Lake trophic state and the limnological effects of omnivorous fish

Ecologists have hypothesized that planktivorous fish have greater effects on the plankton and water quality of oligotrophic lakes than eutrophic lakes. We tested this hypothesis in a tank-mesocosm experiment of factorial design in which five biomass levels of filter-feeding omnivorous gizzard shad (Dorosoma cepedianum) were cross-classified with two levels of lake trophic state achieved by filling tank-mesocosms with water and plankton transported by truck from two lakes with different trophic states. The presence of gizzard shad significantly increased total phosphorus, primary productivity, chlorophyll, and particulate phosphorus (PP) 2–20 and 20–200 μm and significantly decreased Secchi depth, cladocerans, copepods and PP > 200 μm. The effects of gizzard shad on chlorophyll, Secchi depth, cladocerans, copepods and PP 2–20 and > 200 μm were dependent on lake trophic state and most intense in the eutrophic lake system. This experiment suggests that filter-feeding omnivorous fish interact synergistically with trophic state so that the limnological effects of omnivorous fish become more intense with increased eutrophication.     

Plankton community structure in fluctuating environments and the role of productivity

Environmental variability in space and time can have significant influences on community structure. Temporal heterogeneity in nutrient supply has been shown in laboratory studies to have strong impacts on the diversity and composition of phytoplankton communities, depending on the scale of fluctuations. This paper extends the work in chemostats in a number of ways: using large-scale field mesocosms with natural plankton communities exposed to various frequencies of vertical mixing, modifying environmental productivity and incorporating higher trophic levels. The first major question and experiment focus on whether vertical mixing at various frequencies, and the associated nutrient pulse, has similar effects in nutrient-rich and nutrient-poor environments for predominantly single trophic level systems. The results indicate that the temporal scale of fluctuation is more of a structuring factor for phytoplankton communities in enriched enclosures, with little response under oligotrophic conditions. The second experiment examines the responsiveness of entire plankton communities (three trophic levels). Major shifts in community structure were absent under both nutrient-rich and nutrient-poor conditions. Responses were seen only in the demography of the top trophic level ( Chaoborus flavicans ). It appears from these experiments that the spatial disruption that accompanies mixing events may be more important than the temporal component (nutrient pulses) for phytoplankton. This appears to be the case only under conditions where natural spatial heterogeneity is high as it is when systems are enriched. When nutrient pulses are small, as they are in oligotrophic systems where recycling is efficient, little phytoplankton community response is observed. Finally, the inclusion of entire plankton food webs here suppressed the effects of the scale of intermittency in water column mixing at both low and high nutrient levels for all but the highest trophic level.     

Cascading effects of generalist fish introduction in oligotrophic lakes

Introduction of strictly planktivorous fish to lakes can alter plankton communities via cascading interactions in food webs. Less is known about the large-scale and long-term effects resulting from the introduction of fish with more generalist feeding habits, and the extent to which these effects depend on lake trophic status. Paleolimnological records of three oligotrophic lakes in Maine, USA were used to analyze the response of plankton communities to the introduction of white perch (Morone americana), a fish that often numerically dominates fish assemblages and switches from strict planktivory to a more generalist diet during ontogeny. After white perch introduction, cladoceran ephippia size increased up to 50 %, suggesting that the most important role of this generalist fish, with respect to water quality, is as a piscivorous trophic link. Algal standing crop declined by a quarter to over a half of pre-introduction levels, suggesting that top-down effects of white perch reduced phytoplankton biomass. In comparing these oligotrophic lakes to prior work in a eutrophic system, white perch introduction had similar effects on zooplankton body size; however, cascading effects to phytoplankton were only observed in low productivity lakes.     

