Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

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.     

Phytoplankton nutrient limitation in Colorado mountain lakes

SUMMARY. 1. Limiting nutrients for phytoplankton were studied experimentally in eight mountain lakes of central Colorado between May and November of 1984.
2. Five categories of phytoplankton limitation were identified: no limitation, N limitation, P limitation, concurrent limitation (stimulation only by simultaneous additions of N and P), and reciprocal limitation (stimulation by addition of either N or P). The phytoplankton communities of three lakes were primarily N-limited, one was primarily phosphorus-limited, and four showed primarily combined limitation (concurrent or reciprocal). Switching between categories of limitation was also observed within lakes. Nitrogen was the most frequently limiting nutrient; N, either alone or in combination with P, accounted for 79% of all observed instances of limitation.
3. Nine indices were tested for effectiveness in predicting phytoplankton limitation by N and P. The best indices for discriminating all limitations were ratios of dissolved inorganic N: total P (84% accuracy) and dissolved inorganic N:total dissolved P (80% accuracy). The effectiveness of these indices may be explained by the degree to which they represent N and P fractions actually available to the phytoplankton.     

Responses of epilimnetic phytoplankton to experimental nutrient enrichment in three small seepage lakes

This paper describes the responses of three epilimnetic phytoplanktoncommunities to experimental nitrogen and phosphorus enrichmentas compared to the phytoplankton community in a fourth, unmanipulated,lake. Increased nutrient inputs increased total phytoplanktonbiomass, primary productivity, chlorophytes, cryptomonads andspecies turnover rates in all three enriched lakes; cyanobacteriaincreased in two of the three enriched lakes. However, nutrientaddition also led to declines in previously dominant dinoflagellatesand chrysophytes, and in species diversity. At the species level,there were large changes in community composition from yearto year in both enriched and reference lakes, suggesting thatphytoplankton community composition is highly dynamic even inthe absence of enrichment. Overall, changes in total biomass,productivity and species diversity were consistent among theenriched lakes, while changes in species composition differeddue to variation in the physical, chemical and biotic environmentof each lake. This suggests that aggregated variates are moreuseful for quantitative prediction of nutrient effects, whilespecies responses can be used to signal qualitative differencesin environmental conditions among lakes. ³Present address: Department of Biological Sciences, DartmouthCollege, 6044 Gilman Laboratory, Hanover, NH 03755-3576, USA     

Disturbance-diversity relationships in two lakes of similar nutrient chemistry but contrasting disturbance regimes

Phytoplankton diversity was studied in two North German lakes of comparable nutrient chemistry but different exposure to winds. In both lakes, phytoplankton was primarily N-limited but diatoms were Si-limited. Plußsee had a very constant mixing depth during summer, while week-to-week changes of several meters were quite common in the more exposed Behler See. In Plußsee, phytoplankton biomass during summer came closer to the carrying capacity as defined by the available total N. In Plußsee there was a marked decline of diversity during the summer maximum of biomass, while this decline was less pronounced in Behler See. It is concluded that disturbances which prevented phytoplankton from reaching the carrying capacity also maintained a high level of diversity. A negative response of diversity to undisturbed conditions became apparent, after phytoplankton biomass had exceeded about 5% of the carrying capacity.     

Driving forces shaping phytoplankton assemblages in two subtropical plateau lakes with contrasting trophic status

1. We studied driving forces shaping phytoplankton assemblages in two subtropical plateau lakes with contrasting trophic status, the oligotrophic deep Lake Fuxian and the eutrophic shallow Lake Xingyun. 2. Phytoplankton samples were taken monthly for a year and phytoplankton species were sorted into the main taxonomic groups and associations proposed by Reynolds. A canonical correspondence analysis (CCA) was used to test the occurrence of these classification schemes and to determine their discriminatory power. 3. The results suggest that the major driving forces in Lake Fuxian were physical variables, and particularly the underwater light climate, whereas, nutrients were the important driving force in Lake Xingyun. 4. Top–down control through zooplankton grazing in Lake Fuxian was hardly ever a significant determinant itself, because of the scarcity of zooplankton and their low grazing efficiency of predation while a dominance of inedible cyanobacteria throughout the year rendered top–down controls ineffective failing in Lake Xingyun. Hence phytoplankton communities in both lakes appear to be regulated primarily by bottom–up controls.     

Nutrient limitation of periphyton growth in arctic lakes in south-west Greenland

Many arctic lakes are oligotrophic systems where phototrophic growth is controlled by nutrient supply. Recent anthropogenic nutrient loading is associated with biological and/or physico-chemical change in several lakes across the arctic. Shifts in nutrient limitation (nitrogen (N), phosphorus (P), or N + P) and associated effects on the growth and composition of algal communities are commonly reported. The Kangerlussuaq region of south-west Greenland forms a major lake district which is considered to receive little direct anthropogenic disturbance. However, long-range transport of pollutant N is now reaching Greenland, and it was hypothesised that a precipitation gradient from the inland ice sheet margin to the coast might also deliver increased N deposition. In situ nutrient bioassays were deployed in three lakes across the region: ice sheet margin, inland (close to Kangerlussuaq) and the coast (near Sisimiut), to determine nutrient limitation of lakes and investigate any effects of nutrients on periphyton growth and community composition. Nutrient limitation differed amongst lakes: N limitation (ice sheet margin), N and P limitation (inland) and N + P co-limitation (coast). Factors including variation in N supply, ice phenology, seasonal algal succession, community structure and physical limnology are explored as mechanisms to explain differences amongst lakes. Nutrient limitation of arctic lakes and associated ecological impacts are highly variable, even across small geographic areas. In this highly sensitive region, future environmental change scenarios carry a strong risk of significantly altering nutrient limitation; in turn, potentially severely impacting lake structure and function.     

