The Effects of Ultraviolet Radiation and Nutrient Additions on Periphyton Biomass and Composition in a Sub‐Alpine Lake (Castle Lake,USA)

Rising levels of ultraviolet radiation (UVR) striking the Earth's surface have led to numerous studies assessing its inhibitory effects on phytoplankton and periphyton in aquatic systems. Mineral nutrients such as nitrogen (N) and phosphorus (P) have been shown to increase aspects of algal metabolism and compensate for UVR inhibition. An in situ substratum enrichment technique and UV shielding was used to assess the effects of nutrient additions on periphyton exposed to different levels of UVR in Castle Lake, California during July‐August, 1997. UV shielding had no effect on total periphyton biomass, but caused shifts in species composition. The dominant periphyton species, Anabaena circinalis RAB., demonstrated sensitivity to ambient levels of UV radiation possibly due to UV inhibition of N2 ‐fixation. Total diatom biovolume decreased when shielded from UVR. Phosphorus additions continually elicited an increase in periphyton biovolume at all levels of analysis. These results suggest an interaction between nutrient status/availability and UV sensitivity.     

Effects of desiccation duration on the community structure and nutrient retention of short and long-hydroperiod Everglades periphyton mats

Responses of periphyton communities to different relevant durations of dry down were assessed. Long-hydroperiod sites within Everglades National Park remain wet for greater than 8 months of the year while short-hydroperiod mats are wet for fewer than 4 months of the year. Dry down duration of long and short-hydroperiod Everglades periphyton was manipulated from 0 to 1, 3, or 8 months after which periphyton was rewetted 1 month and examined for algal species composition. The effects of desiccation and rewetting on periphyton nutrient retention were also assessed. Relative abundance of diatoms declined from an average of 47% in the long-hydroperiod community at the start of the experiment to 24% after 1 month of desiccation and only 12% after 8 months of desiccation. Short-hydroperiod periphyton contained a lower proportion of diatoms at the outset (3%), which declined to less than 1% after the 8-month desiccation treatment. A significant increase in the filamentous cyanobacteria Schizothrix calcicola occurred in long-hydroperiod periphyton mats during this same period, but not in short-hydroperiod mats. Long-hydroperiod periphyton communities had a greater response to desiccation overall, but short-hydroperiod community structure responded to desiccation more rapidly. Because short-hydroperiod communities dry frequently, they appear to cope better to desiccating conditions than long-hydroperiod periphyton communities. This is indicated by the dominance of desiccation resistant algal taxa such as the cyanobacterial filaments S. calcicola and Scytonema hofmanni. Long-hydroperiod periphyton mat communities converge compositionally to short-hydroperiod periphyton communities after prolonged desiccation. Desiccation and rewetting caused long-hydroperiod periphyton to flux greater concentrations of nutrients than short-hydroperiod periphyton. Significant increases in efflux occurred from 1 to 8 months for total phosphorus (TP) and from 1 to 3 and 8 months for total nitrogen (TN) and total organic carbon (TOC). Thus, changes in periphyton mat community structure and function with altered hydroperiod may have long-term ecosystem effects.     

Linking limitation to species composition: importance of inter- and intra-specific variation in grazing resistance

Short-term responses of producers highlight that key nutrients (e.g., N, P)—or combinations of these nutrients—limit primary production in aquatic and terrestrial ecosystems. These discoveries continue to provide highly valuable insights, but it remains important to ask whether nutrients always predominantly limit producers despite wide variation in nutrient supply and herbivory among systems. After all, predictions from simple food chain models (derived here) readily predict that limitation by grazers can exceed that by nutrients, given sufficient enrichment. However, shifts in composition of producers and/or increasing dominance of invulnerable stages of a producer can, in theory, reduce grazer limitation and retain primacy of nutrient limitation along nutrient supply gradients. We observed both mechanisms (inter- and intra-species variation in vulnerability to herbivory) working in a two-part mesocosm experiment. We incubated diverse benthic algal assemblages for several months either in the presence or absence of benthic macro-grazers in mesocosms that spread a broad range of nutrient supply. We then conducted short-term assays of nutrient and grazer limitation on these communities. In the “historically grazed” assemblages, we found shifts from more edible, better competitors to more resistant producers over enrichment gradients (as anticipated by the food web model built with a tradeoff in resistance vs. competitive abilities). However, contrary to our expectations, “historically ungrazed” assemblages became dominated by producers with vulnerable juvenile forms but inedible adult forms (long filaments). Consequently, we observed higher resource limitation rather than grazer limitation over this nutrient supply gradient in both “historically grazed” (expected) and “historically ungrazed” (not initially expected). Thus, via multiple, general mechanisms involving resistance to grazing (changes in species composition or variation in stage-structured vulnerability), producer assemblages should remain more strongly or as strongly limited by nutrients than grazers, even over large enrichment gradients. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecosystem-phase interactions: aquatic eutrophication decreases terrestrial plant diversity in California vernal pools

