ISSN 0097-8078, Water Resources, 2006, Vol. 33, No. 1, pp. 97–103. © Pleiades Publishing, Inc., 2006.Original Russian Text © S.V. Aleksandrov, O.A. Dmitrieva, 2006, published in Vodnye Resursy, 2006, Vol. 33, No. 1, pp. 104–110.

97

INTRODUCTION

The growing anthropogenic impact on water bodiesin the recent 50 years made it necessary to monitor theirconditions and search for objective criteria of waterquality [1]. Studying the primary production (PP), bio-mass, and species composition of phytoplankton iswidely used for the assessment of the biological pro-ductivity of water bodies and the sanitary–hygienicstate of natural waters, as well as for the developmentof methods for the prediction and control of fish pro-duction [1, 6].

Kursiu Marius Lagoon—a semiclosed freshwaterlagoon—lies in the eastern part of the southern BalticSea [18, 24]. This lagoon is the largest in the Baltic Seain terms of its area and water volume. Its main morpho-metric and hydrological characteristics are given inTable 1.

The lagoon and its watershed area lie in the regionwith developed industry, agriculture, and navigation.Now, by its hydrochemical and biological characteris-tics, Kursiu Marius Lagoon can be characterized as ahighly eutrophic water body. The decline in industrialproduction and fertilizer application in the 1990s didnot result in a substantial improvement of the environ-mental conditions. As before, high P and N concentra-tions are recorded in water. The eutrophication of thebay affects all trophic layers, first of all, the populationsof bacterio-, phyto-, and zooplankton. Considerablewater heating in summer facilitates mass developmentof blue-green algae, which has resulted in the recentyears in a “hyperblooming” of the lagoon. It is worthmentioning that six out of nine cases of hyperblooming(phytoplankton biomass > 100 g/m

3

) took place in therecent 10 years [28]. The eutrophication processes aremost pronounced in the southern (Russian) part of thelagoon, where bottom sediments are composed of silts

rich in organic matter, there is no seawater inflow, theeffect of rivers is insignificant, and, hence, waterexchange is slower.

The state of ichthyofauna in this region is as yet sta-ble. Kursiu Marius Lagoon features high fish produc-tion and belongs to the most important fishery waterbodies in the Baltic region [23], which makes it neces-sary to perform regular comprehensive biological stud-ies. The PP of phytoplankton in the Russian part of thelagoon (occupying 75% of its total area) is still poorlyknown. Observations were made only in 1974–1976[12]. In this work, we present the results of PP studies

Primary Production and Phytoplankton Characteristicsas Eutrophication Criteria of Kursiu Marios Lagoon,

the Baltic Sea

S. V. Aleksandrov and O. A. Dmitrieva

Atlantic Research Institute of Fishery and Oceanography, ul. Dm. Donskogo 5, Kaliningrad, 236000 Russia

Received October 4, 2004

Abstract

—The results of studies of phytoplankton production, biomass, and species composition conducted inKursiu Marius Lagoon in 2001, 2002 are presented. A significant effect of blue-green alga blooming on thetrophic status and processes in the coastal part of Kursiu Marius Lagoon is revealed. It is shown that the massdevelopment of algae in combination with setup phenomena can cause alga accumulation in the coastal zone,oxygen deficiency, and fish kills.

DOI:

10.1134/S0097807806010118

WATER QUALITY AND PROTECTION:ENVIRONMENTAL ASPECTS

Table 1.

Main morphometric and hydrological characteris-tics of Kursiu Marius Lagoon

Description Value Measure-ment units

Total area 1584 km

2

Russian part area 1202 The same

Volume 6.2 km

3

Mean depth 3.8 m

Maximum depth 5.8 The same

Watershed area 100485 km

2

Continental runoff 20.8 (76.6) km

3

/year (%)

Inflow from the sea through the strait

5.1 (18.4) The same

Precipitation 1.3 (5.0)

"

Discharge into the sea through the strait

26.2 (96.6)

"

Evaporation 3.4 (3.4)

"

Water exchange rate 4.2 Year

–1

Salinity 0.1 ‰

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ALEKSANDROV, DMITRIEVA

conducted in Kursiu Marius Lagoon in recent years,which enable the assessment of the present-day biolog-ical productivity and the environmental conditions inthis water body.