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.     

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.     

Size fractionated phytoplankton production in two humic shallow lakes with contrasting coverage of free floating plants

We conducted a 1-year survey in two humic shallow lakes from the floodplain of the Lower Paraná River, Laguna Grande Lake (LGL) and a relictual oxbow lake (ROL). We aimed to test two hypotheses: (1) the efficiency in light use of picoplankton (0.2–3 μm) is greater as light restriction increases and (2) the contribution of picoplankton to the total productivity is higher when the total photosynthetic biomass is lower. We performed P–E curves for picoplankton and nano- and microplankton (>3 μm) using the ¹⁴C assimilation technique. The light environments of the water bodies differed mainly owing to the development of free floating plants on the surface of the ROL and the dominance of phytoplankton in LGL. Primary productivity patterns in LGL were seasonality driven whilst in the ROL they were related to the coverage of floating macrophytes, which promoted light limitation and a lower productivity. In LGL, nano- and microplankton were in general more productive and the relative contribution of picoplankton to the total phytoplankton production decreased with the increase in total photosynthetic biomass. Hence, our study extends previously observed patterns to subtropical shallow lakes, where seasonality and free floating plants may influence the dynamics of phytoplankton production.     

Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Physiological indicators of nutrient deficiency in phytoplankton in southern Chilean lakes

We assessed the nutrient status of phytoplankton in 28 lakes in southern Chile using two types of physiological indicators: specific alkaline phosphatase activity, and the elemental composition (carbon, nitrogen, and phosphorus) of seston. Alkaline phosphatase activity ranged from 0.001 to 0.11 mol P g chl^–1 h^–1, with P-deficiency indicated in about one-half the study lakes. C:N ranged from 3.9 to 24, C:P ranged from 86 to 919, and N:P ranged from 8.7 to 99. C:P and N:P ratios greater than the Redfield ratio were common, suggesting P deficiency in many of the lakes. C:N ratios were not generally indicative of N deficiency. Previous studies have suggested N may be the primary limiting nutrient in southern Chilean lakes, but our results indicate that P should not be discounted as a limiting nutrient.     

Evidence for extensive but variable nutrient limitation in New Zealand lakes

Nutrient limitation causes reduced growth of organisms, which can translate into far-reaching consequences for populations, communities, and ecosystems. Phosphorus (P) limitation, in particular, is associated with reductions in organismal growth because ribosomes, upon which growth depends, require abundant phosphorus to be produced. Chromosomes are also relatively rich in P, meaning that organisms with relatively high chromosome complements (e.g. polyploids) might be especially dependent on abundant environmental P. Here we address the likelihood of nutrient limitation in multiple populations of Potamopyrgus antipodarum, a New Zealand freshwater snail featuring wide ploidy variation. We found that some form of P limitation is very likely in many, but not all, populations of this snail that we surveyed. We also detected extensive across-population variation in P and nitrogen (N) content and N and P limitation and co-limitation in the algae that P. antipodarum eat. Accordingly, we then experimentally evaluated how P and N alone and together influenced growth rate in P. antipodarum. We found that response to nutrients differed by lineage and that dietary P content was more important than dietary N content as a determinant of growth rate, a trait closely tied to fitness in P. antipodarum. The widespread likelihood of (1) P limitation and (2) variation in dietary P availability across New Zealand lakes, along with (3), evidence for lineage-level variation in sensitivity to P limitation, sets the stage for the possibility that variation in nutrient availability contributes to the distribution and maintenance of ploidy variation in P. antipodarum.     

Primary productivity,phytoplankton and limiting nutrient factors in labrador lakes

The primary productivity of some lakes and reservoirs in western Labrador was measured by the ¹⁴C method in order to determine the range of productivities and the effects of impoundment. No primary productivity data previously existed for this part of Canada. Both the primary productivity and standing crops of phytoplankton were found to be low in a newly impounded lake but later rose to levels greater than in surrounding natural lakes. In nutrient enrichment experiments, carbon was never found to be limiting but phosphorus stimulated primary productivity when added alone or in combination with nitrogen.     