广东省大中型供水水库营养现状及浮游生物的响应

于2000年调查了广东省18座大中型供水水库的水质现状并探讨了浮游生物对营养水平的响应。总氮、总磷、透明度和叶绿素a分别为0．15～7.15mg／L、0．003～0．387mg／L、0．4～6．3m和0．6～32．3ug／L。总氮、总磷、透明度均与叶绿素a呈较高的相关性。根据这4个因子的综合加权营养状态指数为23．7～55．1,季节差异不大,大多数水库处于中营养状态。金藻在中-富及富营养型水库中没有分布,而蓝藻、绿藻、硅藻和甲藻在调查水库中均有比较广的营养生态位,但它们的密度及相对优势度在各营养型水库中有一定的差异。高营养水平水库有较高的细胞密度和叶绿素a含量。营养水平较低的水库浮游植物以硅藻-甲藻、硅藻-绿藻或金藻-硅藻为主;营养水平较高的水库以蓝藻-硅藻或蓝藻-绿藻为主,并有较高的裸藻密度。浮游动物基本上以桡足类为优势种群,但在中-富营养和富营养型水库中,哲水蚤种类比低营养型水库中少。枝角类优势种类在各营养型水库差别不大。轮虫对水体营养水平的响应相对比较显著。低营养水平水库的轮虫以广营养型、中营养型或寡中营养型种类为主,种类数目比较少;富营养和中-富营养型水库的轮虫以喜在中营养到富营养条件下生长的种类为主,且轮虫种类数目比较多。     

Prey size diversity hinders biomass trophic transfer and predator size diversity promotes it in planktonic communities

Body size exerts multiple effects on plankton food-web interactions. However, the influence of size structure on trophic transfer remains poorly quantified in the field. Here, we examine how the size diversity of prey (nano-microplankton) and predators (mesozooplankton) influence trophic transfer efficiency (using biomass ratio as a proxy) in natural marine ecosystems. Our results support previous studies on single trophic levels: transfer efficiency decreases with increasing prey size diversity and is enhanced with greater predator size diversity. We further show that communities with low nano-microplankton size diversity and high mesozooplankton size diversity tend to occur in warmer environments with low nutrient concentrations, thus promoting trophic transfer to higher trophic levels in those conditions. Moreover, we reveal an interactive effect of predator and prey size diversities: the positive effect of predator size diversity becomes influential when prey size diversity is high. Mechanistically, the negative effect of prey size diversity on trophic transfer may be explained by unicellular size-based metabolic constraints as well as trade-offs between growth and predation avoidance with size, whereas increasing predator size diversity may enhance diet niche partitioning and thus promote trophic transfer. These findings provide insights into size-based theories of ecosystem functioning, with implications for ecosystem predictive models.     

Effects of water-column mixing on bacteria,phytoplankton, and rotifers under different levels of herbivory in a shallow eutrophic lake

Water-column mixing is known to have a decisive impact on plankton communities. The underlying mechanisms depend on the size and depth of the water body, nutrient status and the plankton community structure, and they are well understood for shallow polymictic and deep stratified lakes. Two consecutive mixing events of similar intensity under different levels of herbivory were performed in enclosures in a shallow, but periodically stratified, eutrophic lake, in order to investigate the effects of water-column mixing on bacteria abundance, phytoplankton abundance and diversity, and rotifer abundance and fecundity. When herbivory by filter-feeding zooplankton was low, water-column mixing that provoked a substantial nutrient input into the euphotic zone led to a strong net increase of bacteria and phytoplankton biomass. Phytoplankton diversity was lower in the mixed enclosures than in the undisturbed ones because of the greater contribution of a few fast-growing species. After the second mixing event, at a high biomass of filter-feeding crustaceans, the increase of phytoplankton biomass was lower than after the first mixing, and diversity remained unchanged because enhanced growth of small fast-growing phytoplankton was prevented by zooplankton grazing. Bacterial abundance did not increase after the second mixing, when cladoceran biomass was high. Changes in rotifer fecundity indicated a transmission of the phytoplankton response to the next trophic level. Our results suggest that water-column mixing in shallow eutrophic lakes with periodic stratification has a strong effect on the plankton community via enhanced nutrient availability rather than resuspension or reduced light availability. This fuels the basis of the classic and microbial food chain via enhanced phytoplankton and bacterial growth, but the effects on biomass may be damped by high levels of herbivory. Received: 3 May 1999 / Accepted: 13 April 2000     

