 a similar, but not so 

severe oxygen depletion caused the total breakdown 

of the Syndosmya alba stocks below the halocline 

as well as that of other species. 

Welsh, B. L. and F. C. Eller (1991) 

Mechanisms controlling summertime 

oxygen depletion in western Long Island 

Sound. Estuaries 14:265-278. 

{Long Island Sound; oxygen; mechanism; 

depletion; hypoxia) 

Physical profile data (salinity, temperature, 

oxygen, and downwelling irradiance) and in situ 

incubation of light and dark bottles were used to 

characterize vertical structure and elucidate 

mechanisms controlling summertime hypoxia in 

western Long Island Sound. The period of oxygen 

depletion corresponded with the period of thermallycontrolled stratification. Bulk density differences 

between surface and bottom waters were only 1.2 to 

2.7 sigma-t units; but they were apparently 

sufficient to resist destratification by winds and 

tides. Thus oxygen depletion was a cumulative 

process through the summer. During the 

stratification period, net oxygen production 

(measured using light BOD bottles) was confined to 

a narrow surface zone of 1.8-4.5 m. Below this 

zone was an intermediate zone of high net oxygen 

uptake, beneath which was a subpycnoclinic zone 

where oxygen uptake was very low. Rates of total 

oxygen uptake (dark bottles) were greatest in the 

surface layer and diminished with depth. There was 

close coupling between physical conditions and 

metabolic structure. Vertical structure patterns of 

oxygen production and removal were strongest in 

calm weather. The location of the intermediate zone 

corresponded with that of the oxycline. The 

thickness of the zone and the steepness of the 

oxycline were determined by the depth and intensity 

of both physical stratification and biological 

production and respiration. The biological structure 

was weakened by physical mixing in the upper 

water column, and the intermediate zone disappeared 

with fall destratification. We hypothesize that 

biological uptake within the water column 

influence oxygen depletion through two 

mechanisms. (1) In bottom waters, uptake rates per 

unit volume are low, but bulk uptake is a 

significant factor in oxygen depletion because of 

the large volumn of water involved. (2) The 

intermediate zone, where respiratory uptake is also 

significant, is strategically located between the 

surface zone of oxygen renewal and the bottom 

zone of depletion, where it constitutes an active 

filter which reinforces the pycnocline as a barrier to 

vertical oxygen dispersion. The magnitude of direct 

oxygen removal in the water column relative to 

removal by sediment oxygen demand and the 

potential effects of this biological filtering 

mechanism are important considerations for 

understanding eutrophic dynamics and managing 

Long Island Sound. Dynamic models which (1) 

underestimate the role of water column uptake and 

(2) incorporate only the two-zone characteristics of 

physical stratification will tend to (a) overestimate 

the contribution of sediments to summertime 

oxygen deficits and (b) overestimate rates of vertical 

dispersion and reventilation of bottom waters. 

Weston, D. P. (1990) Quantitative 

examination of macrobenthic community 

changes along an organic enrichment 

gradient. Marine Ecology Progress Series 

61:233-244. 

{abundance; biomass; macrobenthos; community; 

trophic; size; richness; distribution; Puget Sound) 

Organic enrichment, both natural and 

anthropogenic, is one of the most common forms 

of disturbance in the benthos. The effects of organic 

enrichment on the benthos in the vicinity of a large 

mariculture facility were examined as a general 

model for enrichment, without the confounding 

effects of toxicants often associated with 

anthropogenic inputs. Stations nearest the facility 

were subject to continuous input of fish feed and 

fecal matter, and gross structural changes in the 

macrofaunal community (e.g. reduced species 

richness, dominance of opportunistic species) were 

similar to those commonly reported for other 

enriched sites. More complex community and 

population responses were indicated by changes in 

body size, vertical distribution of infauna and 

patterns of trophic dominance. Enriched areas are 

generally assumed to be characterized by 

macrofauna with small body sizes, and mean 

individual size did, in fact, decrease with proximity 

to the farm. At stations farthest from the farm, 

however, size distributions were skewed by a few 

large individuals, indicating that trends in mean 

individual size are susceptible to the 

methodological limitations inherent in adqeuately 

sampling rare individuals. While interspecific 

measures of animal size decreased with increasing 

enrichment, intraspecific measures indicated a 

tendency for larger individuals to occur at the most 
enriched sites. Enrichdd areas may represent 

