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1.  Interactive effects of chemical and biological controls on food-web composition in saline prairie lakes 
Aquatic Biosystems  2012;8:29.
Salinity is restricting habitatability for many biota in prairie lakes due to limited physiological abilities to cope with increasing osmotic stress. Yet, it remains unclear how salinity effects vary among major taxonomic groups and what role other environmental parameters play in shaping food-web composition. To answer these questions, we sampled fish, zooplankton and littoral macroinvertebrates in 20 prairie lakes (Saskatchewan, Canada) characterized by large gradients in water chemistry and lake morphometry. We showed that salinity thresholds differed among major taxonomic groups, as most fishes were absent above salinities of 2 g L-1, while littoral macroinvertebrates were ubiquitous. Zooplankton occurred over the whole salinity range, but changed taxonomic composition as salinity increased. Subsequently, the complexity of fish community (diversity) was associated with large changes in invertebrate communities. The directional changes in invertebrate communities to smaller taxa indicated that complex fish assemblages resulted in higher predation pressure. Most likely, as the complexity of fish community decreased, controls of invertebrate assemblages shifted from predation to competition and ultimately to productivity in hypersaline lakes. Surprisingly, invertebrate predators did not thrive in the absence of fishes in these systems. Furthermore, the here identified salinity threshold for fishes was too low to be a result of osmotic stress. Hence, winterkill was likely an important factor eliminating fishes in low salinity lakes that had high productivity and shallow water depth. Ultimately, while salinity was crucial, intricate combinations of chemical and biological mechanisms also played a major role in controlling the assemblages of major taxonomic groups in prairie lakes.
PMCID: PMC3547742  PMID: 23186395
Lake food-web; Diversity; Fish; Invertebrates; Salinity; Winter kill
2.  Isotopic evidence for the spatial heterogeneity of the planktonic food webs in the transition zone between river and lake ecosystems 
PeerJ  2013;1:e222.
Resources and organisms in food webs are distributed patchily. The spatial structure of food webs is important and critical to understanding their overall structure. However, there is little available information about the small-scale spatial structure of food webs. We investigated the spatial structure of food webs in a lake ecosystem at the littoral transition zone between an inflowing river and a lake. We measured the carbon isotope ratios of zooplankton and particulate organic matter (POM; predominantly phytoplankton) in the littoral zone of a saline lake. Parallel changes in the δ 13C values of zooplankton and their respective POMs indicated that there is spatial heterogeneity of the food web in this study area. Lake ecosystems are usually classified at the landscape level as either pelagic or littoral habitats. However, we showed small-scale spatial heterogeneity among planktonic food webs along an environmental gradient. Stable isotope data is useful for detecting spatial heterogeneity of habitats, populations, communities, and ecosystems.
PMCID: PMC3869182  PMID: 24392286
Food web; Plankton; Lake; Isotope; Spatial scale; Zooplankton; Phytoplankton; Carbon; Heterogeneity
3.  Assessing element-specific patterns of bioaccumulation across New England lakes 
The Science of the Total Environment  2012;421-422:230-237.
Little is known about differences among trace elements in patterns of bioaccumulation in freshwater food webs. Our goal was to identify patterns in bioaccumulation of different elements that are large and consistent enough to discern despite variation across lakes. We measured methylmercury (MeHg) and trace element (As, Cd, Hg, Pb, and Zn) concentrations in food web components of seven New England lakes on 3–5 dates per lake, and contrasted patterns of bioaccumulation across lakes, metals and seasons. In each lake, trace element concentrations were compared across trophic levels, including three size fractions of zooplankton, planktivorous fish, and piscivorous fish. The trophic position of each food web component was estimated from N isotope analysis. Trace element concentrations varied widely among taxa, lakes and sampling dates. Yet, we identified four consistent patterns of bioaccumulation that were consistent across lakes: (1) MeHg concentration increased (i.e., was biomagnified) and Pb concentration decreased (i.e., was biodiminished) with increased trophic position. (2) Zinc concentration (as with MeHg) was higher in fish than in zooplankton, but overall variation in Zn concentration (unlike MeHg) was low. (3) Arsenic and Cd concentrations (as with Pb) were lower in fish than in zooplankton, but (unlike Pb) were not significantly correlated with trophic position within zooplankton or fish groups. (4) Average summer concentrations of As, Pb, Hg, and MeHg in zooplankton significantly predicted their concentrations in either planktivorous or piscivorous fish. Our secondary goal was to review sampling approaches in forty-five published studies to determine the extent to which current sampling programs facilitate cross-lake and cross-study comparisons of bioaccumulation. We found that studies include different components of the food web and sample too infrequently to enable strong cross-lake and cross-study comparisons. We discuss sampling strategies that would improve our capacity to identify consistent patterns of bioaccumulation and drivers of elevated trace element concentrations under naturally high levels of variability.
PMCID: PMC3306538  PMID: 22356871
aquatic food web; biomagnification; fish; mercury; trace metal; trophic transfer; zooplankton
4.  Chemical and physical properties of some saline lakes in Alberta and Saskatchewan 
Saline Systems  2008;4:3.