Eutrophication has long been known to negatively affect aquatic and terrestrial ecosystems worldwide. In freshwater ecosystems, excessive nutrient input results in a shift from vascular plant dominance to algal dominance, while the nutrient-species richness relationship is found to be unimodal. Eutrophication studies are usually conducted in continuously aquatic or terrestrial habitats, but it is unclear how these patterns may be altered by temporal heterogeneity driven by precipitation and temperature variation. The California vernal pool (CVP) ecosystem consists of three distinct phases (aquatic, terrestrial, and dry) caused by variation in climatic conditions. The purpose of this study was to test the hypothesis that resource addition during the aquatic phase results in increased algal abundance, which reduces vascular plant cover and richness of the terrestrial phase upon desiccation. We used mesocosms layered with CVP soil, in which treatments consisted of five levels of nitrogen and phosphorous added every 2 weeks. Resource addition increased available phosphorus levels and algae cover during the aquatic phase. Increased algal crusts resulted in decreased vascular plant percent cover and species richness. Few significant patterns were observed with individual plant species and total biomass. The phosphorus-plant richness relationship was not significant, but species composition was significantly different among the low and high treatment comparisons. These results highlight a neglected effect of eutrophication in seasonal habitats. Interactions among ecosystem phases clearly require more attention empirically and theoretically. Management and restoration of temporally heterogeneous habitat, such as the endemic-rich CVP, need to consider the extensive effects of increased nutrient input.     

养分资源脉冲供给对几种微藻种间竞争的影响

以淡水生态系统常见的5种微藻为研究对象,通过稳态条件下单一培养的方式获取各微藻在氮素或磷素缺乏条件下对应的生长特征参数和R*值,同时将5种微藻在养分资源脉冲供给的方式下进行混合培养,进而检测养分资源脉冲供给对几种微藻种间竞争的影响作用,并与基于稳态条件下所预测的微藻种间竞争结果进行比较。研究结果显示,在稳态条件下具有最小R*值的纤细角星鼓藻在与其它微藻的种间竞争过程中始终保持优势地位,而其余4种微藻在混合培养状态下的相对比重亦与其稳态条件下的R*值紧密相关。然而,资源脉冲供给条件下的竞争优势种纤细角星鼓藻能与其它种群数量较小的微藻共存。此外,在氮素和磷素的不同供给比例情况下,对应微藻群落的结构也会发生相应的变化。实验养分资源脉冲作用下多种微藻的共存现象与自然水体中的观测现象相一致,显示了资源脉冲可能是维持生物多样性水平的一个重要机制。     

The effects of variable nutrient additions on a pond mesocosm community

The effects of nutrient additions on aquatic systems have been frequently studied. Typically, these studies report an increase in algal biomass and a decrease in species diversity in response to nutrient increases. However, it is not clear why comparable aquatic communities respond differently to nutrient additions of similar magnitudes. We tested the effects of the rate and amount of nutrient load on community structure in 760 l mesocosms; treatments manipulated the total amount of nutrients that entered an aquatic system (small versus large load) and the temporal pattern in which these nutrients entered the system (annually, monthly, or weekly). We found that the effects of the loading rate of nutrients were at least as important as the total amount of the nutrients for several response variables. Although these effects were manifested in several ways, the response to the different rates was most prominent within groups of the primary producers, which showed large shifts in composition and abundance. Handling editor: L. M. Bini     

Phytoplankton population dynamics in perennially ice-covered Lake Fryxell, Antarctica