increased food resources, and thus increased 

potential growth and attainment of larger body sizes 

in those species capable of exploiting such 

habitats. Organically enriched areas are generally 

considered to be characterized by infauna living at 

or near the sediment-water interface. Increasing 

organic enrichment resulted in the loss of large, 

deep-dwelling species and dramatically altered the 

vertical biomass profiles, but, because these 

individuals were numerically few, had little effect 

on the vertical abundance profiles. Changes in 

trophic dominance did occur along the trophic 

groups and assign species to them. The utility of 

trophic grouping approaches in identification of 

enrichment-induced disturbances is thus limited. 

Whiteledge, T. E. (1985) Nationwide Review 

of Oxygen Depletion and Eutrophication 

in Estuarine and Coastal Waters. NOAA, 

{oxygen; estuary; coastal; nutrients; production; 

United States; hypoxia; depletion} 

No abstract 

Widbom, B. and R. Elmgren (1988) Response 

of benthic meiofauna to nutrient 

enrichment of experimental marine 

ecosystems. Marine Ecology Progress Series 

42:257-268. 

{benthic; abundance; composition; meiofauna; 

nematode; polychaete; harpacticoid; bivalve; 

ostracod; kinorhynch; MERL) 

The long-term (2.4 yr) response of benthic 

meiofauna to eutrophication of experimental marine 

ecosystems was studied at the Marine Ecosystems 

Research Laboratory, Graduate School of 

Oceanography, University of Rhode Island (USA). 

Ammonium, phosphate and silicate were added 

daily in the mesocosms in a logarithmic 

progression (OX, lX, 2X, 4X, 8X, 16X, and 32X), 

with the 1X addition being N = 2.88, P = 0.225, Si 

= 0.205 mmol m-2 d -I. Phytoplankton production 

and biomass in the tanks increased with increasing 

nutrient enrichment. The benthic community gave 

a quantitatively less marked response to the gradient 

of nutrient input. The meiofauna showed 

remarkably little response in terms of biomass and 

abundance, but significant effects were found on 

major taxa, leading to a changed meiofauna 

community structure. Nematode and juvenile 

polychaete abundance increased with increasing 

nutrient input, especially in early summer, whereas 

kinorhynchs, ostracods, harpacticoids and juvenile 

bivalve decreased. The lack of a positive biomass 

response of the total meiofauna in the enriched 

tanks suggests that the meifauna was limited not 

only by the availability ot food, but also by biotic 

interactions. 

Wilcox, W. H. (1979) The effect of nitrogen 

and phosphorus enrichment on salt 

marsh. Dissertation for Ph.D., The University of 

Tennessee. 

{salt marsh; biomass; dominance; periphyton; 

composition; Florida) 

Fertilization of a Batis maritima and Salicornia 

virginica dominated salt marsh in Florida, with a 

combined nitrogen and phosphorus fertilizer applied 

at four rates (50:8; 100:16; 150:24 and 200:32 

kgthdyr N:P) produced the following responses: 1) 

dominance shifted to Batis, 2) Batis aerial biomass 

increased from 428 glm2 (control) to 1062 glm2 

(highest rate) while Salicomia showed no positive 

response to fertilization, 3) highest fertilization 

plots had a 9% higher decomposition rate than 

control plots (0.00728lday versus 0.00668/day), 4) 

no luxury consumption of nutrient was measured, 

5) Batis stem biomass reached an asymptote of 750 

g/m2 (295 glm2 control) after 2 years fertilization 

at 150:24 and 200:32 kglhdyr N:P, 6) Batis 

biomass at 328 glm2 (133 glm2 control) had not 

reached an asymptote after two years fertilization. 