Zooplankton biomass dynamics in oligotrophic versus eutrophic conditions: a test of the PEG model

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Microalgal plankton composition in shallow coastal inlets in contrasting trophic and alternative community states

As the trophic state of the environment changes, communities develop into divergent states. These community states are conventionally reflected through primary producers, because they are directly affected by nutrient availability. Studies of submerged macrophytes often focus on community composition to decipher the vegetative (community) state of the environment, while planktic microalgae are usually viewed more cursorily. Although microalgal plankton composition has been related to the trophic state of shallow temperate lakes, corresponding qualitative knowledge is lacking for shallow inlets in the sea. We assessed the composition of microalgal plankton in relation to that of submerged macrophytes in shallow inlets in the northern Baltic Sea during one ice-free season. Microalgal plankton composition varied distinctively among inlets in different trophic and vegetative states especially during early and mid-season, before becoming comparably uniform. These patterns were consisted both inside and outside of macrophyte beds and during day and night. Local and diurnal variation was comparably high in eutrophic and charophyte-dominated inlets, but only during early season. Microalgal plankton composition not only reflects the state of littoral communities in varying trophic conditions, but it may also be important for the whole trophic structure of those communities.     

Net heterotrophy in Faroe Islands clear-water lakes: causes and consequences for bacterioplankton and phytoplankton

1. Five oligotrophic clear‐water lakes on the Faroe Islands were studied during August 2000. Algal and bacterial production rates, community respiration, and CO₂ saturation were determined. In addition, we examined the plankton community composition (phytoplankton and heterotrophic nanoflagellates) and measured the grazing pressure exerted by common mixotrophic species on bacteria. 2. High respiration to primary production (6.6–33.2) and supersaturation of CO₂ (830–2140 μatm) implied that the lakes were net heterotrophic and that the pelagic heterotrophic plankton were subsidised by allochthonous organic carbon. However, in spite of the apparent high level of net heterotrophy, primary production exceeded bacterial production and the food base for higher trophic levels appeared to be mainly autotrophic. 3. We suggest that the observed net heterotrophy in these lakes was a result of the oligotrophic conditions and hence low primary production in combination with an input of allochthonous C with a relatively high availability. 4. Mixotrophic phytoplankton (Cryptomonas spp., Dinobryon spp. and flagellates cf. Ochromonas spp.) constituted a large percentage of the plankton community (17–83%), possibly as a result of their capacity to exploit bacteria as a means of acquiring nutrients in these nutrient poor systems.     

Interactive effects of warming,eutrophication and size structure: impacts on biodiversity and food‐web structure

Warming and eutrophication are two of the most important global change stressors for natural ecosystems, but their interaction is poorly understood. We used a dynamic model of complex, size‐structured food webs to assess interactive effects on diversity and network structure. We found antagonistic impacts: Warming increases diversity in eutrophic systems and decreases it in oligotrophic systems. These effects interact with the community size structure: Communities of similarly sized species such as parasitoid–host systems are stabilized by warming and destabilized by eutrophication, whereas the diversity of size‐structured predator–prey networks decreases strongly with warming, but decreases only weakly with eutrophication. Nonrandom extinction risks for generalists and specialists lead to higher connectance in networks without size structure and lower connectance in size‐structured communities. Overall, our results unravel interactive impacts of warming and eutrophication and suggest that size structure may serve as an important proxy for predicting the community sensitivity to these global change stressors.     

Predators, resources, and trophic chains in the regulation of plankton population and biomass in oligothrophic lakes

