Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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Nutrient limitation of phytoplankton and periphyton growth in upland lakes

SUMMARY 1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were visited three times between April and August 2000. On each occasion water chemistry was measured and phytoplankton bioassays were performed in the laboratory to assess growth‐rate and yield limitation by phosphorus and nitrogen. In addition, yield limitation of periphyton growth was investigated twice, in situ, using nutrient‐diffusing substrata. 2. Over the whole season the percentage frequency of P, N and co‐limitation was 24, 13 and 63%, respectively, for phytoplankton rate limitation and 20, 22 and 58%, respectively, for phytoplankton yield limitation. 3. A clear response of periphyton yield to nutrient additions was found in 75% of all cases and of these, co‐limitation was most common (54%). Average percentage frequency for P and N limitation was 26 and 20%, respectively. 4. Phytoplankton and periphyton showed seasonal changes in nutrient limitation within sites. In particular, co‐limitation became progressively more common as the season progressed. 5. The response of phytoplankton growth rate to ammonium and nitrate addition was identical, but ammonium was a slightly better source of nitrogen than nitrate for phytoplankton yield on 7% and for periphyton yield on 10% of the occasions. However, the magnitude of the effect was small. 6. The concentration of dissolved inorganic nitrogen (DIN) and the molar ratio of DIN to total dissolved phosphorus (TDP), appeared to be the main environmental factors controlling the extent of nitrogen or phosphorus limitation at a given site. Nitrogen limitation was more likely than phosphorus limitation where the DIN was <6.5 mmol m^?3 and the ratio of DIN : TDP was <53. Co‐limitation was the most likely outcome at a DIN concentration <13 mmol m^?3 and at a DIN : TDP molar ratio <250. Above these values phosphorus limitation was most likely. 7. The relatively high frequency of nitrogen limitation and co‐limitation at higher N : P ratios than previously reported, may result from the inability of nitrogen‐fixing cyanobacteria to thrive in these upland lakes where pH and the concentration of phosphorus tended to be low and where flushing rates tended to be high.     

Assessing the sensitivity of alpine lakes and ponds to nitrogen deposition in the Canadian Rocky Mountains

To demonstrate the sensitivity of aquatic ecosystems to forecasted increases in nitrogen deposition along the eastern ranges of the Canadian Rocky Mountains, we conducted midsummer limnological surveys of 29 remote alpine lakes and ponds via helicopter in 2007. Chemical analysis of water and in vitro nitrogen-enrichment bioassays of phytoplankton collected from each site were performed to estimate nutrient limitation. Use of a common chemical index for nutrient limitation (total dissolved inorganic nitrogen: total phosphorus; DIN:TP) together with supportive experimental evidence revealed nitrogen limitation in only 14% of the cases. Shallow (≤1 m maximum depth) ponds were more likely to be nitrogen-limited than lakes, especially as the former exhibited a significantly lower mean DIN:TP ratio of 7.4 during the late summer. Chemical and bioassay-based inferences of nitrogen limitation agreed in 74.5% of the cases, owing mainly to evidence of phosphorus limitation of the surveyed lake ecosystems (mean DIN:TP = 18) being supported by nonsignificant responses of their phytoplankton to nitrogen amendment. Our findings reveal that increased nitrogen deposition should not result in immediate widespread eutrophication of the Canadian Rockies; however, certain alpine ponds appeared nitrogen-limited, making them sensitive early indicators of the potential effects of anthropogenic nitrogen deposition in remote mountainous regions.     

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.     

Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

Nutrient limitation of phytoplankton communities in Subarctic lakes and ponds in Wapusk National Park, Canada

We determined the limiting nutrient of phytoplankton in 21 lakes and ponds in Wapusk National Park, Canada, using nutrient enrichment bioassays to assess the response of natural phytoplankton communities to nitrogen and phosphorus additions. The goal was to determine whether these Subarctic lakes and ponds were nutrient (N or P) limited, and to improve the ability to predict future impacts of increased nutrient loading associated with climate change. We found that 38% of lakes were not limited by nitrogen or phosphorus, 26% were co-limited by N and P, 26% were P-limited and 13% were N-limited. TN/TP, DIN/TP and NO₃ ⁻/TP ratios from each lake were compared to the Redfield ratio to predict the limiting nutrient; however, these predictors only agreed with 29% of the bioassay results, suggesting that nutrient ratios do not provide a true measure of nutrient limitation within this region. The N-limited lakes had significantly different phytoplankton community composition with more chrysophytes and Anabaena sp. compared to all other lakes. N and P limitation of phytoplankton communities within Wapusk National Park lakes and ponds suggests that increased phytoplankton biomass may result in response to increased nutrient loading associated with environmental change.     