Measurement of initial changes in snail numbers 

and periphyton biomass and species composition, 

in response to fertilization could not be repeated, 

probably due to a drying trend in the marsh and a 

drastic reduction in snail and periphyton numbers. 

Wittberg, M. and W. Hunte (1992) Effects of 

eutrophication and sedimentation on 

juvenile corals. I. Abundance, mortality 

and community structure. Marine Biology 

112:131-138. 

{sedimentation; corals; abundance; mortality; 

community structure; reef; size} 

This study investigated effects of eutrophication 

and sedimentation on juvenile abundance, juvenile 

mortality and community structure of scleractinian 

corals on fringing reefs on the west coast of 

Barbados, West Indies, in 1989. Juvenile abundance 

was lower on eutrophiclhigh-sediment reefs than 

less eutrophicllow-sediment reefs, but juvenile size 

was larger on the former. The larger size could 

result from size-selective mortality against smaller 

juveniles on the eutrophic reefs, or from lower 

recruitment to the eutrophic reefs, or from faster 

growth on the eutrophic reefs. Juvenile mortality 

was higher on the eutrophic reefs than the less 

eutrophic reefs and may result from increased 

smothering of corals by eutrophic reefs, probably 

in response to elevated nutrients andlor because 

grazers (Diadema antillarum; herbivorous fish) were 

less common on eutrophic reefs. Juvenile 

community structure on all reefs was dominated by 

Type 1 corals (high recruitment, high natural 

mortality), but Type 2 corals (low recruitment, low 

natural mortality) became more common in adult 

communities on the less eutrophic reefs. This 

transition in community structure did not occur on 
the eutrophic reefs, adult community structure 

continuing to be dominated by Type 1 corals. The 

fact that the pattern of relative abundance of species 

in the juvenile community is maintained in the 

adult community on the eutrophic reefs suggests 

that juvenile mortality rates of different species are 

similar on eutrophic reefs, and hence that 

differences in adult community structure between 

eutrophic and less eutrophic reefs may be largely 

explained by interspecifics in juvenile mortality 

becoming smaller on eutrophic reefs. 

Wolff, W. J. (1988) Impact of pollution of 

the Wadden Sea. In: W. Salomons, B. L. 

Bayne, E. K. Duursma and U. Forstner (ed.) 

Pollution of the North Sea. An Assessment. 

Springer-Verlag, Berlin. 

{phytoplankton; production; microphytobenthos; 

Dutch Wadden Sea; biomass; macrobenthos; 

review) 

No abstract 

Wong, P. S. (1987) The occurrence and 

distributiuon of red tides in Hong Kong - 

Applications in red tide management. 

In: T. Okaichi, D. M. Anderson and T. Nemoto 

(ed.) Red Tides: Biology, Environmental Science, 

and Toxicology. Elsevier, New York. 

{red tide; bloom) 

The occurrence of red tides in Hong Kong is 

reviewed based on reports received by the 

Aquaculture and Fisheries Department. Interesting 

patterns of distribution of some common red tide 

species in Hong Kong are obtained. Data on 

seasonal occurrence and distribution of each species, 

particularly the toxic ones, would enable better 

planning of preventing measures or management 

actions in different coastal regions. Knowledge of 

toxicity of different species would facilitate the 

initial assessment of risks involved. Further 

suggest the possibility of development a system for 

forecasting red tide occurrences. 

Wu, R. S. S. (1982) Periodic defaunation 

and recovery in a subtropical epibenthic 

community, in relation to organic 

pollution. Journal of Experimental Marine 

Biology and Ecology 64253-269. 