The Northern Great Plains of Canada are home to numerous permanent and ephemeral athalassohaline lakes. These lakes display a wide range of ion compositions, salinities, stratification patterns, and ecosystems. Many of these lakes are ecologically and economically significant to the Great Plains Region. A survey of the physical characteristics and chemistry of 19 lakes was carried out to assess their suitability for testing new tools for determining past salinity from the sediment record.
Data on total dissolved solids (TDS), specific conductivity, temperature, dissolved oxygen (DO), and pH were measured in June, 2007. A comparison of these data with past measurements indicates that salinity is declining at Little Manitou and Big Quill Lakes in the province of Saskatchewan. However salinity is rising at other lakes in the region, including Redberry and Manito Lakes.
The wide range of salinities found across a small geographic area makes the Canadian saline lakes region ideal for testing salinity proxies. A nonlinear increase in salinity at Redberry Lake is likely influenced by its morphometry. This acceleration has ecological implications for the migratory bird species found within the Redberry Important Bird Area.
PMCID: PMC2365950  PMID: 18430240
5.  Lake morphometry and wind exposure may shape the plankton community structure in acidic mining lakes 
Limnologica (Online)  2010;40(2):161-166.
Acidic mining lakes (pH <3) are specific habitats exhibiting particular chemical and biological characteristics. The species richness is low and mixotrophy and omnivory are common features of the plankton food web in such lakes. The plankton community structure of mining lakes of different morphometry and mixing type but similar chemical characteristics (Lake 130, Germany and Lake Langau, Austria) was investigated. The focus was laid on the species composition, the trophic relationship between the phago-mixotrophic flagellate Ochromonas sp. and bacteria and the formation of a deep chlorophyll maximum along a vertical pH-gradient. The shallow wind-exposed Lake 130 exhibited a higher species richness than Lake Langau. This increase in species richness was made up mainly by mero-planktic species, suggesting a strong benthic/littoral – pelagic coupling. Based on the field data from both lakes, a nonlinear, negative relation between bacteria and Ochromonas biomass was found, suggesting that at an Ochromonas biomass below 50 μg C L−1, the grazing pressure on bacteria is low and with increasing Ochromonas biomass bacteria decline. Furthermore, in Lake Langau, a prominent deep chlorophyll maximum was found with chlorophyll concentrations ca. 50 times higher than in the epilimnion which was build up by the euglenophyte Lepocinclis sp. We conclude that lake morphometry, and specific abiotic characteristics such as mixing behaviour influence the community structure in these mining lakes.
PMCID: PMC3512575  PMID: 23225914
Acidic lake; Mining lake; Plankton; Mixotrophy; Rotifers; Flagellates; Ochromonas; Lepocinclis; Chlamydomonas; Deep chlorophyll maximum
6.  Mercury Concentrations in Lentic Fish Populations Related to Ecosystem and Watershed Characteristics 
Ambio  2010;39(1):14-19.
Predicting mercury (Hg) concentrations of fishes at large spatial scales is a fundamental environmental challenge with the potential to improve human health. In this study, mercury concentrations were examined for five species across 161 lakes and ecosystem, and watershed parameters were investigated as explanatory variables in statistical models. For all species, Hg concentrations were significantly, positively related to wetland coverage. For three species (largemouth bass, pike, and walleye), Hg concentrations were significantly, negatively related to lake trophic state index (TSI), suggestive of growth biodilution. There were no significant relationships between ecosystem size and mercury concentrations. However, Hg concentrations were strongly, positively related to ecosystem size across species. Scores of small or remote lakes that have never been tested could be prioritized for testing using models akin to those presented in this article. Such an approach could also be useful for exploring how Hg concentrations of fishes might respond to natural or anthropogenic changes to ecosystems over time.
PMCID: PMC3357655  PMID: 20496648
Contaminant; Ecology; Fish tissue; Food chain; GIS; Methylation; Wisconsin
7.  Spatial patterns of mercury in biota of Adirondack, New York lakes 
Ecotoxicology (London, England)  2011;20(7):1543-1554.
We studied the spatial distribution patterns of mercury (Hg) in lake water, littoral sediments, zooplankton, crayfish, fish, and common loons in 44 lakes of the Adirondacks of New York State, USA, a region that has been characterized as a “biological Hg hotspot”. Our study confirmed this pattern, finding that a substantial fraction of the lakes studied had fish and loon samples exceeding established criteria for human and wildlife health. Factors accounting for the spatial variability of Hg in lake water and biota were lake chemistry (pH, acid neutralizing capacity (ANC), percent carbon in sediments), biology (taxa presence, trophic status) and landscape characteristics (land cover class, lake elevation). Hg concentrations in zooplankton, fish and common loons were negatively associated with the lake water acid-base status (pH, ANC). Bioaccumulation factors (BAF) for methyl Hg (MeHg) increased from crayfish (mean log10 BAF = 5.7), to zooplankton (5.9), to prey fish (6.2), to larger fish (6.3), to common loons (7.2). MeHg BAF values in zooplankton, crayfish, and fish (yellow perch equivalent) all increased with increasing lake elevation. Our findings support the hypothesis that bioaccumulation of MeHg at the base of the food chain is an important controller of Hg concentrations in taxa at higher trophic levels. The characteristics of Adirondack lake-watersheds (sensitivity to acidic deposition; significant forest and wetland land cover; and low nutrient inputs) contribute to elevated Hg concentrations in aquatic biota.