Phytoplankton were collected over five austral summers (1987–88through 1991–92) to examine seasonal and annual fluctuationsin species composition and biovolume in Lake Fryxell, a perenniallyice-covered lake located in the Dry Valleys of South VictoriaLand, Antarctica. Lake Fryxell has perennial gradients of salinity,dissolved oxygen and nutrients. We found that algal speciesdiversity was low (56 taxa were collected), confirming the resultsof previous short-term studies. The phytoplankton consistedprimarily of cryptophyte and chlorophyte flagellates, and filamentouscyanobacteria. The presence of filamentous cyanobacteria, whichhave not been reported as abundant in this lake by previousworkers, may represent a significant ecological change. Eachaustral summer, one dominant species contributed >70% ofthe total biovolume; Chroomonas lacustris was dominant in 1987–88,while Cryptomonas sp. dominated the phytoplankton in the remaining4 years. No species succession occurred during the austral summer.Some common taxa were vertically stratified (Oscillatoria limnetica,Phormidium angustissimum, Pyramimonas sp., Oscillatoria sp.),while others showed no distinct vertical stratification (Chlamydomonassubcaudata, Cryptomonas sp.). The stratification of the phytoplanktonreflects the gradients of nutrients and light, and the stabilityof the water column.     

Microscale measurements reveal contrasting effects of photosynthesis and epiphytes on frictional drag on the surfaces of filamentous algae

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Emergence of nutrient co‐limitation through movement in stoichiometric meta‐ecosystems

Evidence that ecosystems and primary producers are limited in their productivity by multiple nutrients has caused the traditional nutrient limitation framework to include multiple limiting nutrients. The models built to mimic these responses have invoked local mechanisms at the level of the primary producers. In this paper, we explore an alternative explanation for the emergence of co‐limitation by developing a simple, stoichiometrically explicit meta‐ecosystem model with two limiting nutrients, autotrophs and herbivores. Our results show that differences in movement rates for the nutrients, autotrophs and herbivores can allow for nutrient co‐limitation in biomass response to emerge despite no local mechanisms of nutrient co‐limitation. Furthermore, our results provide an explanation to why autotrophs show positive growth responses to nutrients despite ‘nominal’ top‐down control by herbivores. These results suggest that spatial processes can be mechanisms for nutrient co‐limitation at local and regional scales, and can help explain anomalous results in the co‐limitation literature.     

Spatial distribution of aquatic marine fungi across the western Arctic and sub‐arctic

Fungi are important parasites of primary producers and nutrient cyclers in aquatic ecosystems. In the Pacific‐Arctic domain, fungal parasitism is linked to light intensities and algal stress that can elevate disease incidence on algae and reduce diatom concentrations. Fungi are vastly understudied in the marine realm and knowledge of their function is constrained by the current understanding of fungal distribution and drivers on global scales. To investigate the spatial distribution of fungi in the western Arctic and sub‐Arctic, we used high throughput methods to sequence 18S rRNA, cloned and sequenced 28S rRNA and microscopically counted chytrid‐infected diatoms. We identified a broad distribution of fungal taxa predominated by Chytridiomycota and Dikarya. Phylogenetic analysis of our Chytridiomycota clones placed Arctic marine fungi sister to the order Lobulomycetales. This clade of fungi predominated in fungal communities under ice with low snowpack. Microscopic examination of fixed seawater and sea ice samples revealed chytrids parasitizing diatoms collected across the Arctic that notably infected 25% of a single diatom species in the Bering Sea. The Pezizomycotina comprised > 95% of eukaryotic sequence reads in Greenland, providing preliminary evidence for osmotrophs being a substitute for algae as the base of food webs.     