{benthic; defaunation; Tolo Harbour; Tolo channel; 

Hong Kong; abundance; biomass; composition; 

oxygen; recovery; diversity) 

A monthly survey was carried out for two years 

on the epifaunal community in Tolo Harbour and 

Tolo Channel, Hong Kong, a subtropical 

embayment subjected to a gradient of organic 

pollution. The number of animals and species, 

biomass and species diversity (Shannon's function 

H' and eveness J), were higher at less polluted 

stations in the outer Harbour and Channel than at 

the polluted stations in the inner Harbour. The 

Channel community was dominated by the crab 

Portumus hastatoides Fabricius and the gastopod 

Nassarius crematus (Hinds). Along an increasing 

gradient of organic pollution, Nassarius crematus 

was gradually replaced by the crab Charybdis 

vadorum (Alcock), and a community dominated by 

Portumus hastatoides and Charybdis vadorum was 

found in the polluted inner Harbour. The abundance 

and dominance of predatory gastropods also showed 

a decrease from the Channel to the inner Harbour, 

reflecting changes in the trophic structure of the 

community in relation to pollution. No significant 

change in the percentage of deposit-feeders was 

found along the pollution gradient. Summer 

mortality of benthos appeared to occur regularly in 

inner Tolo Harbour, and was attributed to summer 

oxygen depletion resulting from eutrophication. 

The benthic community was, however, soon 

restored to its original state by rapid winter 

recolonization. It is postulated that the cyclic 

phenomenon of summer mortality followed by 

winter recovery may be a common characteristic in 

subtropical benthic communities subjected to a 

high level of organic pollution. 

Wulff, F. and A. Stigebrandt (1989) A timedependent budget model for nutrients in 

the Baltic Sea. Global Biogeochemical Cycles 

3:63-78. 

{nitrogen; phosphorus; nutrient; loading; model; 

budget; Baltic) 

Overall budgets for nutrient and humus are 

described for the Baltic Sea as well as for the 

subsystems, i.e., the Baltic proper, the Bothnian 

Bay and the Bothnian Sea. The residence times for 

total phosphorus, total nitrogen, silicate and humus 

are 13.3, 5.5, 11.2, and 9.6 years respectively, 

compared to 2 1.8 years for a conservative substance 

(salt). About 90% of the nutrient losses are due to 

biogeochemical sinks within the Baltic Sea. Thus 

only about 10% is exported to external areas (the 

Kattegatmelt Sea). For humus the corresponding 

figures are about 75 and 25%, respectively. This 

means that the Baltic Sea to a large extent can be 

regarded as a closed system and perturbations in the 

water exchange with the North Sea should have 

little effect on the nutrient budgets. This sinks are 

parameterized by an expression borrowed from 

limnology where the net nutrient loss is a function 

of the winter surface concentration. A budget model 

is run in a prognostic, hindcast mode with the 

assumed time-dependent phosphorus and nitrogen 

loading of the Baltic proper. The computed 

development of the winter surface concentrations of 

total P and total N for the period 1950-1988 

appears quite realistic. The possibility of having 

variable sinks which are functions of the surface 

winter concentrations of nitrogen and phosphorus is 

described using calculations based on data from the 

different Baltic subareas. Such sinks should 
significantly decrease the winter N:P ratio in the 

surface water when the nutrient loading increases 

with time. With better description of in particular 

the pools of nutrients in the sediment, it would be 

possible to model future changes of nutrient 

concentrations in the water column in relation to 

loading. 

Wulff, F., A. Stigebrandt, and L. Rahm (1990) 

Nutrient dynamics of the Baltic Sea. 

Ambio 19:126-133. 

{nitrogen; phosphorus; nutrient; Baltic) 

A prerequisite for understanding the large-scale 

eutrophication of the Baltic Sea is an understanding 

of the factors responsible for regional and long-term 

variations of nutrients. This article summarizes 

recent studies on the changes in overall total 

amounts and the distribution pattern of nutrients. 

The total amounts of phosphorus and nitrogen have 

increased over the last few decades. On the other 

hand, silicate levels have been decreasing, 

indicating a higher net primary production and 

sedimentation of diatoms. A series of models has 

been used to test our present understanding of the 

critical processes controlling nutrient and oxygen 

conditions. It is shown that a few morphometric 

factors are essential, like shallow sill depth at the 

entrance of estuarine-like sea and shallow mean 

depth. It is also shown that phosphorus, nitrogen 

and silicate differ in terms of residence times and 

the fate of these nutrients take place in the 

sediments. A model showing the long-term 

behavior of this system in response to external 

loading illustrates the importance of these 

processes. 