Electronic supplementary material
The online version of this article (doi:10.1007/s10646-011-0717-y) contains supplementary material, which is available to authorized users.
PMCID: PMC3175042  PMID: 21691858
Spatial distribution; Methylmercury; Bioaccumulation; Aquatic biota; pH; Acid neutralizing capacity
8.  Shoreline urbanization interrupts allochthonous subsidies to a benthic consumer over a gradient of lake size 
Biology Letters  2011;7(4):551-554.
The role of resource subsidies across ecosystem boundaries has emerged as an important concept in contemporary ecology. For lake ecosystems, this has led to interest in quantifying the contribution of terrestrial allochthonous carbon to aquatic secondary production. An inverse relationship between habitat area and the role of allochthonous subsidies has been documented on marine islands and assumed for lakes, yet there have been no tests of this pattern among benthic (lake bottom) consumers. Here, we used carbon stable isotopes to trace terrestrial allochthonous and benthic autochthonous carbon use by the crayfish Pacifastacus leniusculus over a gradient of lake area, productivity and urbanization. Consistent with findings from terrestrial islands, habitat size dictated the importance of allochthonous subsidies, as P. leniusculus transitioned from using predominantly terrestrial carbon in small lakes to an increased reliance on autochthonous production in larger lakes. However, shoreline urbanization interacted with this pattern, particularly for small lakes where greater urbanization resulted in reduced use of allochthonous resources. As such, we provide, to our knowledge, the first confirmation of the predicted relationship between habitat size and importance of allochthonous subsidies to lake benthic consumers, but found that urbanization can interfere with this pattern.
PMCID: PMC3130242  PMID: 21389015
crayfish; lakes; resource subsidy; stable isotopes; urbanization
9.  Spatial and temporal variation in mercury bioaccumulation by zooplankton in Lake Champlain (North America) 
Trophic transfer of Hg across lakes within a region has been related to multiple environmental factors, but the nature of these relationships across distinct basins within individual large lakes is unknown. We investigated Hg bioaccumulation in zooplankton in basins of differing trophic status in Lake Champlain (Vermont, USA) to determine the strongest predictors of Hg bioaccumulation. Zooplankton were sampled in Malletts Bay (oligotrophic) and Missisquoi Bay (eutrophic) in 2005–2008. Zooplankton in the eutrophic basin had lower concentrations of total Hg and MeHg than those in the oligotrophic basin in all years but 2007, when no bloom occurred in Missisquoi. In addition, Hg concentrations in seston and small zooplankton, sampled during 2009 at 12 sites spanning the lake, decreased with increasing phytoplankton and zooplankton biomass. Thus, Hg bioaccumulation in zooplankton across basins in Lake Champlain is related to trophic status, as observed previously in multiple lake studies.
PMCID: PMC3356698  PMID: 21995871
Methylmercury; Biomass dilution; Zooplankton; Trophic; Lake Champlain
10.  Bottom-Up versus Top-Down Control of Hypo- and Epilimnion Free-Living Bacterial Community Structures in Two Neighboring Freshwater Lakes▿†‡ 
Applied and Environmental Microbiology  2011;77(11):3591-3599.
Bacterioplankton plays a central role in the microbial functioning of lacustrine ecosystems; however, factors that constrain its structural variation are still poorly understood. Here we evaluated the driving forces exerted by a large set of environmental and biological parameters on the temporal and spatial dynamics of free-living bacterial community structures (BCS) in two neighboring perialpine lakes, Lake Bourget and Lake Annecy, which differ in trophic status. We analyzed monthly data from a 1-year sampling period at two depths situated in the epi- and hypolimnia for each lake. Overall, denaturing gradient gel electrophoresis (DGGE) revealed significant differences in the BCS in the two lakes, characterized by a higher number of bands in the oligotrophic ecosystem (i.e., Lake Annecy). The temporal dynamics of BCS differed greatly between depths and lakes, with temporal scale patterns being much longer in the mesotrophic Lake Bourget. Direct-gradient multivariate ordination analyses showed that a complex array of biogeochemical parameters was the driving force behind BCS shifts in both lakes. Our results indicated that 60 to 80% of the variance was explained only by the bottom-up factors in both lakes, indicating the importance of nutrients and organic matter from autotrophic origin in controlling the BCS. Top-down regulation by flagellates together with ciliates or viruses was found only in the hypolimnion and not in the epilimnion for both lakes and explained less than 18% of the bacterial community changes during the year. Our study suggests that the temporal dynamics of the free-living bacterial community structure in deep perialpine lakes are dependent mainly on bottom-up factors and to a lesser extent on top-down factors, whatever the specific environmental conditions of these lakes.
