Nutrient Elimination in an Underwater Pre-Dam

The nutrient elimination was investigated in the underwater pre-dam Haselbach (UWHA) situated in the mouth region of the Saidenbach reservoir (Saxony, Germany). The pronounced thermal stratification with the depth of the thermocline determined by the top height of the underwater dam (4,8 m below surface) resulted in epilimnetic inflow and hydraulic short circuit between inlet and main basin of the reservoir in summer. A simple model was developed for the approximation of nutrient elimination. Epilimnetic nitrate was used as a quasi-conservative discharge tracer. Compared with a “normal” pre-dam of the same size, the estimated mean phosphorus elimination of less than about 20% in UWHA was low. However, the UWHA proved to be an efficient settling basin for allochthonous particulate P.     

广东省公平水库与星湖生态特征的对比分析

在采样和测定分析的基础上,对营养盐水平和水深较为接近的公平水库和星湖的生态特征进行对比分析。结果表明,公平水库和星湖的生态特征差异显著,其主要原因是两者水动力过程的不同。公平水库水力滞留时间是星湖的1／2,水柱全年混合较好,而星湖在夏季则有明显分层现象。公平水库总氮、总磷浓度的最高值出现在丰水期的7月,枯水期明显低于丰水期,营养盐主要来源于外源．而星湖主要受内源循环的控制,总氮和总磷浓度的最高值出现在丰水期刚开始的4月,7月总氮、总磷浓度有所下降,在分层现象消失的10月,总磷浓度明显升高。公平水库水滞留时间短,水位波动较大。不利于蓝藻形成优势,其浮游植物的群落结构类型表现为硅藻-绿藻-蓝藻型,星湖水滞留时间长,水体相对稳定,为蓝藻占优势提供了条件,浮游植物群落结构类型表现为蓝藻-绿藻-硅藻型。     

Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

Influences of the lowland river Spree on phytoplankton dynamics in the flow-through Lake Müggelsee (Germany)

The influences of imports of nutrients and planktonic algae from the River Spree on the dynamics of phytoplankton were examined in the shallow, eutrophic Müggelsee, which has a retention time of only 42 days. Phytoplankton biomass and nutrient concentrations were measured in both the lake and its inflow from 1980–1990. On a long-term average, mean biomass as well as vitality of most dominant phytoplankton populations in the lake were not significantly different from those in the river. Nevertheless, during distinct periods the external rates of biomass change of single lake populations (due to dilution or enrichment) were as high as the lake internal ones. The import of inocula populations from the river probably induced the formation of the typical community structure in the lake. Growth and decay of phytoplankton populations in the river strongly influenced the load of dissolved nutrients and thus indirectly the dynamics of planktonic algae in the downstream lake. For example, intensive assimilation of phosphorus by riverine algae in spring intensified the P-shortage and supported possible P-limitation of algal growth in the lake at that time. In years with high vernal biomass of centric diatoms in the river, and thus diminished import of dissolved silicon, the growth of diatoms was suppressed and that of cyanobacteria was favoured in the lake during summer.     

Increased nutrient loading and rapid changes in phytoplankton expected with climate change in stratified South European lakes: sensitivity of lakes with different trophic state and catchment properties

We hypothesised that increasing winter affluence and summer temperatures, anticipated in southern Europe with climate change, will deteriorate the ecological status of lakes, especially in those with shorter retention time. We tested these hypotheses analysing weekly phytoplankton and chemistry data collected over 2 years of contrasting weather from two adjacent stratified lakes in North Italy, differing from each other by trophic state and water retention time. Dissolved oxygen concentrations were higher in colder hypolimnia of both lakes in the second year following the cold winter, despite the second summer was warmer and the lakes more strongly stratified. Higher loading during the rainy winter and spring increased nutrient (N, P, Si) concentrations, and a phytoplankton based trophic state index, whilst the N/P ratio decreased in both lakes. The weakened Si limitation in the second year enabled an increase of diatom biovolumes in spring in both lakes. Chlorophyll a concentration increased in the oligo-mesotrophic lake, but dropped markedly in the eutrophic lake where the series of commonly occurring cyanobacteria blooms was interrupted. The projected increase of winter precipitation in southern Europe is likely to increase the nutrient loadings to lakes and contribute to their eutrophication. The impact is proportional to the runoff/in-lake concentration ratio of nutrients rather than to the retention time, and is more pronounced in lakes with lower trophy.     

