Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

太平洋中西部海域浮游植物营养盐的潜在限制

2009年8月至9月期间在太平洋西部N1站和中部N2站进行现场营养盐加富培养实验。结果显示:N1站,浮游植物生物量对N或者P添加都有较强的响应,其中N+P+Si组和N+P组浮游植物长势迅速,叶绿素a从初始的0.03μg/L分别达到2.12μg/L和1.83μg/L,同时P先于N和Si之前被耗尽;说明N1站为N、P共同限制,P是首要限制因子。而N2站,浮游植物生物量仅对N、P共同添加有明显响应,N先于P和Si被浮游植物消耗殆尽。利用培养过程中营养盐比值变化推断,N1站浮游植物以低于Redfield ratio(16N∶1P)吸收N和P;而N2站浮游植物以高于Redfield ratio(16N∶1P)吸收N和P。这可能解释了太平洋西部的寡营养盐海域为潜在P限制,而在太平洋中部海域则为潜在N限制。     

An analysis,by bioassay,of the factors which limit algal growth in the P. K. le Roux Impoundment,Orange River,South Africa

Factors which limit algal growth in a turbid man-made impoundment were investigated using natural phytoplankton populations. Results indicate that light was the primary limiting factor with nutrients possibly limiting growth in the event of an increase in water transparency. These results differ from another study which used Selenastrum capricornutum bioassays of water from other areas of the Orange River. The value of natural phytoplankton bioassay techniques is supported.     

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.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Characterisation of the type and extent of nutrient limitation in grassland vegetation using a bioassay with intact sods

An important methodological problem in plant ecology concerns the way in which the type and extent of nutrient limitation in terrestrial communities should be assessed. Conclusions on nutrient limitation have been founded mainly on soil extractions, fertiliser trials and tissue nutrient concentrations. In order to avoid some of the problems associated with these methods, we employed a special technique using intact sods which rooted both in the intact soil and in a nutrient solution, from which N, P and K were omitted stepwise. The method was applied to hay-field communities which differed in their history of fertiliser application. Four fields were compared which were not fertilised for 2, 6, 19 or 45 years, while hay making continued. This was done to restore former species-rich grassland communities. We tested the hypothesis that the increase in species diversity in these grasslands was attended by an increase in the number of limiting nutrients.We observed clear shifts in the type and extent of nutrient limitation. Fields which were recently fertilised were characterised by nitrogen and potassium limitation, while phosphorus limitation increased in importance towards the later stages of succession. In the last field (45 years unfertilised) N, P and K equally limited production at the community level. These conclusions differed from those drawn from a fertiliser trial in these same four fields, which failed to detect phosphorus limitation.It is concluded that the use of this method provides a valuable extra source of information while studying relationships between nutrient limitation and species diversity in grassland communities.     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

浙江紧水滩水库浮游植物群落结构季节变化特征

于2010年1、3、5、7、9、11月6次对紧水滩水库采样调查,并对浮游植物种类鉴定与数量统计,分析了浮游植物的优势种、多样性和群落结构季节变化特征.其结果为:共鉴定浮游植物284种,隶属7门105属.绿藻门最多,共51属139种,其次是硅藻门19属67种,蓝藻门22属52种,金藻门4属9种,甲藻门5属8种,裸藻门2属5种,隐藻门2属4种.浮游植物细胞丰度在1.04× 105-3.70×106个/L之间,平均丰度9.63×105个/L.多样性指数H'值为1.76-4.64,平均值3.09,丰富度指数D为0.48-2.80,平均值1.62,均匀度指数.J为0.51-1.26,平均值0.91.根据TSI(∑)并结合浮游植物群落结构对水质评价,紧水滩水库水质属于中-富营养状态.     

南海北部浮游植物生长对营养盐的响应

2004年夏季作者在南海北部海域研究了浮游植物生长的营养动力学,结合物理-化学过程对浮游植物生物量分布的影响与机制进行了研究,阐明了水平对流和中尺度涡对营养盐分布的影响及浮游植物生长和现存生物量对其的响应。受西南季风和东向沿岸流作用所形成的Ekman输送的影响,南海北部海岸带表层海水作离岸运动,使深层富含营养盐的冷水爬坡涌升到表层来补充,激发浮游植物生物量迅速增长。海区反气旋涡使海水辐聚下沉,造成水体具高温、低盐、高溶解氧浓度、低营养盐浓度和低浮游植物生物量。同时通过现场营养盐加富试验,发现该海域营养盐是浮游植物生长的主要限制因子,而且是多种营养元素共同限制了浮游植物的生长,添加单一的营养盐并不能促进浮游植物的生长。在生物量出现增长的试验组中,营养盐添加不仅促使浮游植物生物量的增长,而且也改变了浮游植物的粒级结构和群落结构。例如,在站S1008,培养前叶绿素a浓度为0.28 mg.m-3,加富培养60 h后浮游植物生物量在NP和NPSi的试验组中有显著的增加,叶绿素a浓度分别达1.07 mg.m-3和1.19 mg.m-3;培养前粒度分级叶绿素a主要以Pico级份占优势,而加富试验结束后,在NP和NPSi的试验组以Nano级份占优势,其它试验组仍以Pico级份占优势;同时,在培养后生物量出现增长的试验组,浮游植物群落的优势类群从甲藻向硅藻演替。     