MATERIALS AND METHODS

Studies of planktonic PP in Kursiu Marius Lagoonwere carried out 1–2 times a month from April toNovember 2001–2002. PP and organic matter destruc-tion were determined at seven standard long-term sta-

tions of AtlantNIRO (Fig. 1). Oxygen modification ofthe light-and-dark-bottle method with a short exposure(3–5 h) was used. For the evaluation of overall PP under1 m

2

, the rate of photosynthesis was measured at fourhorizons of the photic zone. The lower boundary of thephotic zone was taken at the depth where 1% of lightthat enters water was recorded [5, 8, 30]. The sampleswere placed into an on-deck flow-through incubator,where light conditions corresponding to the samplinghorizons were simulated with the use of light filters.Organic matter destruction and chlorophyll a content

Fig. 1.

Layout of phytoplankton and PP observation stations in Kursiu Marius Lagoon.

BALTIC

SEA

LITHUANIA

RUSSIA

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PRIMARY PRODUCTION AND PHYTOPLANKTON CHARACTERISTICS 99

were measured simultaneously at two horizons of thephotic zone and in the bottom layer. The values of pro-duction obtained at short-term exposure were convertedto the per-day value with allowance made for the pho-tosynthetically active radiation measured within a day(by the ratio of the photosynthetically active radiationfor the exposure period and for the day). The photosyn-thetically active radiation was measured from the sun-rise to the sunset at a half-hour interval by using a TKA-LYUKS luxmeter (the measurement range of 400–700 nm). The daily destruction was evaluated takinginto account the exposure time. Data of the oxygenmethod were converted into carbon units with the use ofa coefficient of 0.32 mg C/mg O

2

.

Phytoplankton samples were taken at all stations in2002. For this purpose, water samples were taken fromdifferent horizons (including bottom layer) and col-lected in one vessel to form an integral sample. Thequantitative processing was performed in accordancewith the conventional procedure in a Nogotta cell [13].

The values of total PP (

Σ

A), destruction (

Σ

R), chl aconcentration, and phytoplankton biomass under thesurface area of 1 m

2

were calculated by using Surfer pro-gram, which allows the construction of distribution mapsof the indices examined over the area of the bay and thecalculation of their mean values. PP and destruction inthe ice-free period (

ΣΣ

A and

ΣΣ

R) were evaluated bygraphic integration method. The values of

ΣΣ

A and

ΣΣ

Rthus obtained were divided by the duration of the ice-freeperiod to obtain mean daily values [16].

For studying the coastal ecosystem of Kursiu Mar-ius Lagoon in the period of blooming of blue-greenalgae, six temporal observational stations were orga-nized on the research base of AtlantNIRO (KurshskayaSpit). At these stations, the concentrations of dissolvedO

2

and Chl a, pH, water temperature and transparency(by Secchi disk) were measured on the daily basis, andphytoplankton samples were taken. PP and planktonrespiration were determined periodically.

RESULTS AND DISCUSSION

Phytoplankton of Kursiu Marius Lagoon has beenstudied for more than 70 years [11, 15, 22, 28, 29]. Interms of phytoplankton species composition, thelagoon is classified as a typical freshwater body; brack-ish-water species are met only in the northern part.According to data of recent years, the community isrepresented by 438 species and intraspecies alga taxa,belonging to eight divisions. Throughout the period ofstudies, a principally invariable model of seasonal phy-toplankton succession was observed: diatom algaedominate in winter and spring, and blue-green algaedominate in summer and autumn. However, significantchanges took place in recent years in the species com-position, population, and biomass of phytoplankton.Among diatoms, the significance of p. Stephanodiscus(S. hantzschii Crun, S. agasisensis Hakanson, etc.) and

Actinocyclus normanii (Greg.) Hust., which were notfound among dominants in the 1950s [22, 28]. In differ-ent months of 2002, these species accounted for 63–97% of diatom biomass. Their appearance in waterbodies accompanies the process of eutrophication, andtheir abundance is typical of eutrophic water bodies[10, 27]. Cold-water species p. Stephanodiscus domi-nate in April and November (their biomass averagedover the lagoon reaches 4 g/m

3

of wet matter), whereasA. normanii become abundant (28–66 g/m

3

) in July–October (this species is a subdominant at the predomi-nance of blue-green algae).