Seasonal Patterns of Nitrogen and Phosphorus Limitation in Four German Lakes and the Predictability of Limitation Status from Ambient Nutrient Concentrations

To identify the seasonal pattern of nitrogen (N) and phosphorus (P) limitation of phytoplankton in four different lakes, biweekly experiments were conducted from the end of March to September 2011. Lake water samples were enriched with N, P or both nutrients and incubated under two different light intensities. Chlorophyll a fluorescence (Chla) was measured and a model selection procedure was used to assign bioassay outcomes to different limitation categories. N and P were both limiting at some point. For the shallow lakes there was a trend from P limitation in spring to N or light limitation later in the year, while the deep lake remained predominantly P limited. To determine the ability of in-lake N:P ratios to predict the relative strength of N vs. P limitation, three separate regression models were fit with the log-transformed ratio of Chla of the P and N treatments (Response ratio = RR) as the response variable and those of ambient total phosphorus:total nitrogen (TN:TP), dissolved inorganic nitrogen:soluble reactive phosphorus (DIN:SRP), TN:SRP and DIN:TP mass ratios as predictors. All four N:P ratios had significant positive relationships with RR, such that high N:P ratios were associated with P limitation and low N:P ratios with N limitation. The TN:TP and DIN:TP ratios performed better than the DIN:SRP and TN:SRP in terms of misclassification rate and the DIN:TP ratio had the highest R² value. Nitrogen limitation was predictable, frequent and persistent, suggesting that nitrogen reduction could play a role in water quality management. However, there is still uncertainty about the efficacy of N restriction to control populations of N₂ fixing cyanobacteria.     

Changes in N and P limitation induced by water level fluctuations in Nature Park Kopački Rit (Croatia): nutrient enrichment bioassay

The importance of nutrients as limiting factors might vary in environments with different limnological characteristics. In this article we evaluate the effect of frequency and duration of flooding on nutrient limitation in a riparian floodplain. Variations in N and P limitations were studied in the period of low (2003) and high (2004) water level in two different floodplain habitat types in the Nature Park Kopački Rit (Croatia), a floodplain area of the Danube River. In 2003 and 2004, the limnological characteristics of floodplain lake (Lake Sakadaš) and the channel (Stara Drava) differed due to their hydrological regimes. Potential for nutrient limitation was determined by DIN:TP and TN:TP ratios, while the actual nutrient limitation was assessed by nutrient enrichment bioassay. A change from non-limited to N-limited conditions in the channel, and consistent actual N limitation in the lake was determined by the nutrient enrichment experiment. Of the two ratios, DIN:TP matched better with the bioassay data. Phosphorus limitation was only occasionally evident. Changes in trophic conditions from hypereutrophy to eutrophy (between low and high water levels) reflected the importance of the hydrologic regime as a factor which can modify the trophic state of Lake Sakadaš. Compared with 2003, the increase of total nitrogen concentrations in 2004 calls attention to the importance of nitrogen inputs from the Danube to the system with excessive phosphorus concentrations.     