Wyatt, T. and J. Horwood (1973) Model which 

generates red tides. Nature 244:23 8-240. 

{red tides; model; bloom; dinoflagellates) 

No abstract 

Yang, D. B. (1987) Nutrient and 

chlorophyll a variations during the red 

tides in Jinhae Bay, Korea. In: T. Okaichi, 

D. M. Anderson and T. Nemoto (ed.) Red Tides: 

Biology, Environmental Science and Toxicology. 

Elsevier, New York. 

{red tides; nutrient; abundance; chlorophyll) 

Tidal cycle time series distributions of nutrients 

and chlorophyll a were measured at a fixed station 

located at mid-channel of Jinhae Bay, Korea. High 

nitrate concentrations were observed at the time of 

low tides whereas high phosphate concentrations 

occurred occasionally at the time of high tides. 

Anoxic bottom water appeared to be the major 

source of high phosphate in the outer bay. 

Chlorophyll a concentrations were positively 

correlated with nitrate concentrations in April 1981, 

May 1982 and June 1983. However, chlorophyll a 

concentrations were also positively correlated with 

phosphate in June and August, 1981 and September 

1982. Despite positive relationships between 

nutrients and chlorophyll a, phytoplankton growth 

is not likely to be limited by these nutrients in 

Jinhae Bay. The role of growth stimulators in 

initiating the extensive blooms of red tide 

organisms in suggested. 

Yi, S. K., J.-S. Hong, and J. H. Lee (1982) A 

study on the subtidal benthic community 

in Ulsan Bay, Korea. Bulletin of the Korea 

Ocean Research and Development Institute 4: 17- 

26. 

{benthic; Ulsan Bay; polychaete; abundance; 

composition) 

A quantitative investigation of the soft-bottom 

macrobenthic community in Ulsan Bay was carried 

out from June, 1980 to March 1981. Of the 127 

species of benthic organisms found, Polychaeta was 

the most dominant taxonomic group with 67 

species which accounted for 88% of the total - 

number of individuals. The number of species and 

individuals decreased from the inner bay toward the 

outer bay stations. At station 1, a sharp drop in the 

number of individuals was noted from June to 

September, and this drop was mainly due to 

decrease in three dominant polychaetes, Cirratulus 

cirratus, Lumbrineris longijolia, and Tharyx sp. 

Based on interstational species similarity, the 

study area can be divided into two zones; Station 

1,2,3, and Station 43. The first zone was 

considered to be under an organic enrichment 

condition. This was supported, to a certain extent, 

by hydrological conditions and species 

composition. 

Zambianchi, E., C. Calvitti, P. Cedcamore, 

F.D1Amico, E. Ferulano, and P. Lanciano (1992) 

The mucilage phenomenon in the Norther 

Adriatic Sea, summer 1989: A study 

carried out with remote sensing 

techniques. In: R. A. Vollenweider, R. 

Marchetti and R. Viviani (ed.) Marine Coastal 

Eutrophication. Elsevier, Amsterdam. 

{Adriatic Sea, gels) 

Observations are presented of the mucilage 

phenomenon which affected the Northern Adriatic 

Sea in the summer of 1989. The observations were 

made with data acquired by different spatial and 

temporal resolution satellite and airborne sensors. 

The results of our study are presented, together with 

data from other sources, and analyzed in order to 

contribute to the understanding of the spatial and 

temporal evolution of the phenomenon. 

Zdanowicz, V. S., D. F. Gadbois, and M. W. 

Newman (1986) Levels of organic and 

inorganic contaminants in sediments and 

fish tissues and prevalences of 

pathological disorders in winter flounder 
from estuaries of the Northeast United 

States, 1984. Oceans '86 578-585. 