The relative strength of "top-down" versus "bottom-up" control of plankton community structure and biomass in two small oligotrophic lakes (with and without fish), located near the Polar circle (Russia), has been investigated for two years, 1996 and 1997. The comparative analyses of zooplankton biomass and species abundance showed strong negative effect of fish, stickeback (Pungitius pungitius L.), on the zooplankton community species, size structure and biomass of particular prey species but no effect on the biomass of the whole trophic level. An intensive predation in Verkhneye lake has lead to: 1) sixfold decline in biomass of large cladoceran Holopedium gibberum comparing to the lake lacking predator, 2) shift in the size mode in zooplankton community and the replacement of the typical large grazers by small species--Bosmina longirostris and rotifers. Their abundance and biomass even increased, demonstrating the stimulating effect of fish on the "inefficient" and unprofitable prey organisms. The analysis of contributions of different factors into the cladoceran's birth rate changes was applied to demonstrate the relative impact of predators and resources on zooplankton abundance. An occasional introduction of the stickleback to Vodoprovodnoye lake (the reference lake in 1996) in summer 1997 lead to drastic canges in this ecosystem: devastating decrease of zooplankton biomass and complete elimination of five previously dominant grazer species. The abundance of edible phytoplankton was slightly higher in the lake with fish in 1996 and considerably higher in the lake where fish has appeared in 1997 showing the prevailing "top-down" control of phytoplankton in oligotrophic ecosystem. The reasons of trophic cascade appearance in oligotrophic lakes are also discussed.     

Lack of Congruence in Species Diversity Indices and Community Structures of Planktonic Groups Based on Local Environmental Factors

The importance of analyzing the determinants of biodiversity and community composition by using multiple trophic levels is well recognized; however, relevant data are lacking. In the present study, we investigated variations in species diversity indices and community structures of the plankton taxonomic groups–zooplankton, rotifers, ciliates, and phytoplankton–under a range of local environmental factors in pond ecosystems. For each planktonic group, we estimated the species diversity index by using linear models and analyzed the community structure by using canonical correspondence analysis. We showed that the species diversity indices and community structures varied among the planktonic groups and according to local environmental factors. The observed lack of congruence among the planktonic groups may have been caused by niche competition between groups with similar trophic guilds or by weak trophic interactions. Our findings highlight the difficulty of predicting total biodiversity within a system, based upon a single taxonomic group. Thus, to conserve the biodiversity of an ecosystem, it is crucial to consider variations in species diversity indices and community structures of different taxonomic groups, under a range of local conditions.     

Zooplanktic indicators of trophy and their food

Regarding specific food requirements zooplankters in oligotrophic forest lakes are compared with indicators of other trophic conditions. Most zooplanktic indicators of eutrophic conditions feed on very small particles, to a large extent on bacteria, while the indicators of oligotrophy may be filtrators of courser particles. Both categories comprise raptors as well. In both extremes of lake types, hypertrophy and ultraoligotrophy, feeders on minute particles increase in relative importance, while moderate eutrophy or oligotrophy render a higher diversity of species and feeding categories.     

Increased Diversity of Predacious <Emphasis Type="Italic">Bdellovibrio</Emphasis>-Like Organisms (BLOs) as a Function of Eutrophication in Kumaon Lakes of India

Predation by Bdellovibrio-like organisms (BLOs) results in bacterial community succession in aquatic ecosystems. The effects of nutrient loading on the distribution and phylogeny of BLOs remain largely unknown. To this end, we present our findings on BLO diversity from four north-Indian lakes that are variable in their trophic status; Nainital is eutrophic, both, Bhimtal and Naukuchiatal are mesotrophic and Sattal remains oligotrophic, respectively. Initially, total heterotrophic bacteria and BLOs were quantified by most probable number (MPN) analyses using Pseudomonas putida and Escherichia coli as prey bacteria. Total bacterial numbers were at least two-logs higher in the eutrophic lake samples compared with oligotrophic lake. Similarly, BLO numbers were approximately 39-fold higher using Pseudomonas sp., which is likely the preferred prey within these lakes. Conversely, significant differences were not observed between mesotrophic and oligotrophic BLO numbers when E. coli was used as the prey. PCR-RFLP of small subunit rDNA (SSU rDNA) of BLOs, followed by cloning, sequencing, and taxonomic categorization revealed distinct differences such that, eutrophic lake consisted of higher BLO diversity compared with mesotrophic and oligotrophic lake, most likely due to both, higher numbers and availability of a diverse population of prey bacteria resulting from nutrient loading in this ecosystem.     