Effects of zooplankton grazing and nutrients on the bloom-forming, N2-fixing cyanobacterium Aphanizomenon in Lake Kinneret

A bloom of the filamentous, N₂-fixing cyanobacterium Aphanizomenonovalisporum Forti occurred for the first time in Lake Kinneretduring late summer and fall 1994. During subsequent years (1995–1999),Aphanizomenon also appeared in late summer and fall, but didnot bloom. In outdoor microcosm experiments, we examined zooplanktongrazing on Lake Kinneret phytoplankton, with and without Aphanizomenonpresent, and the effects of N, P and N:P ratios on phytoplanktongrowth. In one-day feeding experiments, clearance and grazingrates of the ambient Lake Kinneret zooplankton assemblage feedingin lake water dominated by Aphanizomenon were 10-fold lowerthan in water without Aphanizomenon. We suspect that the lowgrazing rates were due to interference caused by the presenceof Aphanizomenon. In 9-day nutrient addition experiments, significantenhancement effects on phytoplankton were detected with additionsof either P or N; a high N:P was better for phytoplankton growththan a low N:P. After 7 days, bottles receiving low P and noN additions were dominated by Oscillatoria sp. and Closteriumacutum; few Aphanizomenon were present. In contrast, bottlesreceiving high P and N additions had large increases of Aphanizomenon,as well as Oscillatoria and Closterium. There was a tendencyfor more green algae and diatoms with increasing N additions.These results provide evidence that (i) non-grazeability ofAphanizomenon enabled it to gain a competitive advantage overgrazeable phytoplankton, and (ii) that nutrient limitation,but not grazing, was probably important in the eventual declineof the Aphanizomenon bloom.     

Effect of temperature on rate of photosynthesis in Antarctic phytoplankton

Summary The rate of photosynthesis of marine antarctic phytoplankton (western Scotia Sea and Bransfield Strait) was determined as a function of temperature, from ambient (-0.8°C to 1.0°C) to 28°C. Photosynthetic rates, based on radiocarbon incorporation during half-day incubations, were increased by as much as 2x with temperatures up to 7°C; at higher temperatures the rates decreased rapidly, so that at 28°C the rates were only 3% of that at ambient temperatures. In antarctic surface waters during the austral summer the rate of photosynthesis by phytoplankton thus is limited by thermodynamic effects on metabolic reactions, in spite of high nutrient concentrations and saturating light levels. The observed rates were in agreement with thermodynamic models of the dependence of phytoplankton growth rate on temperature.     

SEASONAL CYCLING OF ALGAL NUTRIENT LIMITATION IN CHAUTAUQUA LAKE,NEW YORK1

Algal nutrient enrichment bioassays were conducted between May 1975 and August 1978 using water samples collected from Chautauqua Lake, New York. Photosynthetic fixation rates of natural phytoplankton assemblages were enhanced by additions of phosphorus and nitrogen, although enrichment with other nutrients had no significant stimulatory effect on algal photosynthesis. Whereas phosphorus stimulated in spring and early summer, both nitrogen and phosphorus enhanced photosynthesis in midsummer and fall. Relative to the effect of phosphorus enrichment, enhancement of photosynthesis by nitrogen during the summer and fall was highest in the northern part of the lake. During the period of ice cover, photosynthesis did not appear to be limited by nutrients in that nutrient additions (P, N, Si, C, Fe, trace metals) did not enhance fixation rates. Observed temporal fluctuations in the response of the algae to P and N correlated with changes in the lake water N:P ratio as well as with temporal changes in dissolved orthophosphate and nitrate-nitrite nitrogen. The N:P ratio decreased drastically in the summer and remained at ca. 10 or less through mid-fall, suggesting that N concentrations were inadequate for the non-N-fixing phytoplankton. Studies over 3 yr indicate that states of P and N limitation undergo time-space fluctuations that occur in a cyclic pattern in the surface waters of Chautauqua Lake.     