PMCID: PMC3127590  PMID: 21478309
11.  Shorter Food Chain Length in Ancient Lakes: Evidence from a Global Synthesis 
PLoS ONE  2012;7(6):e37856.
Food webs may be affected by evolutionary processes, and effective evolutionary time ultimately affects the probability of species evolving to fill the niche space. Thus, ecosystem history may set important evolutionary constraints on community composition and food web structure. Food chain length (FCL) has long been recognized as a fundamental ecosystem attribute. We examined historical effects on FCL in large lakes spanning >6 orders of magnitude in age. We found that food chains in the world’s ancient lakes (n = 8) were significantly shorter than in recently formed lakes (n = 10) and reservoirs (n = 3), despite the fact that ancient lakes harbored much higher species richness, including many endemic species. One potential factor leading to shorter FCL in ancient lakes is an increasing diversity of trophic omnivores and herbivores. Speciation could simply broaden the number of species within a trophic group, particularly at lower trophic levels and could also lead to a greater degree of trophic omnivory. Our results highlight a counter-intuitive and poorly-understood role of evolutionary history in shaping key food web properties such as FCL.
PMCID: PMC3368915  PMID: 22701583
12.  Predicting Maximum Lake Depth from Surrounding Topography 
PLoS ONE  2011;6(9):e25764.
Information about lake morphometry (e.g., depth, volume, size, etc.) aids understanding of the physical and ecological dynamics of lakes, yet is often not readily available. The data needed to calculate measures of lake morphometry, particularly lake depth, are usually collected on a lake-by-lake basis and are difficult to obtain across broad regions. To span the gap between studies of individual lakes where detailed data exist and regional studies where access to useful data on lake depth is unavailable, we developed a method to predict maximum lake depth from the slope of the topography surrounding a lake. We use the National Elevation Dataset and the National Hydrography Dataset – Plus to estimate the percent slope of surrounding lakes and use this information to predict maximum lake depth. We also use field measured maximum lake depths from the US EPA's National Lakes Assessment to empirically adjust and cross-validate our predictions. We were able to predict maximum depth for ∼28,000 lakes in the Northeastern United States with an average cross-validated RMSE of 5.95 m and 5.09 m and average correlation of 0.82 and 0.69 for Hydrological Unit Code Regions 01 and 02, respectively. The depth predictions and the scripts are openly available as supplements to this manuscript.
PMCID: PMC3184154  PMID: 21984945
13.  Is the Relationship between Body Size and Trophic Niche Position Time-Invariant in a Predatory Fish? First Stable Isotope Evidence 
PLoS ONE  2010;5(2):e9120.
Characterizing relationships between individual body size and trophic niche position is essential for understanding how population and food-web dynamics are mediated by size-dependent trophic interactions. However, whether (and how) intraspecific size-trophic relationships (i.e., trophic ontogeny pattern at the population level) vary with time remains poorly understood. Using archival specimens of a freshwater predatory fish Gymnogobius isaza (Tanaka 1916) from Lake Biwa, Japan, we assembled a long-term (>40 years) time-series of the size-dependence of trophic niche position by examining nitrogen stable isotope ratios (δ15N) of the fish specimens. The size-dependence of trophic niche position was defined as the slope of the relationship between δ15N and log body size. Our analyses showed that the slope was significantly positive in about 60% of years and null in other years, changing through time. This is the first quantitative (i.e., stable isotope) evidence of long-term variability in the size-trophic relationship in a predatory fish. This finding had implications for the fish trophic dynamics, despite that about 60% of the yearly values were not statistically different from the long-term average. We proposed hypotheses for the underlying mechanism of the time-varying size-trophic relationship.
PMCID: PMC2817743  PMID: 20161751
14.  Population-Level Metrics of Trophic Structure Based on Stable Isotopes and Their Application to Invasion Ecology 
PLoS ONE  2012;7(2):e31757.
Biological invasions are a significant driver of human-induced global change and many ecosystems sustain sympatric invaders. Interactions occurring among these invaders have important implications for ecosystem structure and functioning, yet they are poorly understood. Here we apply newly developed metrics derived from stable isotope data to provide quantitative measures of trophic diversity within populations or species. We then use these to test the hypothesis that sympatric invaders belonging to the same functional feeding group occupy a smaller isotopic niche than their allopatric counterparts. Two introduced, globally important, benthic omnivores, Louisiana swamp crayfish (Procambarus clarkii) and carp (Cyprinus carpio), are sympatric in Lake Naivasha, Kenya. We applied our metrics to an 8-year data set encompassing the establishment of carp in the lake. We found a strong asymmetric interaction between the two invasive populations, as indicated by inverse correlations between carp abundance and measures of crayfish trophic diversity. Lack of isotopic niche overlap between carp and crayfish in the majority of years indicated a predominantly indirect interaction. We suggest that carp-induced habitat alteration reduced the diversity of crayfish prey, resulting in a reduction in the dietary niche of crayfish. Stable isotopes provide an integrated signal of diet over space and time, offering an appropriate scale for the study of population niches, but few isotope studies have retained the often insightful information revealed by variability among individuals in isotope values. Our population metrics incorporate such variation, are robust to the vagaries of sample size and are a useful additional tool to reveal subtle dietary interactions among species. Although we have demonstrated their applicability specifically using a detailed temporal dataset of species invasion in a lake, they have a wide array of potential ecological applications.