{organic; inorganic; contaminants; sewage; 

pathology; winter flounder; disease) 

The Benthic Surveillance Project of NOAA's 

National Status and Trends Program seeks to 

determine prevalences of pathological disorders in 

demersal finfishes and levels of organic and 

inorganic contaminants in sediments and fish 

tissues and evaluate possible correlations between 

biological effects and contamination. More than 

fifty of the Nation's estuaries and other coastal 

water bodies are under study. Preliminary findings 

from the first year of sampling in the Northeast 

region of the U.S. reveal that sediments from 

Raritan Bay, Boston Harbor and Salem Harbor 

contained substantially elevated concentrations of 

primary sewage related organic and inorganic 

contaminants. PCB levels in livers of winter 

flounder exceeded 10 uglg in specimens from 

Boston Harbor, but were 3.2 uglg or less 

elsewhere. Distributions of certain pathological 

conditions, such as the presence of gigantic cells in 

the tubular epithelium of winter flounder kidney, 

paralleled the distributions of sediment 

contaminants, whereas distribution of other 

pathological conditions, such as winter flounder 

pancreatitis, did not. 

Zingone, A., M. Monitresor, and D. Marino (1990) 

Summer phytoplankton physiognomy in 

coastal waters of the Gulf of Naples. 

P.S.Z.N.I: Marine Ecology 11: 157-172. 

{phytoplankton; Gulf of Naples; nutrients; 

abundance; diversity; bloom) 

Thirteen sampling cruises were conducted at 

weekly intervals in the inner part of the Gulf of 

Naples in the summer of 1983 to investigate the 

effects of excess nutrient inputs on phytoplankton 

communities. High surface phytoplankton 

concentrations (up to 1.15 x lo8 cells 1-l) were 

recorded, particularly near Naples harbour and along 

the eastern coast, two locations that receive most of 

the area's sewage and industrial discharge. 

Phytoplankton populations were generally 

dominated by small species, mainly diatoms, which 

were associated with small phytoflagellates. 