Searching for allelopathic effects of submerged macrophytes on phytoplankton—state of the art and open questions

Allelopathy, here defined as biochemical interactions between aquatic primary producers, has always been intriguing as a process explaining the dominance of certain plant or algal species over others. Negative chemical interference has been invoked as one of the steering mechanisms behind mutual dominance of either submerged macrophytes or phytoplankton in shallow eutrophic lakes. Yet, despite much effort, convincing evidence for allelopathic interactions in situ is still missing. Also, laboratory approaches often lack reality. Inspired by a series of talks at the Shallow Lakes 2005 meeting in Dalfsen, the Netherlands, we argue that there is circumstantial but strong evidence that allelopathic interference between submerged macrophytes and phytoplankton may indeed exist in aquatic ecosystems despite the problems associated with research in this field. We first discuss experimental approaches combining laboratory and field studies, based on examples presented at this meeting. We then discuss the impact of nutrient status of both producing and target organism and biotic factors such as herbivory or pathogens that might affect allelopathy. Further topics are the potential seasonality of effects and the species-specificity of certain allelochemicals. We conclude with some thoughts why a final proof for allelopathy in situ might remain difficult or even inaccessible in some cases, and why we nevertheless should not abandon this idea.     

Microcommunities and microgradients: Linking nutrient regeneration,microbial mutualism,and high sustained aquatic primary production

Nutrient regeneration is essential to sustained primary production in the aquatic environment because of coupled physical and metabolic gradients. The commonly evaluated ecosystem perspective of nutrient regeneration, as is illustrated among planktonic paradigms of lake ecosystems, functions only at macrotemporal and spatial scales. Most inland waters are small and shallow. Consequently, most organic matter of these waters is derived from photosynthesis of emergent, floating-leaved, and submersed higher plants and microflora associated with living substrata and detritus, including sediments, as well as terrestrial sources. The dominant primary productivity of inland aquatic ecosystems is not planktonic, but rather is associated with surfaces. The high sustained rates of primary production among sessile communities are possible because of the intensive internal recycling of nutrients, including carbon. Steep gradients exist within these attached microbial communities that (a) require rapid, intensive recycling of carbon, phosphorus, nitrogen, and other nutrients between producers, particulate and dissolved detritus, and bacteria and protists: (b) augment internal community recycling and losses with small external inputs of carbon and nutrients from the overlying water or from the supporting substrata; and (c) encourage maximal conservation of nutrients. Examples of microenvironmental recycling of carbon, phosphorus, and oxygen among epiphytic, epipelic, and epilithic communities are explained. Recalcitrant dissolved organic compounds from decomposition can serve both as carbon and energy substrates as well as be selectively inhibitory to microbial metabolism and nutrient recycling. Rapid recycling of nutrient and organic carbon within micro-environments operates at all levels, planktonic as well as attached, and is mandatory for high sustained productivity.     

Control of phytoplankton–bacteria interactions by stoichiometric constraints

In aquatic ecosystems, phytoplanktonic organisms are the major primary producers and bacteria the major decomposers. The interactions between phytoplankton and bacteria may be dependent on nutrient resources. Anthropogenic inputs, by modifying nutrient status and stoichiometry of lakes, might induce changes in these interactions, and thus, could have many consequences on some ecological processes such as primary production or importance of microbial recycling activity.
To test this hypothesis, we grew an axenic strain of a green alga, Scenedesmus obliquus , in a range of stoichiometric situations, in absence and in presence of a natural bacterial community. Here, we show that different phytoplankton limiting factors can generate between algae and bacteria either competition for nutrients in phosphorus-limited conditions, commensalism in nitrogen-limited conditions, or mutualism in eutrophic nutrient-unlimited conditions. Causes of these different interaction types are discussed, in particular the hypothesis that in very eutrophic systems with high primary production, mutualism between algae and bacteria could be due to CO₂ supply by heterotrophic respiration to inorganic carbon limited algae. Some probable consequences for aquatic ecosystems functioning are proposed.     

Determination of tipping points for aquatic plants and water quality parameters in fish pond systems: A multi-year approach