Starting from June, mass development of blue-greenalgae (Aphanizomenon flos-aquae (L) Ralf, Microcys-tis seruginosa Kutz. emend. Elenk., etc.) is observed.This process later passes into water blooming. Thepotentially toxic Aph. flos-aquae species, which is per-manently present in phytoplankton during the year butrapidly increases its biomass (by 100–1000 times)when water temperature (t) reaches 20

°

C. Bloomingcommonly lasts until late October–early Novemberuntil t decreases to 7–8

°

C. The predominance of Aph.flos-aquae was recorded by all researchers; however, itsbiomass has increased in the recent decades by morethan an order of magnitude—from 34 g/m

3

in the 1930sand 12 g/m

3

in the 1950s to 120–240 g/m

3

in the mid-1990s [28]. In the period of blooming in 2002, phy-toplankton biomass averaged over the lagoon varied indifferent months from 35 to 640 g/m

3

. At the values assuch as these, secondary biological pollution takesplace in water bodies [19, 20]. A distinct peak in PP andphytoplankton biomass is observed in the period ofblooming in Kursiu Marius Lagoon; this peak coincidesin time with water heating to 20–22

°

C (Figs. 2, 3). Thisphenomenon is typical of eutrophic water bodies [21].A similar model of seasonal dynamics of PP wasalready recorded in 1974–1976 [12]. A specific featureof 2001 and 2002 was a high rate of primary productionwithin a considerable part of vegetation period. In par-ticular, in 2002, values of PP close to maximum wererecorded for three months (from July to September),which can be attributed to the hydrometeorological fea-tures of this year, e.g., high

t

(20

°

C and more) persisteduntil mid-September. Only after cooling and a temper-ature drop down to 7–8

°

C, which was recorded inNovember 2001 and October 2002, blooming of blue-green algae in the lagoon ceased and organic matter pro-duction by phytoplankton abruptly decreased (Fig. 2).

Water temperature was the factor that had the stron-gest effect on PP. The correlation coefficient between

Σ

A and t in 2001 and 2002 amounted to 0.92 and 0.85,and that between the rate of photosynthesis at the depthwith optimal light conditions (A

opt

) and t is equal to0.87 and 0.71, respectively.

Observations made in the lagoon in 2002 two timesa month revealed another feature of phytoplanktondevelopment—peak-type regime of its functioning. Atmaximum water heating in summer, periods of very

100

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ALEKSANDROV, DMITRIEVA

high phytoplankton biomass gave place to periods of itsrapid decline (the biomass decreased to 35 g/m

3

and85 mg Chl a/m

3

on the average for the lagoon), afterwhich a new considerable increase in biomass tookplace (up to 640 g/m

3

of wet matter and 700 mg Chla/m

3

). These abrupt changes in phytoplankton biomass(by almost 20 times) could take place within one week.Hypereutrophic water bodies, one of which is KursiuMarius Lagoon, feature instability at phytoplanktonpeaks. Periods of high biomass are replaced by periodsof abrupt decline because of self-shadowing and poi-soning by metabolic products [21]. The manifoldincrease in phytoplankton biomass and the rate of pho-tosynthesis (from 4 to 16 g C/(m

3

day)) had only aslight effect on the value of PP in the water column. As

the biomass increases from 35 to 640 g/m

3

, phytoplank-ton production under unit surface area increased onlyby 20% (from 4.3 to 5.6 g C/(m

2

day). This is due to thefact that the extremely high phytoplankton concentra-tion brought about self-shadowing of cells and waterturbidity decreased (from 0.55–0.6 to 0.1–03 m), thusreducing the depth of the photic zone.