Phosphorus and nitrogen limitation of phytoplankton growth in eutrophic Lake Inba, Japan

Lake Inba is one of the most eutrophic lakes in Japan. In this study, field sampling and nutrient enrichment bioassays were conducted to determine the seasonal patterns of nutrient limitation for phytoplankton growth in this lake. Phytoplankton biomass increased significantly with the additions of phosphorus (P) on almost all sampling dates, indicating P limitation of phytoplankton growth from spring to autumn. However, nitrogen (N) limitation was also observed during summer (i.e., 19 August). On 10 August, a typhoon struck Lake Inba. After this event, dissolved inorganic nitrogen (DIN) and phosphorus concentrations increased, probably because of increased river discharge. At the same time, phytoplankton growth in the control treatment became relatively high, with the addition of neither P nor N stimulating the growth. However, 10 days after the typhoon, the phytoplankton growth rate in the control treatment decreased, with only the addition of N having a significant positive effect on phytoplankton growth. N limitation during summer is caused by the low concentrations of DIN, as well as changes in the N:P ratio due to allochthonous nutrient loads. These results indicate that a reduction of both P and N input is necessary to control phytoplankton blooms in Lake Inba.     

Physical factors and dissolved reactive silica affect phytoplankton community structure and dynamics in a lowland eutrophic river (Po river,Italy)

We tested the hypothesis that species composition and persistence of phytoplankton communities in nutrient rich lowland rivers depends mainly on physical factors. The study aimed to analyse the effects of water discharge, temperature and chemistry on phytoplankton dynamic and species composition in the lowland reach of the eutrophic Po river (Italy). Both taxonomical and morpho-functional methods were used. True planktonic and tychoplanktic (i.e. detached taxa of benthic origin that remain in suspension) species were found, among which only a few taxa and functional groups prevailed. Diatoms were the most abundant, with a clear dominance of species either sensitive to the onset of water stratification or well adapted to turbid waters. Phytoplankton abundance, biomass and chlorophyll-a followed similar trends, attaining the highest values in summer, at low discharge rates. Correlation and multivariate analysis revealed that the development of a stable phytoplankton community was mainly controlled by water discharge rates. Namely, changes in water flow rates induced major variations in the community structure. The seasonal succession of phytoplankton assemblages was also related to water temperature and dissolved reactive silica availability to some extent overlapping flow effects.     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。     

Growth rate of alpine phytoplankton assemblages from contrasting watersheds and N‐deposition regimes exposed to nitrogen and phosphorus enrichments

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Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Estimation of export production in the coastal Baltic Sea: effect of resuspension and microbial decomposition on sedimentation measurements

Hydrography, nutrient concentrations, primary production and sedimentation of particulate matter were studied during spring, late summer and autumn in the coastal area of the northern Baltic Sea, SW Finland. Vernal phytoplankton productivity peak and biomass maximum in early May were followed by high sedimentation rates of organic matter at the end of May. In summer, sedimentation rates of organic material were generally low. The decay rates of organic carbon in the sediment traps, estimated by measuring oxygen uptake of settled organic matter, varied between 0.005 to 0.08 d^–1 and were on average 0.02 d^–1. Decomposition of organic matter inside the sediment traps was mainly controlled by temperature, while also organic contents of settled material were significant. Microbial decomposition decreased sedimentation rates of organic carbon and nitrogen on average by 11% and 15%, respectively, during the whole study period of ca. 6 months. Resuspension of organic matter from sediment surface was estimated to contribute ca. 17 and 24% of the total sedimentation of organic carbon during spring and summer, respectively. Export production (i.e. primary sedimentation of organic carbon corrected by decomposition) was estimated to be 32% of the net primary production during the whole productive season and 42% in spring when the flux of primary settling material was greatest. Sedimentation of the spring bloom was the major annual supply of organic matter to the benthos (>80% of the total primary sedimentation).     

Bacterioplankton and Phytoplankton Populations in a Rapidly-Flushed Eutrophic Reservoir

Densities of bacterioplankton and phytoplankton were studied in relation to water retention time, temperature, nutrient concentrations, and other variables in a eutrophic montane reservoir (Bluestone Lake, West Virginia, U.S.A.). Weak temperature stratification occurred occasionally during the summer, but the reservoir did not become anaerobic. Water retention times were short (3.9 to 9.7 days between June and December), and differences in flushing rate within that range were important in regulating phytoplankton populations. Green algae and planktonic diatoms were dominant in summer during periods when water residence times were shorter. The occurrence of blue-green algal blooms in fall was related to increased water residence times. Retention time was highly correlated with the percent composition of phytoplankton represented by Anabaena spp. and Microcystis sp. (r=0.898, p<0.001). Advective effects limited phytoplankton biomass when retention time was shortest, but were not as important when retention times were longer. Peaks in bacterial densities often coincided with decreases in viable algal cell densities, so bacterioplankton were less directly related to retention time.     