Species diversity values were relatively high 

(H'S3.62) in most samples. Throughout the 

sampling period a high spatial and temporal 

variability for phytoplankton abundances and 

species composition was observed. 
Appendix: Keyword List 

abiotic Black Sea decomposition 

abnormal 

abnormalities 

Abra 

abundance 

acid 

acidity 

activity 

Adriatic 

Adriatic Sea 

Aegean Sea 

Alexandrium 

algae 

algal bloom 

algal growth 

alkaline phosphatase 

Ammonia toxicity 

ammonia 

ammonium 

ammoniumlnitrate ratio 

Ampelisca abdita 

amphipod 

annelids 

anoxic 

aquaculture 

aquatic vegetation 

archipelago 

archipelago sea 

Arkona 

Arkona Sea 

Asia 

assay 

assemblage 

assessment 

Atlantic menhaden 

Atlantic shelf 

atmosphere 

auhwuchs 

Aurococcus 

Australia 

autoecology 

bacteria 

bacterioplankton 

Baltic 

Barbados 

beach 

benthic 

benthic algae 

benthic communities 

benthic fauna 

benthos 

Berrow Flat 

bioassay 

biogeochemical cycling 

bioindicator 

biomass 

bird 

bivalve 

bivalves 

Blanca Bay 

bloom 

blue-green algae 

Bohai Bay 

Boston Harbor 

Brachionus 

brown tide 

brown tide bloom 

budget 

Burgas Bay 

Buzzard Bay 

Cadiz Bay 

California 

capacity 

Capitella 

carbon 

14c production 

causes 

change 

Charlotte Harbor 

Chesapeake Bay 

chironomonids 

chl a 

chlorophyll 

chlorophyll a 

Ch~ysochromulina 

ciliates 

Cladophora 

classification 

climate 

C02 

coast 

coastal 

coccolithophores 

colored tides 

community 

community structure 

comparison 

concept 

contaminants 

copepod 

copepods 

copper 

coral 

coral reef 

corals 

Corophium 

Crassostrea 

criteria 

crustacea 

culture 

cyanobacteria 

cycle 

cycles 

cycling 

cyst 

decapod 

decrease 

defaunation 

demand 

demersal 

denitrification 

density 

deoxygenation 

depletion 

deposit-feeder 

depth penetration 

desert 

development 

diatom 

diatoms 

Dictyocha 

die1 

dinoflagellate 

dinoflagellates 

Dinophysis 

disappearance 

discharge 

disease 

dissolved oxygen 

distribution 

diversity 

Dogger Bank 

DOM 

dominance 

dredging 

Dunaliella 

Dutch Wadden Sea 

dy narnics 

dystrophic 

ecosystem 

ecosystem health 

eelgrass 

effects 

effluent 

effluents 

egg 

egg production 

Eh 

Elefsis Bay 

Ems estuary 

energetics 

energy flow 

enrichment 

environment 

environmental condition 

epifauna 

epiphytes 

epiphytic 

Erie 

estuaries 

estuarine ponds 

estuary 

eutrophic 

eutrophic marine ecosystems 
eutrophicated 

eutrophication 

evidence 

excretion 

Exuviaella 

Far East 

fate 

fats 

fauna 

faunal composition 

feeding 

filamentous 

Finland 

fish 

fish farming 

fish kill 

fish yield 

fishkill 

fjord 

flagellates 

floating 

Florida 

flounder 

fluorescence 

flux 

food 

food web 

Forth estuary 

fouling 

framework 

freshwater 

Fucus 

Furcellaria 

gels 

general model 

German Bight 

gonad index 

gonads 

Gonyaulax 

Gotland Sea 

grazing 

Greece 

green algae 

green tide 

Greifswald Bay 

growth 

growth form 

growth potential 

growth rate 

Gulf of Maine 

Gulf of Naples 

Gulf of Patras 

Gymnodinium 

H2S 

habitat 

hatpacticoid 

Hawaii 

health 

Helgoland 

herring 

heterotrophic uptake 

historical 

Hong Kong 

Hudson River 

Hudson River estuary 

humic 

hydrogen sulfide 

hydrogen sulphide 

hydrographic 

hydrographic d ynarnics 

hydrography 

hydromedusae 

hypoxia 

impacts 

increase 

index 

indicator 

Indo-Pacific 

infauna 

inorganic 

inorganic nutrients 

interface 

intertidal 

Irish Sea 

iron 

Italy 

Japan 

Kanaohe Bay 

Kastela Bay 

Kattegat 

Kelatella 

kelp 

Kiel Bay 

Kiel Bight 

kinorhynch 

lagoon 

Lake Kinneret 

Lake Ontario 

lake 

lakes 

Laminaria 

larval development 

Lebanon 

length 

life cycle 

life history 

light 

limitation 

loading 

lobster 

log-normal 

Long Island 

Long Island Sound 

Louisiana 

Macoma 

macroalgae 

macroalgal mats 

macrobenthos 

macrofauna 

macronutrient 

macrophytes 

macrozoobenthos 

Madmck 

management 

mariculture 

marine 

Marsdiep 

mhsh 

mats 

Matsushima Bay 

maturation 