Blooming of blue-green algae in the lagoon affectedthe state of its coastal zone. Coastal areas covered bymacrophytes in Kursiu Marius Lagoon serve as spawn-ing grounds and fattening areas of young and adult fish[18]. In particular, the concentration of ichthyoplankton(larvae of roach, bream, and other fish species) in Julyis several hundreds of times higher than in other partsof the lagoon [9]. Concentrations of Chl a and dissolvedO

2

recorded in the coastal part of the lagoon in theperiod of blooming are given in Fig. 4. The abundanceof blue-green algae in combination with setup phenom-ena resulted in accumulation of phytoplankton in thecoastal zone in late July–August 2002, when the con-centration of Chl a reached anomalously high values(up to 22 500 mg/m

3

) [2]. The result was that the rate oforganic matter destruction became many times largerthan the rate of phytoplankton production, whichcaused a deficiency in the concentration of dissolved O

2

(up to its complete absence) and fish kill (primarily,young), first in small bays and next in the coastal zone.This effect was local and controlled by wind directionand speed. In late July–August 2002, at steady easternwind, accumulation and decomposition of algae with aconsequent fish kill was observed at the western shoreof the lagoon. The lack of dissolved O

2

in the coastalzone (500 m from the shore) was observed within along time (up to eight days). In this period, the coastalzone was low in phytoplankton, only small amounts ofp. Microcystis (N. aeruginosa, M. wesenbergii), thebest adapted to the development under the conditions ofO

2

deficiency, were met here [19]. Wind-induced accu-mulation of algae formed hazardous conditions for

Fig. 2.

(

1

) Total phytoplankton production, (

2

) organic mat-ter destruction, and (

3

) Chl a concentration in the photiclayer of Kursiu Marius Lagoon in (a) 2001 and (b) 2002.

Fig. 3.

Phytoplankton biomass and Chl a concentration inthe water column in 2002. (1) Cyanophyta, (2) Bacillario-phyta, (3) other species, (4) Chl a concentration.

Fig. 4.

Concentrations of (

1

) O

2

and (

2

) Chl a in the coastalzone of Kursiu Marius Lagoon (500 m from the macrophytebelt) in the period of water blooming (July–August 2002).

6543210

300

200

100

0

g C

/(m

2

day)

mg

Chl

a/m

2

250

200

150

100

50

0

6

5

4

3

2

10

Apr. May June July Aug. Sept. Oct. Nov.

1 2 3

(‡)

(b)

350

300

250

200

150

100

500

Apr. May June July Aug. Sept. Oct. Nov.

100

200

300

400

0

1 2 3 4

Biomass, g/m

3

mg Chl a/m

3

16

12

8

4

030 1 3 5 7 9 11 15 17 19 21 23 25

Date

22500 1200

1000

800

600

400

200

0

g O

2

/m

3

mg Chl a/m

3

1 2

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PRIMARY PRODUCTION AND PHYTOPLANKTON CHARACTERISTICS 101

aquatic organisms inhabiting the areas overgrown bymacrophytes and small bays. Phytoplankton biomasshere reached several tens and even hundreds kg of wetmass per m

3

, as a result of which O

2

concentrationremained at the level close to zero for a long time afterthe decay of algae and aquatic organisms could not sur-vive in the conditions that formed. Similar phenomenawere observed before during water blooming inDnieper reservoirs, where wind-induced accumulationof phytoplankton in the coastal zone also brought aboutextremely high concentrations of it and hence, local fishkills [4, 19]. The prevention of this adverse phenome-non in water bodies as large as Kursiu Marius Lagoonseems unlikely now [3].