Origin and succession of phytoplankton in a river-lake system (Spree,Germany)

The River Spree (Germany) flows through an impoundment and several shallow lakes in its middle and lower course. In this river-lake system, the seasonal and longitudinal dynamics of dominant phytoplankton populations were studied in relation to retention time of water, mixing conditions and nutrient supply from 1988–92. Some phytoplankton species populated the same river section for weeks or months each year at their season. Such stable populations have to origin from river zones functioning like mixed reactors. In the Spree system, centric diatoms originated from an impoundment and filamentous cyanobacteria from a flushed lake with longer retention time of water. Downstream, biomass and composition of phytoplankton altered nearly simultaneously along the system.The fate of planktonic organisms washed from mixed reactors into the flow depended on the conditions at the zones of origin. During spring, populations dominating phytoplankton communities of the well-mixed lakes grew further under river conditions. However the biomass of summer species, adapted to intermittent stratification, was halved along the river course. These seasonal differences were probably caused by lower maximum growth rates of summer species and enhanced losses (photorespiration, sedimentation or grazing of benthic filter feeders, but not of zooplankton) of algal populations under river conditions in summer.Phytoplankton assimilation, settlement of diatoms, or denitrification caused declining (probably growth limiting) concentrations of dissolved inorganic phosphorus (spring), silicon (early summer) or nitrogen (summer) along the river course, respectively. The minimum content of DRP was often followed by a clear-water phase. Reduced DSi supply selected against diatoms and additional DIN shortage favoured N₂-fixing cyanobacteria in the last lake of the system.R-strategists (sensu Reynolds) were selected in both the flushed, shallow lakes and the lowland river. In general, the biomass of cyanobacteria increased within the lakes and declined along the river course. Some diatom populations grew in the river, but were grazed or settled down in the lakes. Beside this general picture, different populations from the same phylogenetic group did not necessarily perform in similar ways.     

Effects of N and P supply on phytoplankton in Bizerte Lagoon (western Mediterranean)

Enriched bottle experiments were conducted in situ during winter (January and February) and summer (July and August) 2001 to examine the effects of nutrient enrichments (+ N, + P and + NP) on phytoplankton in Bizerte Lagoon, Tunisia. Chlorophyll a (Chl a), ranging from 3.05 μg L⁻¹ in winter to 4.52 μg L⁻¹ in summer, was dominated by the small size-faction (<5 μm) during both seasons. However, the contribution of the large size-fraction (5-200 μm) to Chl a increased from winter (26%) to summer (37%). Similarly, the carbon biomass of the 5-200 μm algae increased during the July/August period that was characterised by the high proliferation of several diatom taxa. In winter, N was the limiting element for phytoplankton growth. Its addition alone (+ N) or with P (+ NP) increased both the <5 μm and 5-200 μm Chl a concentrations. There was no change in the phytoplankton size structure, with the small cells dominating the final algal biomass in all treatments after 5 days. In summer, N and P limited the phytoplankton, but small and large algae exhibited diverse responses to different nutrient enrichments: addition of P increased the Chl a only in the 5-200 μm fraction, the + N treatment enhanced both size classes, and the NP fertilisation mostly stimulated the biomass of large cells. Consequently, the N and P addition in summer was followed by a significant change in the phytoplankton size structure, since both size-fractions contributed equally to the final Chl a biomass. Within the 5-200 μm algal community, various taxa had diverse responses to the nutrient supply during both seasons, leading to a change in the final community composition. The autotrophic flagellates appeared to grow well under N-deficient conditions. In contrast, diatom growth and biomass were mostly stimulated by the N enrichment while dinoflagellates exhibited the highest increase in their growth and biomass with P fertilisation. Our results suggest that the increasing anthropogenic supply of nutrients in the lagoon may influence algal dynamics as well as productivity in different ways depending on the nutrient composition.     