The influence of a dissolved inorganic nitrogen gradient on phytoplankton community dynamics in a chain of lakes

In a chain of lakes along which nutrient availability varies in a gradient, we performed factorial nutrient enrichment experiments to determine if nitrogen limitation was the principal factor controlling the differences in phytoplankton biomass, photosynthetic productivity, diversity, and species composition among two of the lakes in the chain. In the least productive lake, East Graham Lake, P and C enrichments (in the absence of N enrichment) had no effect on biomass and diversity, whereas within two weeks the N enrichments (alone or in any combination with P and/or C) increased the biomass and decreased the diversity of East Graham Lake phytoplankton to levels similar or identical to those in more productive Shoe Lake. Short-term ¹⁴C photosynthetic rates in East Graham Lake water also responded only to N in the third week. However, photosynthesis was stimulated by P in the first week, and a few species did increase in numbers with P enrichment, suggesting that some degree of P limitation remains in addition to the strong N limitation in East Graham Lake. A number of species responded individually to the enrichments in a manner similar to that of the overall community, and a strong overlapping of discriminant analysis scores for N-enriched East Graham Lake with those of Shoe Lake was consistent with our prediction that the community structure of N-enriched East Graham Lake water would shift toward that of Shoe Lake. However, many species did not respond consistently with these results, and the nutrients tested were clearly not a major factor in the differences in abundance of those species among the two lakes. The results support the argument that overall biomass production and diversity of the phytoplankton community in a lake can be a relatively simple function of a single most-limiting nutrient. However, many of the species responses also confirm that, while nutrient availability is an important factor in the control of the species composition of the community, other factors are likely to prevent reliable predictions of all species effects on the basis of nutrient availability alone.     

A bioassay to assess the potential effects of sediment resuspension on phytoplankton community composition

A laboratory assay (SAGA or Sediment Algal Growth Assay) was developed to assess the potential impact of sediment resuspension on the structure of phytoplankton communities, and to evaluate the effectiveness of various sediment treatments in decreasing the abundance of blue-green algae in the event of sediment resuspension during storms. In assays with sediment from eutrophic Akanoi Bay, Lake Biwa, Japan, 7–11 species of phytoplankton seeded from the sediments grew during the 3-week assay indicating that sediment resuspension has the potential to increase both phytoplankton biomass and species diversity. Treatment of sediments with Ca(NO3)2 substantially decreased phytoplankton biomass (measured as chlorophyll concentration) in assays with sediments from Akanoi Bay and the North Basin of Lake Biwa. Further, among various oxidation treatments of sediments, Ca(NO3)2 was most effective in decreasing or preventing filamentous blue-green algal growth in N- and P-replete media. In contrast, when sediments were added to P-limited phytoplankton dominated by green algae and diatoms, no growth of blue-green algae occurred regardless of sediment treatment. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ecology of the phytoplankton of the River Moselle: effects of disturbances on community structure and diversity

A data set on community composition of the phytoplankton of the River Moselle (France) has been used for testing the intermediate disturbance hypothesis (IDH). After a short presentation of the ecology of the river and of its phytoplankton, the main changes in composition and diversity of the suspended algal assemblage are described. It is emphasized that discharge fluctuations, related to weather changes, play a key role, but that biotic factors such as grazing and parasitism may also influence diversity in stable summer conditions.     

Effects of upstream lakes and nutrient limitation on periphytic biomass and nitrogen fixation in oligotrophic, subalpine streams