Because of the long (from July to October) bloom-ing of blue-green algae in Kursiu Marius Lagoon, veryhigh production characteristics averaged over vegetationseason were recorded here in 2001 and 2002 (Table 2).Considering that phytoplankton photosynthesis in thelagoon virtually ceases in late November and amountsto about 0.003 g C/(m

3

day) in winter [12], we canassume that the value of PP during vegetation season isequal to the annual production. The obtained values ofPP, phytoplankton biomass and Chl a concentration aretypical of water bodies in a hypereutrophic state [5, 8, 14].They are significantly higher than the production valuesrecorded in previous studies in Kursiu Marius Lagoon. In1974–1976, the mean value of

ΣΣ

Ä

was estimated at 304[12], while in 2001 and 2002, at 535–620 g C/m

2

year)(Table 2). Thus, based on trophic classifications ofwater bodies, the status of Kursiu Marius Lagoon canbe estimated as eutrophic with transition into a hyper-eutrophic phase in the periods of blue-green algablooming. Notwithstanding the “hyperblooming” ofphytoplankton, the values of daily assimilation number(DAN), averaged over the vegetation season, were sim-ilar (Table 2) and coincided with the mean value(30 mg C/mg Chl) available in literature for fresh andsalt water bodies in moderate and northern latitudes[5, 25, 26].

In addition to the high rate of organic matter produc-tion, a high rate of its mineralization was recorded inthe lagoon in all years of its studies (Table 2). The larg-est values of destruction, reflecting the consumption forthe exchange of the entire planktonic community in2001 and 2002 (as in 1974–1976) were recorded fromJune to September. As in the case of PP, temperaturewas the factor that had the largest effect on the rate ofdestruction. The correlation coefficient between t and

Σ

R

in the photic layer and in water column in 2001 wasequal to 0.77 and 0.72 and in 2002, 0.80 and 0.87,respectively.

Because of the high rate of photosynthesis, whichwas due to blue-green alga blooming, organic matterdestruction in the photic layer throughout the vegeta-tion period (except for June 2001) was much lower thanPP (Fig. 2). The ratio

ΣΣ

Ä/

ΣΣ

R

in the photic layer forthe entire vegetation season amounted to 1.2 in 1974–

1976, 1.5 in 2001, and 2.4 in 2002. The larger value ofthis ratio in the recent years as compared with 1974–1976 can be accounted for by longer blooming of blue-green algae, especially in 2002.

In 2001, organic matter destruction in water columnwas somewhat greater than the phytoplankton produc-tion (Table 2). However, these characteristics werealmost the same in 2002. The values of destructionhigher than primary production of plankton is a com-mon phenomenon, which was recorded in many high-production and medium-production reservoirs in theformer USSR, as well as in some bays in the Baltic Sea(Gdansk Bay, Pomorskaya Bay, etc.) [5, 17, 31, 32].Water bodies are open systems; therefore, destructioninvolves not only autochthonous organic matter, butthat delivered from the watershed as well. It is alsoworth mentioning that in this study we take intoaccount only phytoplankton production and do not con-sider the production of macrophytes, periphyton, andphytobenthos.

The final phase of production process in the waterbody is fish production. Fish productivity is related toPP via a series of intermediate links; however, it isdirectly dependent on the phytoplankton “yield.” In theliterature, PP is most often correlated with fish catch

Y

f

[5–7, 25, 26]. The authors compared PP for KursiuMarius Lagoon with data on

Y

f

presented by Zapbaltry-bvod. Fish catch in 2001 and 2002 amounted to 0.05%of the annual PP of phytoplankton. This relatively low

Y

f

/

ΣΣ

A

ratio as compared with data published in the lit-

Table 2.

Characteristics of the trophic status of Kursiu Mar-ius Lagoon

CharacteristicsPeriod of studies

2001 2002

Water transparency, m 0.60 (0.45–0.90) 0.55 (0.20–0.90)

Photic layer depth, m 1.55 (1.2–2.4) 1.50 (0.55–2.45)

Concentration ofChl

a

, mg/m

3

83 (25–465) 103 (32–697)

DAN, mg C/mg Chl 33 (7–57) 57 (7–57)

A

opt

, g C/(m

3

day) 2.4 (0.4–4.6) 3.0 (0.3–16)

Σ

A

, g C/(m

2

day) 2.2 (0.2–4.8) 2.0 (0.1–5.6)

Σ

R

, g C/(m

2

day) 1.5 (0.3–3.3) 0.9 (0.1–2.6)

Σ

R

*, g C/(m

2

day) 3.0 (0.9–6.0) 2.3 (0.2–11.9)

ΣΣ

A

, g C/(m

2

day) 535 620

ΣΣ

R

, g C/(m2 day) 354 263

ΣΣR*, g C/(m2 day) 785 670

ΣΣA/ΣΣR 1.5 2.4

ΣΣA/ΣΣR* 0.7 0.9

Phytoplanktonbiomass, g/m3

– 71 (10–640)

* For the entire water column.