PMCID: PMC3283663  PMID: 22363724
15.  Methane Carbon Supports Aquatic Food Webs to the Fish Level 
PLoS ONE  2012;7(8):e42723.
Large amounts of the greenhouse gas methane (CH4) are produced by anaerobic mineralization of organic matter in lakes. In spite of extensive freshwater CH4 emissions, most of the CH4 is typically oxidized by methane oxidizing bacteria (MOB) before it can reach the lake surface and be emitted to the atmosphere. In turn, it has been shown that the CH4-derived biomass of MOB can provide the energy and carbon for zooplankton and macroinvertebrates. In this study, we demonstrate the presence of specific fatty acids synthesized by MOB in fish tissues having low carbon stable isotope ratios. Fish species, zooplankton, macroinvertebrates and the water hyacinth Eichhornia crassipes were collected from a shallow lake in Brazil and analyzed for fatty acids (FA) and carbon stable isotope ratios (δ13C). The fatty acids 16∶1ω8c, 16∶1ω8t, 16∶1ω6c, 16∶1ω5t, 18∶1ω8c and 18∶1ω8t were used as signature for MOB. The δ13C ratios varied from −27.7‰ to −42.0‰ and the contribution of MOB FA ranged from 0.05% to 0.84% of total FA. Organisms with higher total content of MOB FAs presented lower δ13C values (i.e. they were more depleted in 13C), while organisms with lower content of MOB signature FAs showed higher δ13C values. An UPGMA cluster analysis was carried out to distinguish grouping of organisms in relation to their MOB FA contents. This combination of stable isotope and fatty acid tracers provides new evidence that assimilation of methane-derived carbon can be an important carbon source for the whole aquatic food web, up to the fish level.
PMCID: PMC3413669  PMID: 22880091
16.  Effects of Temperature, Salinity and Fish in Structuring the Macroinvertebrate Community in Shallow Lakes: Implications for Effects of Climate Change 
PLoS ONE  2012;7(2):e30877.
Climate warming may lead to changes in the trophic structure and diversity of shallow lakes as a combined effect of increased temperature and salinity and likely increased strength of trophic interactions. We investigated the potential effects of temperature, salinity and fish on the plant-associated macroinvertebrate community by introducing artificial plants in eight comparable shallow brackish lakes located in two climatic regions of contrasting temperature: cold-temperate and Mediterranean. In both regions, lakes covered a salinity gradient from freshwater to oligohaline waters. We undertook day and night-time sampling of macroinvertebrates associated with the artificial plants and fish and free-swimming macroinvertebrate predators within artificial plants and in pelagic areas. Our results showed marked differences in the trophic structure between cold and warm shallow lakes. Plant-associated macroinvertebrates and free-swimming macroinvertebrate predators were more abundant and the communities richer in species in the cold compared to the warm climate, most probably as a result of differences in fish predation pressure. Submerged plants in warm brackish lakes did not seem to counteract the effect of fish predation on macroinvertebrates to the same extent as in temperate freshwater lakes, since small fish were abundant and tended to aggregate within the macrophytes. The richness and abundance of most plant-associated macroinvertebrate taxa decreased with salinity. Despite the lower densities of plant-associated macroinvertebrates in the Mediterranean lakes, periphyton biomass was lower than in cold temperate systems, a fact that was mainly attributed to grazing and disturbance by fish. Our results suggest that, if the current process of warming entails higher chances of shallow lakes becoming warmer and more saline, climatic change may result in a decrease in macroinvertebrate species richness and abundance in shallow lakes.
PMCID: PMC3290630  PMID: 22393354
17.  A cascade of evolutionary change alters consumer-resource dynamics and ecosystem function 
It is becoming increasingly clear that intraspecific evolutionary divergence influences the properties of populations, communities and ecosystems. The different ecological impacts of phenotypes and genotypes may alter selection on many species and promote a cascade of ecological and evolutionary change throughout the food web. Theory predicts that evolutionary interactions across trophic levels may contribute to hypothesized feedbacks between ecology and evolution. However, the importance of ‘cascading evolutionary change’ in a natural setting is unknown. In lakes in Connecticut, USA, variation in migratory behaviour and feeding morphology of a fish predator, the alewife (Alosa pseudoharengus), drives life-history evolution in a species of zooplankton prey (Daphnia ambigua). Here we evaluated the reciprocal impacts of Daphnia evolution on ecological processes in laboratory mesocosms. We show that life-history evolution in Daphnia facilitates divergence in rates of population growth, which in turn significantly alters consumer-resource dynamics and ecosystem function. These experimental results parallel trends observed in lakes. Such results argue that a cascade of evolutionary change, which has occurred over contemporary timescales, alters community and ecosystem processes.