High levels of nutrients in fish ponds by fish farming may cause significant eutrophication leading to a loss in species richness and a decrease of cover of aquatic plants to phytoplankton dominance. This shift can be represented by a tipping point where a significant change in the state of the ecosystem is observed such as a change from high to low aquatic plants species richness and cover. A total of 100 fish ponds were studied during five years in the Dombes region, France, to determine tipping points in aquatic plant richness and cover using chlorophyll α (CHL), water transparency, Total N (TN) and Total P (TP) gradients with two statistical methods. The relationships between tipping points, nutrient loads and yearly variations in weather conditions were also evaluated. Looking at the five years data, tipping points were observed in aquatic plant richness at 6 and 60 μg/L for CHL, and at 3.90 mg/L for TN concentration; as well as at 70 cm for water transparency, but no tipping point was found with TP. For aquatic plant cover, tipping points were observed at 11 μg/L for CHL, 2.42 mg/L for TN, 0.05 mg/L for TP, and at 62 cm for water transparency. These tipping points showed a significant decrease of aquatic plant species richness and cover, linked to the nutrient concentrations which drive the competition between the primary producers phytoplankton and aquatic plants. However, tipping points could vary significantly between years. The inter-annual variability may be due to an early occurrence of phytoplankton blooms in some ponds in a year preventing the establishment of aquatic plants, and thus influencing the value of tipping points. Weather conditions influence the competition between primary producers by impacting chlorophyll α and nutrients concentrations. When weather conditions supported increased nutrient concentrations, the development of phytoplankton and aquatic plants was facilitated and tipping points in aquatic plant richness and cover occurred with relatively high values. Thus, a significant decrease of plant cover and richness occurred at higher level of nutrients compared to the other years. In these cases, aquatic plants dominated over phytoplankton for the spring period, and also often during summer. In conclusion, tipping points observed are mainly linked to the competition between aquatic plants and phytoplankton. In shallow and eutrophic systems like fish ponds where nutrients are not a limiting resource, weather conditions act temporarily during spring as the main regulator of this competition.     

In situ microcosm experiments on the influence of nitrate and light on phytoplankton community composition

Phytoplankton blooms are fundamental features of coastal ecosystems, but the processes that select for blooms of certain species are not well understood. The aim of this work was to investigate experimentally the interaction of light and nutrients (nitrate) in structuring phytoplankton community composition in Pelorus Sound, New Zealand. Microcosm experiments were conducted in situ nine times throughout the year, providing controls and treatments for increased nutrients and decreased light. Nitrate availability was found to be limiting to phytoplankton growth during spring and summer. Small- to medium-sized, chain-forming diatom taxa such as Chaetoceros sp., Skeletonema sp., Pseudonitzschia sp. and Thalassiosira sp. responded most rapidly to nitrate enrichment, increasing their biovolume up to 32-fold during the 5-day experiments. A long-term phytoplankton monitoring database showed that these taxa have historically dominated the phytoplankton assemblage, suggesting that intense competition for nitrate is a key component in structuring the phytoplankton community. Many of the taxa that were able to withstand light reduction in the shaded treatments were rare historically in Pelorus Sound, suggesting that light is secondary to nitrate availability in structuring the phytoplankton community composition in this coastal embayment.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

Calanoid copepods are major components of most lacustrine ecosystems and their grazing activities may influence both phytoplankton biomass and species composition. To assess this we conducted four seasonal, in situ, grazing experiments in eutrophic Lake Rotomanuka, New Zealand. Ambient concentrations of late stage copepodites and adults of calanoid copepods (predominantly Calamoecia lucasi, but with small numbers of Boeckella delicata) were allowed to feed for nine days on natural phytoplankton assemblages suspended in the lake within 1160 litre polyethylene enclosures. The copepods reduced the total phytoplankton biomass of the dominant species in all experiments but were most effective in summer (the time of highest grazer biomass) followed by spring and autumn. In response to grazing pressure the density of individual algal species showed either no change or a decline. There were no taxa which increased in density in the presence of the copepods. The calanoid copepods suppressed the smallest phytoplankton species (especially those with GALD (Greatest Axial Linear Dimension) < µm) and there appeared to be no selection of algae on the basis of biovolume. Algal taxa which showed strong declines in abundance in the presence of the copepods include Cyclotella stelligera, Coelastrum spp., Trachelomonas spp., Cryptomonas spp., and Mallomonas akrokomos. Calanoid copepods are considered important grazers of phytoplankton biomass in this lake. The study supports the view that high phytoplankton:zooplankton biomass ratios and large average algal sizes characteristic of New Zealand lake plankton may, at least partly, be caused by year round grazing pressure on small algae shifting the competitive balance in favour of larger algal species.     

