Grazer control of nutrient availability in the periphyton

Summary Benthic algal assemblages are regulated by both abiotic (e.g., nutrient) and biotic (e.g., grazing) constraint. The objective of this study was to determine how changes in these two factors affected the structure of an algal assemblage in an ephemeral stream. Coverslips were incubated for 21 days in enclosures containing one of three nutrient environments (ambient, phosphorus-enriched, or phosphorus and nitrogen enriched) and one of four densities of the snail Gonibasis (0, 40, 80, or 120 snails/m²) and examined directly to enumerate the algal assemblage. The effect of grazing on algal biomass was dependent on the nutrient environment. An overstory of diatoms was susceptible to removal by grazing and was not strongly affected by nutrient enrichment. An understory of Stigeoclonium was more resistant to grazing and responded strongly to nutrient enrichment only in the presence of grazers. Snail grazers may mediate nutrient availability to the understory indirectly by removing overlying cells or by direct excretion of nutrients. Multiple interactions occur between benthic herbivores and algae, and, as shown here, some of them are positive and involve modifications of the nutrient environment.     

An experimental investigation of phytoplankton nutrient limitation in two contrasting low arctic lakes

We investigated whether phytoplankton communities in two lakes in SW Greenland were phosphorus or nitrogen limited. The study lakes have contrasting water chemistry (mean conductivities differ ten fold) and are located near Kangerlussuaq, SW Greenland (~67°N, 51°W). A microcosm nutrient enrichment experiment was performed in June 2003 to determine whether nitrate or phosphate addition stimulated phytoplankton growth. Samples were analysed for species composition, biomass, and alkaline phosphatase activity (APA). Initially, both lakes had extremely low total phosphorus but high total nitrogen concentrations and high APA, suggesting that the phytoplankton were phosphorus limited prior to the start of the experiment. The phytoplankton composition and biomass (mainly Ochromonas spp.) responded to phosphate but not to nitrate addition. In both lakes, chlorophyll a increased significantly when phosphate was added. Furthermore, APA was significantly lower in the two lakes when phosphate was added compared to the control and the nitrogen addition treatment. The dominance of mixotrophic phytoplankton and high DOC values suggest that these lakes may be regulated by microbial loop processes.     

Light and nutrient control phytoplankton biomass responses to global change in northern lakes

Global change affects terrestrial loadings of colored dissolved organic carbon (DOC) and nutrients to northern lakes. Still, little is known about how phytoplankton respond to changes in light and nutrient availability across gradients in lake DOC. In this study, we used results from whole‐lake studies in northern Sweden to show that annual mean phytoplankton biomass expressed unimodal curved relationships across lake DOC gradients, peaking at threshold DOC levels of around 11 mg/L. Whole‐lake single nutrient enrichment in selected lakes caused elevated biomass, with most pronounced effect at the threshold DOC level. These patterns give support to the suggested dual control by DOC on phytoplankton via nutrient (positively) and light (negatively) availability and imply that the lakes' location along the DOC axis is critical in determining to what extent phytoplankton respond to changes in DOC and/or nutrient loadings. By using data from the large Swedish Lake Monitoring Survey, we further estimated that 80% of northern Swedish lakes are below the DOC threshold, potentially experiencing increased phytoplankton biomass with browning alone, and/or combined with nutrient enrichment. The results support the previous model results on effects of browning and eutrophication on lake phytoplankton, and provide important understanding of how northern lakes may respond to future global changes.     

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.     