Aquatic macrophytes as limnological indicators

Species of submersed and floating-leaved aquatic macrophytes have been place in a series based on their patterns of occurrence in an ordination of floristic lists. Two chemical parameters from lake water analyses are correlated with the species assemblage in individual lakes and trophic categories are defined on the quantitative chemical characteristics of lake waters. The range and limiting tolerance of solute content for many aquatic species are described and related to these trophic categories. Restiction towards eutrophic conditions is considered as an obligate relatinship reflecting physiological demands. Some dystrophic and oligotrophic species are shown to have wide tolerance and are thought to be excluded from sites of higher trophic status by competition rather than physiological limitation.     

Patterns in freshwater diatom taxonomic distinctness along an eutrophication gradient

1. A variety of species richness measures have been used to assess the effects of environmental degradation on biodiversity. Such measures can be highly influenced by sample size, sampling effort, habitat type or complexity, however, and typically do not show monotonic responses to human impact. In addition to being independent of the degree of sampling effort involved in data acquisition, effective measures of biodiversity should reflect the degree of taxonomical relatedness among species within ecological assemblages and provide a basis for understanding observed diversity for a particular habitat type. Taxonomic diversity or distinctness indices emphasize the average taxonomic relatedness (i.e. degree of taxonomical closeness) between species in a community. 2. Eutrophication of freshwater ecosystems, mainly due to the increased availability of nutrients, notably phosphorus, has become a major environmental problem. Two measures of taxonomic distinctness (Average Taxonomic Distinctness and Variation in Taxonomic Distinctness) were applied to surface sediment diatoms from 45 lakes across the island of Ireland to examine whether taxonomic distinctness and nutrient enrichment were significantly related at a regional scale. The lakes span a range of concentrations of epilimnic total phosphorus (TP) and were grouped into six different types, based on depth and alkalinity levels, and three different categories according to trophic state (ultra‐oligotrophic and oligotrophic; mesotrophic; and eutrophic and hyper‐eutrophic). 3. The taxonomic distinctness measures revealed significant differences among lakes in the three different classes of trophic state, with nutrient‐rich lakes generally more taxonomically diverse than nutrient‐poor lakes. This implies that enrichment of oligotrophic lakes does not necessarily lead to a reduction in taxonomic diversity, at least as expressed by the indices used here. Furthermore, taxonomic distinctness was highly variable across the six different lake types regardless of nutrient level. 4. Results indicate that habitat availability and physical structure within the study lakes also exert a strong influence on the pattern of taxonomic diversity. Overall the results highlight problems with the use of taxonomic diversity measures for detecting impacts of freshwater eutrophication based on diatom assemblages.     

Viral production, decay rates, and life strategies along a trophic gradient in the North Adriatic Sea

Although the relationships between trophic conditions and viral dynamics have been widely explored in different pelagic environments, there have been few attempts at independent estimates of both viral production and decay. In this study, we investigated factors controlling the balance between viral production and decay along a trophic gradient in the north Adriatic basin, providing independent estimates of these variables and determining the relative importance of nanoflagellate grazing and viral life strategies. Increasing trophic conditions induced an increase of bacterioplankton growth rates and of the burst sizes. As a result, eutrophic waters displayed highest rates of viral production, which considerably exceeded observed rates of viral decay (up to 2.9 x 10(9) VLP liter(-1) h(-1)). Viral decay was also higher in eutrophic waters, where it accounted for ca. 40% of viral production, and dropped significantly to 1.3 to 10.7% in oligotrophic waters. These results suggest that viral production and decay rates may not necessarily be balanced in the short term, resulting in a net increase of viruses in the system. In eutrophic waters nanoflagellate grazing, dissolved-colloidal substances, and lysogenic infection were responsible together for the removal of ca. 66% of viral production versus 17% in oligotrophic waters. Our results suggest that different causative agents are primarily responsible for the removal of viruses from the water column in different trophic conditions. Factors other than those considered in the past might shed light on processes responsible for the removal and/or decay of viral particles from the water column.     