mechanism 

Mediomastus ambiseta 

Mediterranean 

meiobenthos 

meiofauna 

MERL 

meroplankton 

mesocosm 

mesoplankton 

mesozooplankton 

metabolism 

methane 

method 

methods 

Mexico 

microalgae 

microflora 

microheterotrophs 

microlayer 

micronutrient 

micronutrients 

microphytobenthos 

microplankton 

microzooplankton 

mining 

model 

modeling 

modelling 

mollusc 

morphotype 

mortality 

mucous aggregates 

mudflat 

mussel 

Mytilus 

NIP ratio 

N20 production 

Nahant Bay 

nanoplankton 

Narragansett Bay 

nematode 

Nephrops norvegicus 

net ecosystem metabolism 

net ecosystem production 

Netherlands 

New Jersey 

New York Bay 

New York Bight 

nitrate 
nitrification 

nitrite 

nitrogen 

Nitzrchia seriata 

Nodularia 

North Sea 

Norway 

Nucula 

Nucula annulata 

nuisance 

nuisance bloom 

nutrient 

nutrient addition 

nutrient enrichment 

nutrient limitation 

nutrient ratios 

nutrients 

oligochaetes 

opportunistic 

organic 

organic carbon 

organic enrichment 

organic matter 

Oslofjord 

ostracod 

oxygen 

oxygen demand 

oxygen uptake 

oyster 

Palmico River estuary 

paper 

particulate 

Pawilucina 

pathology 

pelagic 

pelagic biology 

pelagic ecosystem 

Peridinium 

periphyton 

PH 

Phaeocystis 

Phaeodactylum -. Philippines 

phosphorus 

photosynthesis 

photosynthetic rate 

Phycodrys 

Phyllophora 

physical energy 

physiology 

phytoplankton 

phytoplankton abundance 

phytoplankton assemblage 

phytoplankton bloom 

phytoplankton species 

composition 

picoalgae 

picoplankton 

pigments 

Pilayella 

plankton 

Po River 

pollution 

polychaete 

polychaetes 

Polydora ligni 

Pornatoschistus 

population 

Portugal 

precipitation 

prediction 

primary production 

primary productivity 

processes 

production 

productivity 

protein 

Providence River 

Puget Sound 

pulp 

Pyrodinium 

ratio 

recolonization 

recovery 

recycling 

Red Sea 

red tide 

red algae 

redox potential 

reef 

regeneration 

relationship 

remedial action 

remineralization 

remote sensing 

reproduction 

research needs 

reservior 

resource 

respiration 

review 

Rhine 

richness 

River Elbe 

rocky shore 

rotifer 

runoff 

salt marsh 

salt pond 

San Francisco Bay 

sand goby 

sandflat 

sandy beaches 

Saronikos Gulf 

Scandinavian 

Scottish coast 

Scottish water 

seagrass 

seasonality 

seaweeds 

secondary 

secondary production 

sediment 

sediment transport 

sedimentation 

sediments 

settlement 

sewage 

shading 

shelf 

shellfish 

silica 

silicate 

silicoflagellate 

silicon 

similarity 

size 

size structure 

Skagerrak 

Skeletonema 

sludge 

smelt 

spatial 

spawning 

species 

species abundance 

species composition 

species richness 

SPM 

stability 

stage 

standing crop 

standing stock 

Strait of Georgia 

strategy 

stratification 

striped bass 

submerged 

submerged plants 

submerged vegetation 

subsurface 

succession 

sulfate reduction 

survival 

susceptibility 

Sweden 

temperature 

temporal 

Thailand 

tidal flat 

tidal marsh 

tide 

tissue 

Tokyo Bay 

tolerance 

Tolo channel 

toxic 

toxic algal exudates 

toxicity 

transition 
transparency 

trend 

trophic 

trophic state index 

trophic status 

turbidity 

Ulsan Bay 

ultrastructure 

Ulva 

United States 

upwelling 

Uronerna 

Vancouver Harbor 

vascular plants 

vegetation 

Venice Lagoon 

Victoria Harbour 

Visakhapatnam Harbour 

Vollenweider 

Wadden Sea 

wastewater 

winter flounder 

yield 

Ythan estuary 

Yugoslavia 

zonation 

zooplankton 

zooxanthellae 

Zostera 

.U.S. GOVERNMENT PRINTING OFFICE 1997/432-031 
OTHER TITLES IN THE 

DECISION ANALYSIS SERIES 

No. 1. Able, Kenneth W. and Susan C. Kaiser. 1994. Synthesis of Summer Flounder Habitat 

Parameters. 

No. 2. Matthews, Geoffrey A. and Thomas J. Minello. 1994. Technology and Success in 

Restoration, Creation, and Enhancement of Spartina alternzflora Marshes in the United States. 2 

vols. 

No. 3. Collins, Elaine V., Maureen Woods, Isobel C. Sheifer and Janice Beattie. 1994. 

Bibliography of Synthesis Documents on Selected Coastal Ocean Topics.