30 Seasonal control of phytoplankton biomass and productivity in coastal british columbia lakes and reservoirs

The algal class Chlorarachniophyceae is comprised of a small group of unicellular eukaryotic algae that are often characterized by an unusual amoeboid morphology. This morphology is hypothesized to be the result of a secondary endosymbiosis in which a green alga was engulfed as prey by a nonphotosynthetic amoeba or amoebaflagellate. Whereas much is known about the phylogenetic relationships of individual chlorarachniophytes to one another, and to possible ancestral host organisms in the genera Cercomonas and Heteromita , little is known about their physiology, particularly that of their lipids. In an initial effort to characterize the lipids of this algal class, seven organisms were examined for their fatty acid and sterol composition. These included Bigelowiella natans, Chlorarachnion globusum, Chlorarachnion reptans, Gymnochlora stellata, Lotharella amoeboformis, Lotharella globosa , and Lotharella sp . Fatty acids associated with chloroplast-associated glycolipids, cytoplasmic membrane-associated phospholipids, and storage triglycerides were characterized. Glycolipid fatty acids were found to be of limited composition, containing principally eicosapentaenoic acid [20:5(n-3)] and hexadecanoic acid (16:0), which ranged in relative percentage from 67–90% and 10–29%, respectively, in these seven organisms. Triglyceride-associated fatty acids were found to be similar. Phospholipid fatty acid composition was more variable. The principal phospholipid fatty acids, 16:0 (25–32%) and a compound tentatively identified as docosapentaenoic acid [22:5(n-3)] (26–35%), were found along with a number of C18 and C20 fatty acids. All organisms contained two sterols as free sterols. These were tentatively identified as 24-ethylcholesta-5,22E-dien-3b-ol (stigmasterol; 70–95%) and 24-methylcholesta-5,22E-dien-3b-ol (brassicasterol; 5–30%).     

Effects of nutrient and light limitation on the biochemical composition of phytoplankton

Three marine phytoplankters (Isochrysis galbana, Chaetoceros calcitrans andThalassiosira pseudonana), commonly used in the culture of bivalve larvae, were grown in batch or semi-continuous cultures. Changes in protein, carbohydrate, lipid and some fatty acids were measured as growth became limited by nitrogen, silicon, phosphorus or light. Under N starvation (2 d) the % lipid remained relatively constant, while% carbohydrate increased and% protein decreased in all 3 species compared to cells growing under no nutrient limitation. Under Si starvation (6 h) there was no change in lipid, protein or carbohydrates. The amount of two fatty acids, 20 : 53 and 22 : 63 remained relatively constant under N, P and Si starvation, exept for a sharp drop in the cells of P-starvedT. pseudonana. However, there were pronounced species differences withI. galbana containing significantly less 20 : 5 3 thanC. calcitrans orT. pseudonana. Under light limitation the amount of lipid per cell showed no consistent trend over a range of irradiances for all 3 species. The amount of N per cell (an index of protein content) as a function of irradiance, was relatively constant forI. galbana andT. pseudonana, while the amount of N per cell was lower under low irradiances forC. calcitrans. These examples of changes in protein, carbohydrate, lipid and certain fatty acids under nutrient (N, Si or P) or light limitation, emphasize the importance of knowing the phase (e.g. logarithmic vs stationary) of the growth curve in batch cultures, since the nutritional value of the phytoplankters could change as cultures become dense and growth is terminated due to nutrient or light limitation.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.     

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。     

灌河口邻近海域春季浮游植物的生态分布及其营养盐限制

2011年4月通过灌河口邻近海域的现场调查及营养加富培养实验,研究了春季灌河口邻近海域浮游植物生态分布特征以及硝酸盐、磷酸盐对浮游植物生长的限制作用,结果表明:共发现浮游植物68种,其中硅藻61种,优势度最高的为中肋骨条藻(Skeletonema costatum,Y=0.53),各个站位浮游植物的丰度介于0.84× 106-2.25×106个/L,均值为1.54×106个/L,种类范围为29-39种,均值为35种,叶绿素a浓度呈现近岸高外海低的特征,在2.66-6.67 μg/L变化,均值为3.89 μg/L,多样性指数介于2.60-3.79,均值为3.20,海域环境基本适宜浮游植物的生长;调查海域磷酸盐浓度的范围为0.35-0.90μmol/L,均值为0.58μmol/L,亚硝酸盐浓度范围为1.57-3.93 μmol/L,均值为3.08 μmol/L,两者分布均具有近岸高外海低的特征;铵盐浓度范围为3.145.43μmol/L,均值为3.95 μmol/L,其分布则是近岸低外海高;硝酸盐浓度严重偏高,在31.21-37.00μmol/L之间变化,均值为34.55 μmol/L,导致调查区域具有高N/P比(42-112),且浮游植物叶绿素a与磷酸盐浓度有显著的正相关(R2=0.80),而与无机氮线性关系不明显(R2=0.11);在P加富培养实验中,磷酸盐在3个培养组(对照,+P,++P)中的比吸收速率分别为0.36、0.43、0.51d-1,加P促进了P本身的吸收,硝酸盐和亚硝酸盐的吸收也得以促进,但没有磷酸盐那么显著,而铵盐浓度基本呈增加趋势,P的添加也促进了藻类的生长,培养结束后叶绿素a浓度最大值分别为77.24、90.57、96.49μg/L.在N加富培养实验中,硝酸盐的比吸收速率分别为0.39、0.049、0.025d-1,加N未促进硝酸盐本身的吸收,磷酸盐浓度在3个实验组变化曲线相似,其吸收也没有得到促进,亚硝酸盐在加N组中浓度是增加的,培养结速后加N组(+N,++N)叶绿素a浓度最大值分别为72.31、69.62μg/L,都小于对照组,N的添加也未促进藻类的生长.上述研究表明了春季灌河口邻近海域浮游植物的生长主要受到P的限制,而不是N限制.     