Bottom–up carbon subsidies and top–down predation pressure interact to affect aquatic food web structure

Human impacts such as eutrophication, overexploitation and climate change currently threaten future global food and drinking water supplies. Consequently, it is important that we understand how anthropogenic resource (bottom–up) and consumer (top–down) manipulations affect aquatic food web structure and production. Future climate changes are predicted to increase the inputs of terrestrial dissolved organic carbon to lakes. These carbon subsidies can either increase or decrease total basal production in aquatic food webs, depending on bacterial competition with phytoplankton for nutrients. This study examines the effects of carbon subsidies (bottom–up) on a pelagic community exposed to different levels of top–down predation. We conducted a large scale mesocosm experiment in an oligotrophic clear water lake in northern Sweden, using a natural plankton community exposed to three levels of glucose addition (0, 420 and 2100 μg C l^?1 total added glucose) and three levels of young‐of‐the‐year perch Perca fluviatilis density (0, 0.56 and 2 individuals m^?3). Bacterioplankton production doubled with glucose addition, but phytoplankton production was unaffected, in contrast to previous studies that have manipulated carbon, nutrients or light simultaneously. This suggests that carbon addition alone is not sufficient to reduce autotrophic production, at least in an oligotrophic lake dominated by mixotrophic phytoplankton. Larval perch grazing did not produce a classical trophic cascade, but substantially altered the species composition of crustacean zooplankton and ciliate trophic levels. Glucose addition increased the biomass of rotifers, thus potentially increasing energy transfer through the heterotrophic pathway, but only when fish were absent. This study illustrates that changes in community structure due to selective feeding by top‐predators can determine the influence of bottom–up carbon subsidies.     

广东省典型水库浮游植物群落特征与富营养化研究

研究了广东省19座主要水库2000年丰水期和枯水期浮游植物状况,并根据浮游植物群落结构和多样性指数对水库进行营养状况评价。结果表明,东江流域的新丰江和白盘珠水库水质良好,属于贫营养型水库;大部分水库为中营养型,而鹤地、契爷石和石岩这3座位于沿海经济发达地区水库属于富营养型。流域上游水库水质明显优于下游水库,山区水库水质优于沿海水库,东江流域水库水质普遍较好。北江流域次之,而粤西沿海和珠江三角洲地区各水库均受到不同程度的污染。     

The mixotroph Ochromonas tuberculata may invade and suppress specialist phago- and phototroph plankton communities depending on nutrient conditions

Mixotrophic organisms combine light, mineral nutrients, and prey as supplementary resources. Based on theoretical assumptions and field observations, we tested experimentally the hypothesis that mixotrophs may invade established plankton communities depending on the trophic status of the system, and investigated possible effects on food web structure, species diversity, and nutrient dynamics. To test our hypothesis, we inoculated the mixotrophic nanoflagellate Ochromonas tuberculata into established planktonic food webs, consisting of specialist phototrophs, specialist phagotrophs, and bacteria at different supplies of soluble inorganic nutrients and dissolved organic carbon. Oligotrophic systems facilitated the invasion of O. tuberculata in two different ways. First, the combination of photosynthesis and phagotrophy gave mixotrophs a competitive advantage over specialist phototrophs and specialist phagotrophs. Second, low nutrient supplies supported the growth of small plankton organisms that fell into the food size spectrum of mixotrophs. Conversely, high nutrient supplies prevented O. tuberculata from successfully invading the food webs. Two important conclusions were derived from our experiments. First, in contrast to a paradigm of ecology, specialization may not necessarily be the most successful strategy for survival under stable conditions. Indeed, the use of several resources with lower efficiency can be an equally, or even more, successful strategy in nature. Second, when limiting nutrients promote the growth of bacterio- and picophytoplankton, invading mixotrophs may have a habitat-ameliorating effect for higher trophic levels, gauged in terms of food quantity and quality. Using given resources more efficiently, O. tuberculata generated higher biomasses and expressed an increased nutritional value for potential planktivores, due to decreased cellular carbon to phosphorus (C:P) ratios compared to specialized plankton taxa. Our findings may help to explain why energy transfer efficiency between phytoplankton and higher trophic levels is generally higher in oligotrophic systems than in nutrient rich environments.