Importance of sediment deposition and denitrification for nutrient retention in floodplain wetlands

Questions: Various floodplain communities may differ in their relative abilities to influence water quality through nutrient retention and denitrification. Our main questions were: (1) what is the importance of sediment deposition and denitrification for plant productivity and nutrient retention in floodplains; (2) will rehabilitation of natural floodplain communities (semi‐natural grassland, reedbed, woodland, pond) from agricultural grassland affect nutrient retention? Location: Floodplains of two Rhine distributaries (rivers Ussel and Waal), The Netherlands. Methods: Net sedimentation was measured using mats, denitrification in soil cores by acetylene inhibition and bio‐mass production by clipping above‐ground vegetation in winter and summer. Results: Sediment deposition was a major source of N and P in all floodplain communities. Highest deposition rates were found where water velocity was reduced by vegetation structure (reedbeds) or by a drop in surface elevation (pond). Sediment deposition was not higher in woodlands than in grassland types. Denitrification rates were low in winter but significantly higher in summer. Highest denitrification rates were found in an agricultural grassland (winter and summer) and in the ponds (summer). Plant productivity and nutrient uptake were high in reedbeds, intermediate in agricultural grasslands, ponds and semi‐natural grasslands and very low in woodlands (only understorey). All wetlands were N‐limited, which could be explained by low N:P ratios in sediment. Conclusions: Considering Rhine water quality: only substantial P‐retention is expected because, relative to the annual nutrient loads in the river, the floodplains are important sinks for P, but much less for N. Rehabilitation of agricultural grasslands into ponds or reedbeds will probably be more beneficial for downstream water quality (lower P‐concentrations) than into woodlands or semi‐natural grasslands.     

The effects of nutrient addition and pH manipulation in bag experiments on the phytoplankton of a small acidic lake in West Virginia,USA

In situ bag experiments were performed during summer and autumn in a small acidic lake, Tibbs Run Lake, West Virginia, USA. The objective was to evaluate phytoplankton responses to pH manipulation and nutrient addition. Increasing the pH from below 4.5 to over 6.3 resulted in great declines in phytoplankton biovolume. There was also a succession from dinoflagellates (Peridinium inconspicuum to small chlorophytes. The trend was more rapid where phosphorus (P) additions were made along with pH enhancement. During summer, P limitation was indicated, while nitrogen (N) appeared to limit production in autumn. In both seasons, nutrient additions greatly altered the phytoplankton composition in high pH treatments, but had no discernable effects at (the natural) low pH. A low pH, P addition treatment in autumn was the single exception. When N was subsequently added, phytoplankton composition changed dramatically, probably because the proceeding P additions caused severe secondary N-limitation. In general, however, the results supported the view that phytoplankton compositional responses to nutrient additions are suppressed in low pH, relative to high pH lake water.     

Response of Sargasso Sea phytoplankton biomass, growth rates and primary production to seasonally varying physical forcing

The response of phytoplankton biomass, growth rates and primaryproduction to seasonally varying physical forcing was studiedat a station southeast of Bermuda over an 18 month period. Phytoplanktongrowth rates and primary production were measured using thepigment-labeling method, and phytoplankton biomass was calculatedfrom these measurements. Phytoplankton carbon biomass variedsystematically over the year. Highest values were observed duringthe winter and spring. Seasonal variations of chlorophyll (Chi)a in the surface layer could primarily be attributed to variationsin phytoplankton biomass and secondarily to photoacclimation.During the summer period, average values of carbon (C)/Chl ratios(g C g^–1 Chi) ranged from 160 at the surface to 33 atthe 1.6% light level, changes attributed to photoacclimationof the phytoplankton, consistent with the observation that phytoplanktonbiomass did not vary as a function of depth. Phytoplankton growthrates in the surface layer did not vary systematically overthe year, ranging from 0.15 to 0.45 day^–1, in spite ofseasonally varying concentrations of nitrate. Growth rates variedas a function of depth from average values of 0.3 day^–1in the surface layer to <0.1 day¹ at the 1.6% light level.Thus, the primary response of the phytoplankton community tonutrient enrichment during the winter period was an increasein phytoplankton biomass rather than an increase in growth rates.A simple nutrient-phyto-plankton-zooplankton model was usedto explore this phenomenon. The model demonstrated that theobserved response of the phytoplankton to nutrient enrichmentis only possible when phytoplankton growth is not severely limitedby nutrients.