Spatial and temporal variation of phytoplankton in two subtropical Brazilian reservoirs

The purpose of this study was to verify the longitudinal distribution of phytoplankton biomass in two subtropical Brazilian reservoirs in the State of Paraná and investigate intervening factors on changes in phytoplankton biomass according to functional groups. In the Capivari and Segredo reservoirs, samples were obtained every 3 months during 2002, along a longitudinal axis (fluvial, transition, and lacustrine zones) at different depths. One hundred and eighteen taxa were identified, with Chlorophyceae as the most specious group. During the study period, both reservoirs had mostly low biomass values (less than 1 mm³ l⁻¹). The short retention time of these reservoirs constituted the principal limiting factor to phytoplankton development. Biomass values above 1 mm³ l⁻¹ were observed in the Capivari fluvial zone in March and in the Segredo lacustrine zone in December, with dominance by Microcystis aeruginosa Kütz (L_M) and Anabaena circinalis Rab. (H₁), respectively. Vertical and horizontal gradients of analyzed abiotic variables and phytoplankton biomass were observed. Considering the phytoplankton biomass values, both reservoirs were oligotrophic for the duration of the study. The Canonical Correspondence Analysis (CCA) evidenced temporal and spatial gradients of phytoplankton biomass; nevertheless, it did not follow the classic model proposed for deep reservoirs, since higher biomass was registered in the lacustrine zone during some months and in fluvial zones during other months. Distinct functional groups of phytoplankton characterized both studied reservoirs. Capivari Reservoir was best characterized by L_M and Y groups, indicative of its greater water column stability and higher phosphorus concentration, whereas Segredo Reservoir was principally characterized by the MP functional group, indicative of its greater mixing zone extension and higher nitrate concentration. The obtained results also evidenced the influence of morphometric conditions and watershed purposes as important structuring factors of phytoplankton biomass in these reservoirs. Handling editor: L. Naselli-Flores     

Defenses in phytoplankton against grazing induced by nutrient limitation, UV-B stress and infochemicals

It is becoming increasingly evident that the efficiency of zooplankton grazing on algae is not only a matter of quantity of the grazer relative to its food. Planktonic primary producers are not defenseless food-particles that are easily harvested by the consumers. Several algal species are able to adjust their phenotype (colony formation, spines, size) in such a way that it results in a reduced grazing pressure. It was recently demonstrated that morphological changes in the cell wall of green algae, induced by nutrient limitation and UV-B stress, may reduce their digestibility. A high fraction of induced cells pass intact and viable through the gut of the zooplankters, such that the grazing impact on the population is strongly reduced. It was also found that the presence of exudates (infochemicals) released by daphnids may change the morphology of algae. Unicellular green algae of the genus Scenedesmus were induced to form eight-cell coenobial types, heavily armed with spines, within three to five days after adding filtered water from an algal culture with Daphnia present. Both defence mechanisms may play an important role in zooplankton production and competition, and may serve as an example of highly efficient strategies to resist heavy grazing pressure.     

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.     