PMCID: PMC3385726  PMID: 22628469
life-history evolution; local adaptation; eco-evolutionary dynamics
18.  Effect of grazers and viruses on bacterial community structure and production in two contrasting trophic lakes 
BMC Microbiology  2011;11:88.
Over the last 30 years, extensive studies have revealed the crucial roles played by microbes in aquatic ecosystems. It has been shown that bacteria, viruses and protozoan grazers are dominant in terms of abundance and biomass. The frequent interactions between these microbiological compartments are responsible for strong trophic links from dissolved organic matter to higher trophic levels, via heterotrophic bacteria, which form the basis for the important biogeochemical roles of microbial food webs in aquatic ecosystems. To gain a better understanding of the interactions between bacteria, viruses and flagellates in lacustrine ecosystems, we investigated the effect of protistan bacterivory on bacterial abundance, production and structure [determined by 16S rRNA PCR-DGGE], and viral abundance and activity of two lakes of contrasting trophic status. Four experiments were conducted in the oligotrophic Lake Annecy and the mesotrophic Lake Bourget over two seasons (early spring vs. summer) using a fractionation approach. In situ dark vs. light incubations were performed to consider the effects of the different treatments in the presence and absence of phototrophic activity.
The presence of grazers (i.e. < 5-μm small eukaryotes) affected viral production positively in all experiments, and the stimulation of viral production (compared to the treatment with no eukaryotic predators) was more variable between lakes than between seasons, with the highest value having been recorded in the mesotrophic lake (+30%). Viral lysis and grazing activities acted additively to sustain high bacterial production in all experiments. Nevertheless, the stimulation of bacterial production was more variable between seasons than between lakes, with the highest values obtained in summer (+33.5% and +37.5% in Lakes Bourget and Annecy, respectively). The presence of both predators (nanoflagellates and viruses) did not seem to have a clear influence upon bacterial community structure according to the four experiments.
Our results highlight the importance of a synergistic effect, i.e. the positive influence of grazers on viral activities in sustaining (directly and indirectly) bacterial production and affecting composition, in both oligotrophic and mesotrophic lakes.
PMCID: PMC3114703  PMID: 21527043
Lakes; microcosm; spring-summer variations; bacterial production; viral production; bacterial community structure; grazers
19.  Saline systems of the Great Plains of western Canada: an overview of the limnogeology and paleolimnology 
Saline Systems  2005;1:10.
In much of the northern Great Plains, saline and hypersaline lacustrine brines are the only surface waters present. As a group, the lakes of this region are unique: there is no other area in the world that can match the concentration and diversity of saline lake environments exhibited in the prairie region of Canada and northern United States. The immense number of individual salt lakes and saline wetlands in this region of North America is staggering. Estimates vary from about one million to greater than 10 million, with densities in some areas being as high as 120 lakes/km2.
Despite over a century of scientific investigation of these salt lakes, we have only in the last twenty years advanced far enough to appreciate the wide spectrum of lake types, water chemistries, and limnological processes that are operating in the modern settings. Hydrochemical data are available for about 800 of the lake brines in the region. Composition, textural, and geochemical information on the modern bottom sediments has been collected for just over 150 of these lakes. Characterization of the biological and ecological features of these lakes is based on even fewer investigations, and the stratigraphic records of only twenty basins have been examined.
The lake waters show a considerable range in ionic composition and concentration. Early investigators, concentrating on the most saline brines, emphasized a strong predominance of Na+ and SO4-2 in the lakes. It is now realized, however, that not only is there a complete spectrum of salinities from less than 1 ppt TDS to nearly 400 ppt, but also virtually every water chemistry type is represented in lakes of the region. With such a vast array of compositions, it is difficult to generalize. Nonetheless, the paucity of Cl-rich lakes makes the northern Great Plains basins somewhat unusual compared with salt lakes in many other areas of the world (e.g., Australia, western United States). Compilations of the lake water chemistries show distinct spatial trends and regional variations controlled by groundwater input, climate, and geomorphology. Short-term temporal variations in the brine composition, which can have significant effects on the composition of the modern sediments, have also been well documented in several individual basins.
From a sedimentological and mineralogical perspective, the wide range of water chemistries exhibited by the lakes leads to an unusually large diversity of modern sediment composition. Over 40 species of endogenic precipitates and authigenic minerals have been identified in the lacustrine sediments. The most common non-detrital components of the modern sediments include: calcium and calcium-magnesium carbonates (magnesian calcite, aragonite, dolomite), and sodium, magnesium, and sodium-magnesium sulfates (mirabilite, thenardite, bloedite, epsomite). Many of the basins whose brines have very high Mg/Ca ratios also have hydromagnesite, magnesite, and nesquehonite. Unlike salt lakes in many other areas of the world, halite, gypsum, and calcite are relatively rare endogenic precipitates in the Great Plains lakes. The detrital fraction of the lacustrine sediments is normally dominated by clay minerals, carbonate minerals, quartz, and feldspars.