Pelagic phytoplankton community change‐points across nutrient gradients and in response to invasive mussels

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Field evidence for stoichiometric relationships between zooplankton and N and P availability in a shallow calcareous lake

1. According to stoichiometric theory, zooplankters have a species‐specific elemental composition. Daphniids have a relatively high phosphorus concentration in their tissues and copepods high nitrogen. Daphniids should, therefore, be more sensitive to phosphorus limitation and copepods more sensitive to nitrogen. A 2‐year study of a shallow marl lake in the west of Ireland investigated whether population fluctuations of the two dominant taxa, Daphnia spp. and the calanoid Eudiaptomus gracilis, were associated with the availability of phosphorus and nitrogen. 2. In accordance with stoichiometric predictions, Daphnia and Eudiaptomus reproduction had contrasting relationships with dietary phosphorus and nitrogen availability. Egg production by Daphnia was negatively associated with the ratio of dissolved inorganic nitrogen (DIN) : total phosphorus (TP) and the ratio of light to TP which was used as an indirect index for seston carbon (C) : phosphorus (P). Conversely calanoid egg production had a positive relationship with the DIN : TP ratio and was unrelated to the estimated C : P (light : TP) ratio. 3. Daphnia biomass was not, however, correlated with phosphorus availability, and neither was calanoid biomass correlated with nitrogen. The high ratio of DIN : TP when Daphnia dominated the zooplankton biomass and the low ratio when calanoids dominated, is consistent with Daphnia acting as a sink for phosphorus and calanoids as a sink for nitrogen and suggests consumer‐driven nutrient recycling.     

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。     

Nutrient stoichiometry of linked catchment‐lake systems along a gradient of land use

1. Catchments export nutrients to aquatic ecosystems at rates and ratios that are strongly influenced by land use practices, and within aquatic ecosystems nutrients can be processed, retained, lost to the atmosphere, or exported downstream. The stoichiometry of carbon and nutrients can influence ecosystem services such as water quality, nutrient limitation, biodiversity, eutrophication and the sequestration of nutrients and carbon in sediments. However, we know little about how nutrient stoichiometry varies along the pathway from terrestrial landscapes through aquatic systems. 2. We studied the stoichiometry of nitrogen and phosphorus exported by three catchments of contrasting land use (forest versus agriculture) and in the water column and sediments of downstream reservoirs. We also related stoichiometry to phytoplankton nutrient limitation and the abundance of heterocystous cyanobacteria. 3. The total N : P of stream exports varied greatly among catchments and was 18, 54 and 140 (molar) in the forested, mixed‐use and agricultural catchment, respectively. Total N : P in the mixed layers of the lakes was less variable but ordered similarly: 35, 52 132 in the forested, mixed‐use and agricultural lake, respectively. In contrast, there was little variation among systems in the C : N and C : P ratios of catchment exports or in reservoir seston. 4. Phytoplankton in the forested lake were consistently N limited, those in the agricultural lake were consistently P limited, and those in the mixed‐use lake shifted seasonally from P‐ to N limitation, reflecting N : P supply ratios. Total phytoplankton and cyanobacteria biomass were highest in the agricultural lake, but heterocystous (potentially N fixing) cyanobacteria were most abundant in the forested lake, corresponding to low N : P ratios. 5. Despite large differences in catchment export and water column N : P ratios, the N : P of sediment burial (integrated over several decades) was very low and remarkably similar (4.3–7.3) across reservoirs. N and P budgets constructed for the agricultural reservoir suggested that denitrification could be a major loss of N, and may help explain the relatively low N : P of buried sediment. 6. Our results show congruence between the catchment export N : P, reservoir N : P, phytoplankton N versus P limitation and the dominance of heterocystous cyanobacteria. However, the N : P stoichiometry of sediments retained in the lakes was relatively insensitive to catchment stoichiometry, suggesting that a common set of biogeochemical processes constrains sediment N : P across lakes of contrasting catchment land use.     

Nutrient-phytoplankton relationships in a tropical meromictic soda lake

Seasonal variation through one year in total nitrogen (TN), total phosphorus (TP), phytoplankton biomass, phytoplankton species composition and other environmental factors were examined in Lake Sonachi, a tropical meromictic soda lake. Mean concentrations of TN and TP were 11 000 µg N l^-1 and 100 µg P l^-1, respectively. Maximum concentrations of TN and TP occurred in the monimolimnion. Phytoplankton biomass ranged from 350 to 1260 mg m^-3. Synechococcus bacillaris, a small coccoid cyanophyte, dominated the phytoplankton. The mean chlorophyll a concentration of 37 mg · m^-3 was a modest value when compared with those of other tropical soda lakes. High TN:TP ratios indicated phosphorus limitation in the lake.     

Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. This synthesis examines 35 long‐term (5–35 years, mean: 16 years) lake re‐oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in‐lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10–15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in‐lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100–150 μg L^?1. This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

The effect of increased nitrogen load on phytoplankton in a phosphorus‐limited lake

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Do Nutrient Limitation Patterns Shift from Nitrogen Toward Phosphorus with Increasing Nitrogen Deposition Across the Northeastern United States?

Atmospheric nitrogen (N) deposition is altering biogeochemical cycling in forests and interconnected lakes of the northeastern US, and may shift nutrient limitation from N toward other essential elements, such as phosphorus (P). Whether this shift is occurring relative to N deposition gradients across the northeastern US has not been investigated. We used datasets for the northeastern US and the Adirondack sub-region to evaluate whether P limitation is increasing where N deposition is high at two geographic scales, based on N:P mass ratios. Using a model-selection approach, we determined that foliar N for dominant tree species and lake dissolved inorganic N (DIN) increased coincident with increasing N deposition, independent of relationships between foliar N or lake DIN and precipitation or temperature. Foliar P also increased with N deposition across the northeastern US for seven of eight deciduous species, but changed less across the Adirondacks. Foliar N:P therefore declined at the highest levels of N deposition for most deciduous species across the region (remaining nearly constant for most conifers and increasing only for black cherry and hemlock), but increased across all species in the Adirondacks. Ratios between DIN and total P (DIN:TP) in lakes were unrelated to N deposition regionally but increased across the Adirondacks. Thus, nutrient limitation patterns shifted from N toward P for dominant trees, and further toward P for predominantly P-limited lakes, at the sub-regional but not regional scale. For the northeastern US overall, accumulated N deposition may be insufficient to drive nutrient limitation from N toward P; alternatively, elements other than P (for example, calcium, magnesium) may become limiting as N accumulates. The consistent Adirondack foliar and lake response could provide early indication of shifts toward P limitation within the northeastern US, and together with regional patterns, suggests that foliar chemistry could be a predictor of lake chemistry in the context of N deposition across the region.     

Determining trophic state in Lake Whatcom,Washington (USA), a soft water lake exhibiting seasonal nitrogen limitation

We evaluated an eleven year data set to assess trophic state and nutrient limitation in Lake Whatcom, an oligotrophic, soft water, chain lake located in the Puget Sound lowlands of Washington (U.S.A.). Although total phosphorus (TP) and soluble reactive phosphate (SRP) concentrations were relatively low throughout the lake, there were significant differences between the northern basin (Site 1) and the other sampling sites (Sites 2–4). Nonparametric correlation coefficients (Kendall's ) were highest between chlorophyll (CHL), Secchi depth (SD), total nitrogen (TN), and dissolved inorganic nitrogen (DIN). Late summer algal biomass correlated best with DIN and TP. Trophic State Indices based on TP, TN, CHL and SD revealed that although algal growth was most likely phosphorus limited throughout the year, the northern basin of the lake may have developed nitrogen co-limitation during late summer and fall. During this period, N/P ratios were often less than 20, and in 1998 the epilimnetic DIN concentrations dropped below 20 g l^–1 while DIN/TP ratios fell below 4. Reviews of the literature suggest that while co-limitation by phosphorus and nitrogen is fairly common in unproductive lakes, the patterns seen in Lake Whatcom were more similar to those reported for eutrophic lakes experiencing secondary nitrogen limitation resulting from excess phosphorus loading.     

TN : TP ratio and planktivorous fish do not affect nutrient-chlorophyll relationships in shallow lakes