Nitrogen deposition and warming – effects on phytoplankton nutrient limitation in subarctic lakes

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N‐deposition (<1 kg N ha^?1 yr^?1) in northern subarctic Sweden. Short‐term bioassay experiments with N‐ and P‐additions revealed that phytoplankton in high‐alpine lakes were more prone to P‐limitation, and with decreasing altitude became increasingly N‐ and NP‐colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N‐deposition over the Swedish mountain areas will, with the exception of high‐alpine lakes, enhance biomass and drive phytoplankton from N‐ to P‐limitation. However, if not accompanied by warming, N‐input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.     

Nutrient regeneration by zooplankton: effects on nutrient limitation of phytoplankton in a eutrophic lake

We tested the hypothesis that excretion of nutrients by zooplanktoncan reduce the severity of nutrient limitation of phytoplankton,and determine whether the phytoplankton community is limitedby nitrogen or phosphorus. In situ experiments were conductedin eutrophic Lake Mendota (Wisconsin, USA) during the summerof 1988, where phytoplankton were limited by N and P, but periodsof nutrient limitation were transitory Increased zooplanktonbiomass and the consequent increased excretion of nutrientsby zooplankton reduced P limitation (as measured by specificalkaline phosphatase activity) in all experiments Excretionof nutrients also reduced N limitation (as measured by ammoniumenhancement response) in one of three experiments. In additionalexperiments in the more highly eutrophic Lake Wingra, excretionof nutrients by zooplankton reduced both N and P limitationThese results support the hypothesis that zooplankton have potentiallyimportant indirect effects on phytoplankton communities throughrecycling of nutrients     

The relative importance of light and nutrient limitation of phytoplankton growth: a simple index of coastal ecosystem sensitivity to nutrient enrichment

Anthropogenic nutrient enrichment of the coastal zone is now a well-established fact. However, there is still uncertainty about the mechanisms through which nutrient enrichment can disrupt biological communities and ecosystem processes in the coastal zone. For example, while some estuaries exhibit classic symptoms of acute eutrophication, including enhanced production of algal biomass, other nutrient-rich estuaries maintain low algal biomass and primary production. This implies that large differences exist among coastal ecosystems in the rates and patterns of nutrient assimilation and cycling. Part of this variability comes from differences among ecosystems in the other resource that can limit algal growth and production – the light energy required for photosynthesis. Complete understanding of the eutrophication process requires consideration of the interacting effects of light and nutrients, including the role of light availability as a regulator of the expression of eutrophication. A simple index of the relative strength of light and nutrient limitation of algal growth can be derived from models that describe growth rate as a function of these resources. This index can then be used as one diagnostic to classify the sensitivity of coastal ecosystems to the harmful effects of eutrophication. Here I illustrate the application of this diagnostic with light and nutrient measurements made in three California estuaries and two Dutch estuaries.     

Basin-scale control on the phytoplankton biomass in Lake Victoria, Africa

The relative bio-optical variability within Lake Victoria was analyzed through the spatio-temporal decomposition of a 1997–2004 dataset of remotely-sensed reflectance ratios in the visible spectral range. Results show a regular seasonal pattern with a phase shift (around 2 months) between the south and north parts of the lake. Interannual trends suggested a teleconnection between the lake dynamics and El-Niño phenomena. Both seasonal and interannual patterns were associated to conditions of light limitation for phytoplankton growth and basin-scale hydrodynamics on phytoplankton access to light. Phytoplankton blooms developed during the periods of lake surface warming and water column stability. The temporal shift apparent in the bio-optical seasonal cycles was related to the differential cooling of the lake surface by southeastern monsoon winds. North-south differences in the exposure to trade winds are supported by the orography of the Eastern Great Rift Valley. The result is that surface layer warming begins in the northern part of the lake while the formation of cool and dense water continues in the southern part. The resulting buoyancy field is sufficient to induce a lake-wide convective circulation and the tilting of the isotherms along the north-south axis. Once surface warming spreads over the whole lake, the phytoplankton bloom dynamics are subjected to the internal seiche derived from the relaxation of thermocline tilting. In 1997–98, El-Niño phenomenon weakened the monsoon wind flow which led to an increase in water column stability and a higher phytoplankton optical signal throughout the lake. This suggests that phytoplankton response to expected climate scenarios will be opposite to that proposed for nutrient-limited great lakes. The present analysis of remotely-sensed bio-optical properties in combination with environmental data provides a novel basin-scale framework for research and management strategies in Lake Victoria.     

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.     

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.