Sediment accumulation in these salt lakes is controlled and modified by a wide variety of physical, chemical, and biological processes. Although the details of these modern sedimentary processes can be exceedingly complex and difficult to discuss in isolation, in broad terms, the processes operating in the salt lakes of the Great Plains are ultimately controlled by three basic factors or conditions of the basin: (a) basin morphology; (b) basin hydrology; and (c) water salinity and composition. Combinations of these parameters interact to control nearly all aspects of modern sedimentation in these salt lakes and give rise to four 'end member' types of modern saline lacustrine settings in the Great Plains: (a) clastics-dominated playas; (b) salt-dominated playas; (c) deep water, non-stratified lakes; and (d) deep water, "permanently" stratified lakes.
PMCID: PMC1315329  PMID: 16297237
20.  Genetic Diversity of Small Eukaryotes in Lakes Differing by Their Trophic Status 
Applied and Environmental Microbiology  2005;71(10):5935-5942.
Small eukaryotes, cells with a diameter of less than 5 μm, are fundamental components of lacustrine planktonic systems. In this study, small-eukaryote diversity was determined by sequencing cloned 18S rRNA genes in three libraries from lakes of differing trophic status in the Massif Central, France: the oligotrophic Lake Godivelle, the oligomesotrophic Lake Pavin, and the eutrophic Lake Aydat. This analysis shows that the least diversified library was in the eutrophic lake (12 operational taxonomic units [OTUs]) and the most diversified was in the oligomesotrophic lake (26 OTUs). Certain groups were present in at least two ecosystems, while the others were specific to one lake on the sampling date. Cryptophyta, Chrysophyceae, and the strictly heterotrophic eukaryotes, Ciliophora and fungi, were identified in the three libraries. Among the small eukaryotes found only in two lakes, Choanoflagellida and environmental sequences (LKM11) were not detected in the eutrophic system whereas Cercozoa were confined to the oligomesotrophic and eutrophic lakes. Three OTUs, linked to the Perkinsozoa, were detected only in the Aydat library, where they represented 60% of the clones of the library. Chlorophyta and Haptophyta lineages were represented by a single clone and were present only in Godivelle and Pavin, respectively. Of the 127 clones studied, classical pigmented organisms (autotrophs and mixotrophs) represented only a low proportion regardless of the library's origin. This study shows that the small-eukaryote community composition may differ as a function of trophic status; certain lineages could be detected only in a single ecosystem.
PMCID: PMC1266003  PMID: 16204507
21.  Sixty years of environmental change in the world's largest freshwater lake – Lake Baikal, Siberia 
Global Change Biology  2008;14(8):1947-1958.
High-resolution data collected over the past 60 years by a single family of Siberian scientists on Lake Baikal reveal significant warming of surface waters and long-term changes in the basal food web of the world's largest, most ancient lake. Attaining depths over 1.6 km, Lake Baikal is the deepest and most voluminous of the world's great lakes. Increases in average water temperature (1.21 °C since 1946), chlorophyll a (300% since 1979), and an influential group of zooplankton grazers (335% increase in cladocerans since 1946) may have important implications for nutrient cycling and food web dynamics. Results from multivariate autoregressive (MAR) modeling suggest that cladocerans increased strongly in response to temperature but not to algal biomass, and cladocerans depressed some algal resources without observable fertilization effects. Changes in Lake Baikal are particularly significant as an integrated signal of long-term regional warming, because this lake is expected to be among those most resistant to climate change due to its tremendous volume. These findings highlight the importance of accessible, long-term monitoring data for understanding ecosystem response to large-scale stressors such as climate change.
PMCID: PMC3597250
Daphnia; global warming; great lakes; high-latitude; long-term ecological research; plankton; Russia; subarctic; time series; trophic structure
22.  Invasive Mussels Alter the Littoral Food Web of a Large Lake: Stable Isotopes Reveal Drastic Shifts in Sources and Flow of Energy 
PLoS ONE  2012;7(12):e51249.
We investigated how establishment of invasive dreissenid mussels impacted the structure and energy sources of the littoral benthic food web of a large temperate lake. We combined information about pre- and postdreissenid abundance, biomass, and secondary production of the littoral benthos with results of carbon and nitrogen stable isotope analysis of archival (predreissenid) and recent (postdreissenid) samples of all common benthic taxa. This approach enabled us to determine the importance of benthic and sestonic carbon to the littoral food web before, and more than a decade after dreissenid establishment. Long term dreissenid presence was associated with a 32-fold increase in abundance, 6-fold increase in biomass, and 14-fold increase in secondary production of the littoral benthos. Dreissenids comprised a large portion of the post-invasion benthos, making up 13, 38, and 56% of total abundance, biomass, and secondary production, respectively. The predreissenid food web was supported primarily by benthic primary production, while sestonic material was relatively more important to the postdreissenid food web. The absolute importance of both sestonic material and benthic primary production to the littoral benthos increased considerably following dreissenid establishment. Our results show drastic alterations to food web structure and suggest that dreissenid mussels redirect energy and material from the water column to the littoral benthos both through biodeposition of sestonic material as well as stimulation of benthic primary production.