1. We conducted bioassays of nutrient limitation to understand how macronutrients and the position of streams relative to lakes control nitrogen (N₂) fixation and periphytic biomass in three oligotrophic Rocky Mountain catchments. We measured periphytic chlorophyll‐a (chl‐a) and nitrogen‐fixation responses to nitrogen (N) and phosphorus (P) additions using nutrient‐diffusing substrata at 19 stream study sites, located above and below lakes within the study catchments. 2. We found that periphytic chl‐a was significantly co‐limited by N and P at 13 of the 19 sites, with sole limitation by P observed at another four sites, and no nutrient response at the final two sites. On average, the addition of N, P and N + P stimulated chl‐a 35%, 114% and 700% above control values respectively. The addition of P alone stimulated nitrogen fixation by 2500% at five of the 19 sites. The addition of N, either with or without simultaneous P addition, suppressed nitrogen fixation by 73% at nine of the 19 sites. 3. Lake outlet streams were warmer and had higher dissolved organic carbon concentrations than inlet streams and those further upstream, but position relative to lakes did not affect chl‐a and nitrogen fixation in the absence of nutrient additions. Chl‐a response to nutrient additions did not change along the length of the study streams, but nitrogen fixation was suppressed more strongly by N, and stimulated more strongly by P, at lower altitude sites. The responses of chl‐a and nitrogen fixation to nutrients were not affected by location relative to lakes. Some variation in responses to nutrients could be explained by nitrate and/or total N concentration. 4. Periphytic chl‐a and nitrogen fixation were affected by nutrient supply, but responses to nutrients were independent of stream position in the landscape relative to lakes. Understanding interactions between nutrient supply, nitrogen fixation and chl‐a may help predict periphytic responses to future perturbations of oligotrophic streams, such as the deposition of atmospheric N.     

Changes in phytoplankton community along a transversal section of the Lower Paraná floodplain,Argentina

Interactions between the main stream of large rivers and their floodplains are usually observed, determining gradual changes in environmental variables along a transversal axis. This paper studies the natural phytoplanktonic community, as well as the main physical and chemical properties from a transversal section of the Lower Paraná River floodplain, encompassing the main channel, a lake and a swamp. In vitro nutrient bioassays for phytoplankton growth were performed in order to evaluate the importance of nutrients as potential limiting factors. A decrease in depth, pH, dissolved oxygen, suspended solids, DIN (dissolved inorganic N), particularly nitrates, and DIN:SRP (soluble reactive P) ratio, was observed from the main channel towards the swamp; as well as an increase in transparency, chlorophyll-a and phytoplankton density. Phytoplankton from the mainstream consisted mostly of typical potamoplanktonic species adapted to grow in turbid environments (mainly Aulacoseira granulata (Müll.) Sim. and its varieties), while in the swamp it was composed mainly of mixotrophic phytoflagellates (euglenoids and cryptomonads), facultative N-heterotrophic diatoms and small-size green algae. Values of both abiotic and biotic variables measured in the lake were always intermediate between the former ones, although closer to those of the river, an account of their permanent connection. Phytoplankton response to N-addition was more frequent in the swamp bioassays than in the other two sites. However, a great increase in algal density in the control flasks, suggests a plentiful source of N at the beginning of the experiments. Light scarcity is more likely to occur than N-limitation in both river and lake due to their low transparency and a high DIN concentration. Although N-limitation may be expected in the swamp, the growth in the controls in spite of the low initial DIN concentration, suggests the existence of an N-supply other than DIN (perhaps organic N). Rather than indicate in situ N-limitation, present results seem, therefore, to suggest that phytoplankton of the swamp mainly composed of mixotrophic algae would be adapted to grow under low DIN concentrations. Possible reasons for the absence of N₂-fixing Cyanobacteria in this environment are discussed.     

Ploidy evolution in the yeast Saccharomyces cerevisiae: a test of the nutrient limitation hypothesis

The nutrient limitation hypothesis provides a nongenetic explanation for the evolution of life cycles that retain both haploid and diploid phases: differences in nutrient requirements and uptake allow haploids to override the potential genetic advantages provided by diploidy under certain nutrient limiting conditions. The relative fitness of an isogenic series of haploid, diploid and tetraploid yeast cells (Saccharomyces cerevisiae), which were also equivalent at the mating type locus, was measured. Fitness was measured both by growth rate against a common competitor and by intrinsic growth rate in isolated cultures, under four environmental conditions: (1) rich medium (YPD) at the preferred growth temperature (30 °C); (2) nutrient poor medium (MM) at 30 °C; (3) YPD at a nonpreferred temperature (37 °C); and (4) MM at 37 °C. In contrast to the predictions of the nutrient limitation hypothesis, haploids grew significantly faster than diploids under nutrient rich conditions, but there were no apparent differences between them when fitness was determined by relative competitive ability. In addition, temperature affected the relative growth of haploids and diploids, with haploids growing proportionately faster at higher temperatures. Tetraploids performed very poorly under all conditions compared. Cell geometric parameters were not consistent predictors of fitness under the conditions measured.     