Priming and microbial nutrient limitation in lowland tropical forest soils of contrasting fertility

Priming is an increase in soil organic carbon decomposition following input of labile organic carbon. In temperate soils where biological activity is limited commonly by nitrogen availability, priming is expected to occur through microbial acquisition of nitrogen from organic matter or stimulated activity of recalcitrant-carbon degrading microorganisms. However, these priming mechanisms have not yet been assessed in strongly weathered tropical forest soils where biological activity is often limited by the availability of phosphorus. We examined whether microbial nutrient limitation or community dynamics drive priming in three lowland tropical forest soils of contrasting fertility (‘low’, ‘mid’ and ‘high’) by applying C₄-sucrose (alone or in combination with nutrients; nitrogen, phosphorus and potassium) and measuring (1) the δ¹³C-signatures in respired CO₂ and in phospholipid fatty acid (PLFA) biomarkers, and (2) the activities of enzymes involved in nitrogen (N-acetyl β-glucosaminidase), phosphorus (phosphomonoesterase) and carbon (β-glucosidase, cellobiohydrolase, xylanase, phenol oxidase) acquisition from organic compounds. Priming was constrained in part by nutrient availability, because priming was greater when sucrose was added alone compared to when added with nutrients. However, the greatest priming with sucrose addition alone was detected in the medium fertility soil. Priming occurred in parallel with stimulated activity of phosphomonoesterase and phenol oxidase (but not N-acetyl β-glucosaminidase); when sucrose was added with nutrients there were lower activities of phosphomonoesterase and phenol oxidase. There was no evidence according to PLFA δ¹³C-incorporation that priming was caused by specific groups of recalcitrant-carbon degrading microorganisms. We conclude that priming occurred in the intermediate fertility soil following microbial mineralization of organic nutrients (phosphorus in particular) and suggest that priming was constrained in the high fertility soil by high nutrient availability and in the low fertility soil by the low concentration of soil organic matter amenable to priming. This first study of priming mechanisms in tropical forest soils indicates that input of labile carbon can result in priming by microbial mineralization of organic nutrients, which has important implications for understanding the fate of organic carbon in tropical forest soils.     

Linking nutrient limitation and water chemistry in upland lakes to catchment characteristics

Overbank sedimentation on river floodplains can result in significant reduction of the suspended sediment load transported by a river and can thus represent an important component of the catchment sediment budget. Such conveyance losses will also exert an important influence on sediment-associated contaminant fluxes and budgets. This contribution reports the results of a study of sediment-associated contaminants (i.e. total-P, Cr, Cu, Pb and Zn) fluxes in the River Swale (1346 km²) and River Aire (1002 km²) in Yorkshire, U.K., aimed at quantifying the role of overbank floodplain sedimentation in the sediment-associated contaminant budgets. The catchment of the River Aire is dominated by urban and industrial land use in its middle and lower reaches, whereas the River Swale drains a largely rural catchment, although there is a legacy of metal mining in its headwaters which impacts on heavy metal transport by the river. The results for the River Swale indicate that the conveyance losses associated with the deposition of sediment-associated contaminants on the floodplains bordering the main river can be as high as 47% of the total flux through the main channel system. Equivalent values for the River Aire range up to 26%. Contrasts between the two rivers reflect both the location of the contaminant sources within the catchments and the relative magnitude of the fine sediment deposition fluxes associated with their floodplains.     

The light : nutrient ratio in lakes: a test of hypothesized trends in bacterial nutrient limitation

The light-to-nutrient hypothesis explores how the balance between energy (as light energy) and nutrients (as total phosphorus) shapes aquatic ecosystem structure and process. The balance of energy and nutrients is thought to regulate ecosystem structure and process such that, in a "high" light-to-nutrient environment, bacteria would probably be driven towards phosphorus (P) limitation, whereas, in a "low" light-to-nutrient environment, bacteria would be driven towards carbon (C) limitation. We assessed the growth limitation of bacteria in two reservoirs of the southern U.S.A. using a mortality-corrected dilution-growth approach. We compared the frequency of P and C growth limitation with the intralake variation in the light-to-nutrient environment. As a metric of the light-to-nutrient environment, we used the ratio of the mean light in the surface mixed layer ( I _m) to the total phosphorus concentration ( TP ). In each lake, bacterial growth was more often P-limited when the I _m :  TP ratio was above the median ratio than below. We believe our data provide the first evidence supporting this aspect of the light-to-nutrient hypothesis.     

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。     

Primary production studies in two linked but contrasting Welsh lakes

SUMMARY. Llyn Padarn and Llyn Peris have distinct phytoplankton populations. During 1975–76, the standing crop measured as chlorophyll- a was 5.5 times greater in Padarn than in Peris and the production rate, determined by the ¹⁴CO₂ method, was faster by 3.4 times. These differences were attributed to the higher concentrations of phosphorus in the lower lake caused by treated sewage effluent. Incident light intensity, which was slightly lower in Peris due to mountain shading, and temperature, which was 1–4°C higher in Padarn, made little significant contribution to these differences during the summer. The reduced transparency of Padarn water, compared with that of Peris, resulted from denser phytopiankton crops in Padarn. During the summer, Padarn exhibited carbon dioxide depletion which correlated with the chlorophyll concentration. Light inhibition at the surfaces of both lakes correlated with solar radiation intensity. However, the relationship between pigment content and maximum photosynthetic rate was poor. Extracellular products accounted for about 16% of the total production in the lakes. Uptake of ¹⁴C-labelled acetate was low compared with that of ¹⁴CO₂ uptake.