Estimating phytoplankton carbon from microscopic counts: an application for riverine systems

Algal biomass, in addition to cell numbers, is a measure of the successful conversion of inorganic to organic carbon. Consequently, carbon is the main currency used in aquatic models and in flux and budget studies. On the other hand, microscopic observation and counts remain the only means for determining species composition and biomass, which is relevant to many aspects of aquatic ecology. In this study, we focus on the way to convert biovolume to carbon biomass for algal assemblages of two rivers, using a computerized system that records dimensions of phytoplankton (Gosselain & Hamilton, 2000). We first compare different equations found in the literature for converting algal cell volume to cellular carbon content. We then evaluate the accuracy of a biomass estimate based on less time-consuming measurements, using pre-determined biovolume values instead of measuring cells in all samples. Biovolume/carbon equations are evaluated using total phytoplankton carbon biomass determined from measured chlorophyll a. Equations established for freshwater taxa seem to provide better estimates of algal biomass in the two case studies presented here, the Rideau and Meuse rivers (Canada and Belgium, respectively) than do more numerous equations defined for marine taxa. Furthermore, equations that make a distinction between diatoms and other algae appear more appropriate than those considering all algal groups as a whole. Finally, mean values of algal biovolumes, determined using sufficient measurements of cell dimensions from representative sampling series, may prove sufficient for carbon estimates of taxa in relatively homogenous size ranges. The careful choice of appropriate volumetric shapes and taxa categories remains of prime importance to get precise results.     

Relative strengths of benthic algal nutrient and grazer limitation along a lake productivity gradient

The relative effects of nutrients and herbivores on primary producers are rarely compared across ecosystems that vary in potential primary productivity. Furthermore, proposed mechanisms to explain such patterns remain understudied. Here, I examine the strength of nutrient and grazer (herbivore) limitation (i.e., the extent to which producers’ growth is limited by insufficient nutrient supply or herbivory) of benthic algae across 13 southwest Michigan lakes that vary widely in productivity (i.e., resource supply). I compare the observed patterns of algal limitation and species composition to those predicted by two simple models: one that includes multiple species and species’ traits (the food-web model) and one that includes no variation in species or traits (the food-chain model). Species in the food-web model are assumed to display a tradeoff between resource competitive ability and resistance to herbivory (the “keystone predator” tradeoff). Among these lakes, benthic algal nutrient limitation was positive and declined significantly along a lake N:P gradient. In contrast, grazer limitation was negative and was not significantly related to any of the measured lake productivity variables. Negative grazer limitation indicated that the removal of grazers caused unexpected declines in algal biomass, which were potentially due to indirect, positive effects (e.g., nutrient recycling) of grazers. Nutrient limitation was significantly stronger than grazer limitation across lakes, which was more consistent with the food-web versus food-chain model. Changes in algal composition were also broadly consistent with predictions of the food-web model in that vulnerable, superior nutrient competitors dominated in low productivity lakes and more grazer-resistant species were observed in high productivity lakes. In general, these results point to the importance of examining limiting factors across systems and the consideration of key species’ traits when predicting and interpreting patterns.     

Benthic algal response to N and P enrichment along a pH gradient

Nutrient enrichment and its effect on benthic algal growth, community composition, and average cell size was assessed across two sites of differing pH within a single habitat. Nutrients were added using in situ substrata, which released either N, P, or no additional nutrients (controls) at each site for 21 days. Upon collection, chlorophyll and biovolume standing stocks of the attached algal microflora were measured. Chlorophyll concentration was different among all treatments, accumulating greatest on P, followed by N, and the least on C substrata (P < 0.001) and was highest at site-2 (P < 0.001), while total algal biovolume was highest on P compared to both N and C substrata (P < 0.05) and did not vary between sites. Increased growth on P substrata was due to the enhanced biovolume of filamentous green algae, although the affected taxa varied between sites. Biovolume to cell density ratios (as a measure of average cell size) were highest on P substrata over both N-enriched and control substrata (P < 0.05) and this pattern was similar between sites. Progression towards a community composed of larger cells following P enrichment observed along this pH gradient, seems to be related to the dominance of larger celled filamentous green algae. Thus, nutrients exhibited greater control on benthic algal growth than did changes in hydrogen ion concentration.Contribution number 581, Great Lakes Environmental Research LaboratoryContribution number 581, Great Lakes Environmental Research Laboratory