Short-term variations in the physiological state of phytoplankton [2pt] in a shallow temperate estuary

Short-term changes in the photosynthetic carbon metabolism and physiological state of phytoplankton were studied over a summer fortnight-long period in the Urdaibai estuary (Bay of Biscay) and related to observed environmental patterns. Day-to-day variability in the hydrographical and biological features of the estuary during the study period was due to changes in meteorological and tidal conditions. Phytoplankton biomass and primary production increased with the improvement of weather, i.e., light conditions, during neap tides. Thus a mixed bloom of cryptophyceans, Euglena sp., and the dinoflagellate Peridinium foliaceum developed in the middle and upper estuary. Photosynthetic responses of phytoplankton were related to the time-scale of changes in light regime. Allocation of photosynthate to major macromolecular classes (LMWM, lipid, polysaccharide, and protein), like phytoplankton biomass and primary production, showed strong spatio-temporal variability. High carbon fixation into low molecular weight metabolites was associated with growth limitation by low light. The relative incorporation of photosynthetic carbon into proteins increased at the beginning of the phytoplankton bloom but overall, it was rather constant. However, carbon allocation into storage products such us lipid or polysaccharide increased when carbon and energy produced under optimal growth conditions exceeded what could be assimilated into protein. These patterns are explained by both spatio-temporal changes in the environmental conditions and species-specific differences. In general, daily variability appeared to be more important than diurnal periodicity in the physiological responses of phytoplankton. Results from this study show that phytoplankton photosynthesis and carbon metabolism are simultaneously affected by biotic and abiotic factors, although short-term light fluctuations may have a major influence on the physiological state of phytoplankton in the Urdaibai estuary.     

Effects of modified nutrient concentrations and ratios on the structure and function of the native phytoplankton community in the Neuse River Estuary, North Carolina, USA

A variety of analyses were used to assess the structure (community composition) and function (assimilation number, nitrogen fixation) of phytoplankton in the Neuse River Estuary (NRE), NC under ambient and modified nutrient concentrations. Dilution bioassays were employed to reduce the concentration of nitrogen (N) or both N and phosphorus (P) and thus compare varied DIN:DIP ratios. Experimental manipulations created conditions that may result from mandated N load reductions to the estuary. We hypothesized that unilateral reduction of N loading to the NRE would increase the activity, abundance and diversity of N₂ fixing cyanobacteria. Changes in phytoplankton primary productivity, N₂ fixation (nitrogenase activity), genetic potential for N₂ fixation (presence of nifH), phytoplankton taxonomic composition (diagnostic photopigment concentration) and abundances of N₂ fixing cyanobacteria (microscopy) were determined. Decreasing ambient DIN:DIP ratios in NRE samples resulted in increased rates of N₂ fixation when seed populations were present and environmental conditions were amenable. Decreasing the DIN:DIP ratio did not lead to an increase in the abundance or diversity of N₂ fixing cyanobacteria. Because N₂ fixing cyanobacteria were only actively fixing nitrogen during periods of low riverine N discharge (summer and early autumn), lowering nutrient ratios may not have a major impact on the NRE. However, the maximum potential amount of N from N₂ fixation was calculated using rates from this study and was found to be approximately 3% of total riverine loading of N to the NRE. Because N₂ fixation occurs farther downstream and later in the year than riverine N loading to the NRE, there is potential for N₂ fixation to modify N dynamics. Analyses of the phytoplankton community as a whole in these relatively short term experiments indicated that reduced DIN:DIP may not have a major impact on their structure and function.     

Effects of Asian dust input on eukaryotic phytoplankton community structure in the open areas in the Northwestern Pacific Ocean

To study the effects of aerosol particulates originated from Asian dust on the growth of eukaryotic phytoplankton in the oligotrophic open ocean, we conducted deck-board incubation experiments in the oligotrophic region of Northwestern Pacific Ocean (NWPO). Our results showed that when dust was added at a concentration of 2?mg/L (Dust-2), the NO₃^–N concentration increased by 3.2 fold, and chlorophyll a (chl a) concentration of nano- and micro-eukaryotic phytoplankton increased, while that of pico-eukaryotic phytoplankton did not change significantly. The microphytoplankton abundance increased but the species number decreased in Dust-2. Community structure of eukaryotic microphytoplankton also changed after dust addition. The abundance of diatoms in Dust-2 (23,072 cells/L) was 4.0 fold of that in the control (5750 cells/L), and 2.4 fold of that in Dust-1 (0.2?mg/L of dust addition) (9425 cells/L) at the 7th day of incubation. Abundance of dinoflagellates decreased in Dust-2, being 42.2% that of the control at the 7th day of the incubation. Effects of dust addition on the growth of phytoplankton differed among the dominant genera: growth of Pseudo-nitzschia and Chaetoceros were promoted while that of Prorocentrum was inhibited, and growth of Thalassiosira, Heterocapsa and Scrippsiella was not influenced significantly. The growth of nano- and pico-eukaryotic phytoplankton was promoted, with the cell abundance in Dust-2 2.4 fold of that in control. Our results indicated that Asian dust-originated aerosol particulates could provide nutrients to the oligotrophic NWPO, increase the marine productivity in the area, and alter the eukaryotic phytoplankton community structure.