PMCID: PMC3524176  PMID: 23284673
23.  Microbiology of Lonar Lake and other soda lakes 
The ISME Journal  2012;7(3):468-476.
Soda lakes are saline and alkaline ecosystems that are believed to have existed throughout the geological record of Earth. They are widely distributed across the globe, but are highly abundant in terrestrial biomes such as deserts and steppes and in geologically interesting regions such as the East African Rift valley. The unusual geochemistry of these lakes supports the growth of an impressive array of microorganisms that are of ecological and economic importance. Haloalkaliphilic Bacteria and Archaea belonging to all major trophic groups have been described from many soda lakes, including lakes with exceptionally high levels of heavy metals. Lonar Lake is a soda lake that is centered at an unusual meteorite impact structure in the Deccan basalts in India and its key physicochemical and microbiological characteristics are highlighted in this article. The occurrence of diverse functional groups of microbes, such as methanogens, methanotrophs, phototrophs, denitrifiers, sulfur oxidizers, sulfate reducers and syntrophs in soda lakes, suggests that these habitats harbor complex microbial food webs that (a) interconnect various biological cycles via redox coupling and (b) impact on the production and consumption of greenhouse gases. Soda lake microorganisms harbor several biotechnologically relevant enzymes and biomolecules (for example, cellulases, amylases, ectoine) and there is the need to augment bioprospecting efforts in soda lake environments with new integrated approaches. Importantly, some saline and alkaline lake ecosystems around the world need to be protected from anthropogenic pressures that threaten their long-term existence.
PMCID: PMC3578565  PMID: 23178675
Lonar Lake; soda lakes; alkaliphiles; stable-isotope probing; microbial prospecting
24.  Influence of Climate Change and Trophic Coupling across Four Trophic Levels in the Celtic Sea 
PLoS ONE  2012;7(10):e47408.
Climate change has had profound effects upon marine ecosystems, impacting across all trophic levels from plankton to apex predators. Determining the impacts of climate change on marine ecosystems requires understanding the direct effects on all trophic levels as well as indirect effects mediated by trophic coupling. The aim of this study was to investigate the effects of climate change on the pelagic food web in the Celtic Sea, a productive shelf region in the Northeast Atlantic. Using long-term data, we examined possible direct and indirect ‘bottom-up’ climate effects across four trophic levels: phytoplankton, zooplankton, mid-trophic level fish and seabirds. During the period 1986–2007, although there was no temporal trend in the North Atlantic Oscillation index (NAO), the decadal mean Sea Surface Temperature (SST) in the Celtic Sea increased by 0.66±0.02°C. Despite this, there was only a weak signal of climate change in the Celtic Sea food web. Changes in plankton community structure were found, however this was not related to SST or NAO. A negative relationship occurred between herring abundance (0- and 1-group) and spring SST (0-group: p = 0.02, slope = −0.305±0.125; 1-group: p = 0.04, slope = −0.410±0.193). Seabird demographics showed complex species–specific responses. There was evidence of direct effects of spring NAO (on black-legged kittiwake population growth rate: p = 0.03, slope = 0.0314±0.014) as well as indirect bottom-up effects of lagged spring SST (on razorbill breeding success: p = 0.01, slope = −0.144±0.05). Negative relationships between breeding success and population growth rate of razorbills and common guillemots may be explained by interactions between mid-trophic level fish. Our findings show that the impacts of climate change on the Celtic Sea ecosystem is not as marked as in nearby regions (e.g. the North Sea), emphasizing the need for more research at regional scales.
PMCID: PMC3472987  PMID: 23091621
25.  Productivity, Disturbance and Ecosystem Size Have No Influence on Food Chain Length in Seasonally Connected Rivers 
PLoS ONE  2013;8(6):e66240.
The food web is one of the oldest and most central organising concepts in ecology and for decades, food chain length has been hypothesised to be controlled by productivity, disturbance, and/or ecosystem size; each of which may be mediated by the functional trophic role of the top predator. We characterised aquatic food webs using carbon and nitrogen stable isotopes from 66 river and floodplain sites across the wet-dry tropics of northern Australia to determine the relative importance of productivity (indicated by nutrient concentrations), disturbance (indicated by hydrological isolation) and ecosystem size, and how they may be affected by food web architecture. We show that variation in food chain length was unrelated to these classic environmental determinants, and unrelated to the trophic role of the top predator. This finding is a striking exception to the literature and is the first published example of food chain length being unaffected by any of these determinants. We suggest the distinctive seasonal hydrology of northern Australia allows the movement of fish predators, linking isolated food webs and potentially creating a regional food web that overrides local effects of productivity, disturbance and ecosystem size. This finding supports ecological theory suggesting that mobile consumers promote more stable food webs. It also illustrates how food webs, and energy transfer, may function in the absence of the human modifications to landscape hydrological connectivity that are ubiquitous in more populated regions.
PMCID: PMC3680379  PMID: 23776641

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