1. In previous work, phytoplankton regulation in freshwater lakes has been associated with many factors. Among these, the ratio of total nitrogen to total phosphorus (TN : TP) has been widely proposed as an index to identify whether phytoplankton are N‐ or P‐limited. From another point of view, it has been suggested that planktivorous fish can be used to control phytoplankton. 2. Large‐scale investigations of phytoplankton biomass [measured as chlorophyll a, (chl‐a)] were carried out in 45 mid‐lower Yangtze shallow lakes to test hypotheses concerning nutrient limitation (assessed with TN : TP ratios) and phytoplankton control by planktivorous fish. 3. Regression analyses indicated that TP was the primary regulating factor and TN the second regulating factor for both annual and summer phytoplankton chl‐a. In separate nutrient–chl‐a regression analyses for lakes of different TN : TP ratios, TP was also superior to TN in predicting chl‐a at all particular TN : TP ranges and over the entire TN : TP spectrum. Further analyses found that chl‐a : TP was not influenced by TN : TP, while chl‐a : TN was positively and highly correlated to TP : TN. 4. Based on these results, and others in the literature, we argue that the TN : TP ratio is inappropriate as an index to identify limiting nutrients. It is almost impossible to specify a ‘cut‐off’ TN : TP ratio to identify a limiting nutrient for a multi‐species community because optimal N : P ratios vary greatly among phytoplankton species. 5. Lakes with yields of planktivorous fish (silver and bighead carp, the species native to China) >100 kg ha^?1 had significantly higher chl‐a and lower Secchi depth than those with yields <100 kg ha^?1. TP–chl‐a and TP–Secchi depth relationships are not significantly different between lakes with yields >100 kg ha^?1 or <100 kg ha^?1. These results indicate that the fish failed to decrease chl‐a yield or enhance Z_SD. Therefore, silver carp and bighead carp are not recommended as a biotic agent for phytoplankton control in lake management if the goal is to control the entire phytoplankton and to enhance water quality.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.     

Phytoplankton nutrient deficiency in lakes of the McMurdo dry valleys, Antarctica

1. The influence of inorganic nitrogen and phosphorus enrichment on phytoplankton photosynthesis was investigated in Lakes Bonney (east and west lobes), Hoare, Fryxell and Vanda, which lie in the ablation valleys adjacent to McMurdo Sound, Antarctica. Bioassay experiments were conducted during the austral summer on phytoplankton populations just beneath the permanent ice cover in all lakes and on populations forming deep-chlorophyll maxima in the east and west lobes of Lake Bonney. 2. Phytoplankton photosynthesis in surface and mid-depth (13 m) samples from both lobes of Lake Bonney were stimulated significantly (P < 0.01) by phosphorus enrichment (2 μM) with further stimulation by simultaneous phosphorus plus NH₄⁺ (20 μM) enrichment. Similar trends were observed in deeper waters (18 m) from the east lobe of Lake Bonney, although they were not statistically significant at P < 0.05. Photosynthesis in this lake was never enhanced by the addition of 20 μM NH₄⁺ alone. Simultaneous addition of phosphorus plus nitrogen stimulated photosynthesis significantly (P < 0.01) in both Lake Hoare and Lake Fryxell. No nutrient response occurred in Lake Vanda, where activity in nutrient-enriched samples was below unamended controls; results from Lake Vanda are suspect owing to excessively long sample storage in the field resulting from logistic constraints. 3. Ambient dissolved inorganic nitrogen (DIN) (NH4⁺+ NO₂^?+ NO₃^?): soluble reactive phosphorus (SRP) ratios partially support results from bioassay experiments indicating strong phosphorus deficiency in Lake Bonney and nitrogen deficiency in Lakes Hoare and Fryxell. DIN : SRP ratios also imply phosphorus deficiency in Lake Vanda, although not as strong as in Lake Bonney. Particulate carbon (PC): particulate nitrogen (PN) ratios all exceed published ratios for balanced phytoplankton growth, indicative of nitrogen deficiency. 4. Vertical nutrient profiles in concert with low advective flux, indicate that new (sensu Dugdale & Goering, 1967) phytoplankton production in these lakes is supported by upward diffusion of nutrients from deep nutrient pools. This contention was tested by computing upward DIN : SRP flux ratios across horizontal planes located immediately beneath each chlorophyll maximum and about 2 m beneath the ice (to examine flux to the phytoplankton immediately below the ice cover). These flux ratios further corroborated nutrient bioassay results and bulk DIN : SRP ratios indicating phosphorus deficiency in Lakes Bonney and Vanda and potential nitrogen deficiency in Lakes Hoare and Fryxell. 5. Neither biochemical reactions nor physical processes appear to be responsible for differences in nutrient deficiency among the study lakes. The differences may instead be related to conditions which existed before or during the evolution of the lakes.