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WATER RESOURCES Vol. 33 No. 1 2006

ALEKSANDROV, DMITRIEVA

erature (0.12–0.15% [6, 25]) can be explained as fol-lows.

Since the 1960s, Kursiu Marius Lagoon is a waterbody with controlled fishery [18]. At present, the majorportion of commercial production (20–30 kg/ha) con-sists of large long-cycle fish, such as benthos-eatingspecies (mostly bream) and predators (zander). Thetransformation of organic matter of phytoplankton intothe production of benthos-eating fish passes through arelatively long detrital food chain (involving bacteriaand zoobenthos), which is accompanied by additionallosses of the primary organic matter. The controlledfishery aimed predominantly at more valuable species(bream, zander) has resulted in a considerable (almosttwofold) decrease in the total fish catch as comparedwith 1920–1930 when the major portion of the catchconsisted of small short-cycle fish (whitebait, smelt,ruff).

In the recent years, a common phenomenon in Kur-siu Marius Lagoon was “hyperblooming” of blue-greenalgae, which almost do not reach the pasture foodchain. In particular, in 2002, when the phytoplanktonbiomass during the ice-free period averaged 71 g/m3,the fraction unavailable for filterers (mostly trichomesAph. flos-aquae) accounted for 69% (46 g/m3). There-fore, the consumption of the major part of phytoplank-ton production in the trophic chain is possible only afterits bacterial decomposition in the form of detritus witha subsequent utilization by benthos and zooplankton.That is, phytoplankton PP in hypereutrophic KursiuMarius Lagoon is utilized mostly via microbial trophicweb, thus leading to a considerable decrease in the effi-ciency of matter and energy transport. The decrease inthe efficiency of transformation of phytoplankton pro-duction toward the upper trophic levels (including fish)with an increase in the trophic status of the water bodymay be a general regularity [3].

CONCLUSIONS

The characteristics of PP, phytoplankton biomass,and Chl a content obtained in 2001 and 2002 for KursiuMarius Lagoon are typical of water bodies in hyper-eutrophic state. The values of these characteristics aremuch larger than the values of production and biomassrecorded in the 1970s. Kursiu Marius Lagoon now canbe classified as an eutrophic water body with passinginto hypereutrophic phase in the periods of blue-greenalga blooming. This conclusion is in agreement withthe estimates of the environmental state of the lagoonbased on hydrochemical and biological characteristics.

The significance of phytoplankton species, theabundance of which is characteristic of eutrophic waterbodies (p. Stephanodiscus, Actinocyclus normanii),increased many times. From June to October, poten-tially toxic species of blue-green algae Aphanizomenonflos-aqua and Mocrocystis aeruginosa rapidly developin Kursiu Marius Lagoon with a result of water bloom-

ing. Phytoplankton biomass in this period reached thelevel at which secondary biological pollution takesplace in water bodies. “Peak-type” regime of phy-toplankton community functioning appears at maxi-mum water heating in summer.

Blue-green alga blooming reflected in the state ofthe coastal zone of Kursiu Marius Lagoon. The massdevelopment of blue-green algae in combination withsetup phenomena resulted in phytoplankton accumula-tion in the coastal zone, oxygen deficiency (up to itscomplete absence), and fish kill, first of all, young. Thisphenomenon was of local character and controlled bythe direction and speed of wind.

ACKNOWLEDGMENTS

The authors are grateful to V.V. Boulion (ZoologicalInstitute, RAS) for his help in the analysis and general-ization of the obtained results and V.A. Sushin,Yu.M. Senin, V.A. Smyslov (AtlantNIRO) for theirhelp in expedition works.

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