A lake as a microcosm: reflections on developments in aquatic ecology

In the present study, we aim at relating Forbes' remarkable paper on "The lake as a microcosm", published 125 years ago, to the present status of knowledge in our own research group. Hence, we relate the observations Forbes made to our own microcosm, Lake Krankesjon in southern Sweden, that has been intensively studied by several research groups for more than three decades. Specifically, we focus on the question: Have we made any significant progress or did Forbes and colleagues blaze the trail through the unknown wilderness and we are mainly paving that intellectual road? We conclude that lakes are more isolated than many other biomes, but have, indeed, many extensions, for example, input from the catchment, fishing and fish migration. We also conclude that irrespective of whether lakes should be viewed as microcosms or not, the paper by Forbes has been exceptionally influential and still is, especially since it touches upon almost all aspects of the lake ecosystem, from individual behaviour to food web interactions and environmental issues. Therefore, there is no doubt that even if 125 years have passed, Forbes' paper still is a source of inspiration and deserves to be read. Hence, although aquatic ecology has made considerable progress over the latest century, Forbes might be viewed as one of the major pioneers and visionary scientists of limnology

Is size-assortative mating important for rapid pigment differentiation in a freshwater isopod?

Identifying mechanisms behind assortative mating is central to the understanding of ecological divergence and speciation. Recent studies show that populations of the freshwater isopod Asellus aquaticus can rapidly become locally differentiated when submerged Chara vegetation expands in lakes. In the novel Chara habitat, isopods have become lighter pigmented and smaller than in ancestral reed stands. In this study, we used a laboratory multiple-choice experiment to investigate assortative mating as a possible prezygotic reproductive barrier between Chara and reed isopods. Mating was assortative when Chara isopods were experimentally mixed with isopods from an adjacent reed site with large-size individuals, suggesting a partial prezygotic reproductive barrier. No deviation from random mating could, however, be detected when Chara isopods were mixed with smaller sized isopods from another reed site. In both experiments, assortative mating was apparently based on size, as Chara isopods were larger and reed isopods smaller in mixed pairs than in assortative pairs. Pigmentation did not have any clear influence on mating. We suggest that divergence in pigmentation evolved through natural selection in conjunction with size-assortative mating indirectly causing assortative mating between Chara and reed isopods. Size-assortative mating is likely a by-product of natural selection, but its importance may hypothetically be transient, if selection erodes the correlation between pigmentation and size over time

Shifts between clear and turbid states in a shallow lake: multi-causal stress from climate, nutrients and biotic interactions

We used long-term monitoring data to assess causes behind a recent shift front a clear to a turbid water state in Lake Takern, Sweden. The lake has a previous record of shifts between clear-water and turbid states. but the causes behind these shifts are not well known. During the recent shift, which occurred in 1995-1997, Submerged vegetation Subsequently declined after a 30-year period of clear-water and abundant vegetation. Among the possible explanations we identify several processes unlikely to have contributed to the recent shift from clear to turbid conditions. including long-term changes in external input of phosphorus. fluctuations in water level. and changes in zooplankton grazing pressure. Instead, likely scenarios to have contributed to the macrophyte decline, and thereby to the shift were: (1) a series of mild winters with short ice cover and absence of winter-kills of fish, leading to high biomasses of benthivorous and planktivorous fish before the shift, and thereby increased bioturbation and internal nutrient recycling, (2) unusually cool and windy springs the years before and during, the shift, leading to unfavourable conditions during the establishing phase of submerged macrophytes. Both shorter periods of ice cover and high wind velocity in winter and spring were associated with climate. approximated by the North Atlantic Oscillation (NAO). We argue that none of these processes alone can force the lake front the clear to the turbid state, but that several stress factors in concert are necessary to initiate a shift. Hence, we conclude that climate variability is likely to have contributed to a multi-causal stress. reducing the resilience of the clear-water state and finally triggering the Shift through inter-year dependent changes; in biomass of submerged macrophytes and fish, organism groups known to have key roles in the dynamics of shallow lakes

Does a long-term oscillation in nitrogen concentration reflect climate impact on submerged vegetation and vulnerability to state shifts in a shallow lake?

Various ecosystems, including shallow lakes, are suggested to possess alternative stable state dynamics. The response of such systems to environmental change is non-linear and not fully reversible, which calls for identification of feedback mechanisms and subtle changes connected to structural stability. Here, we used a 25-year data series on water chemistry to make inferences on processes prior to a recent shift from a clear to a turbid state in Lake Takern, Sweden. Before the shift, annual concentration of total organic nitrogen (TON) described a cyclic pattern, with a periodicity of eight years. Annual TON was negatively correlated with the magnitude of a summer decline in calcium carbonate, treated as a proxy of the seasonal production of submerged vegetation. Cross-correlations of TON and the north Atlantic oscillation (NAO) indicated a possible connection to climate. The strongest correlation was obtained by a three-year lag of the NAO index. Using a set of linear time series models, the most parsimonious model was a 3(rd) order autoregressive model with NAO, delayed three years, as a covariate. Analyses of seasonality indicated that the delayed NAO signal was strongly correlated with summer TON. Also, the autocorrelation function was very similar for these two time series, and autoregressive models including NAO as a covariate were strongly supported (sum of Akaike weights = 0.93). These results indicate that climate may have contributed to the regime shift through lowered macrophyte production at the time of the shift, and therefore most likely also a depleted resilience of the clear water state. The delayed effect of climate is suggested to result from indirect and inter-year dependent response of submerged vegetation to fish kills during harsh winters

Seasonal changes of mechanisms maintaining clear water in a shallow lake with abundant Chara vegetation

The study is based on monitoring data on the seasonal variation during four (1996-1999) vegetation periods, as well as long-term summer data on submerged vegetation, nutrients, light, phyto-plankton and zooplankton in Lake Krankesjon, a shallow, calcium-rich, moderately eutrophic take in southern Sweden. The lake has been in the clear water state with abundant submerged vegetation since the end of the 1980s. Somewhat lower summer biomass of submerged macrophytes during 1997 and 1999 indicates a temporary instability of the clear water state. During these 2 years, summer transparency was about 1.2-2.1 m, while concentrations of total phosphorus and chlorophyll (Chl) a were about 26-40 and 8-18 mug 1(-1), respectively. Summer biomass of submerged macrophytes was higher during 1996 and 1998. In both years, a distinct increase in light availability and decrease in concentrations of nutrients and chlorophyll occurred simultaneously with the development of dense Chara vegetation. Summer values for transparency were about 2.0-2.5 m, while concentrations of total phosphorus and Chl a were about 20-32 and 4-11 mug 1(-1), respectively. Summer biomass of crustacean zooplankton was below 250 mug 1(-1) during all 4 years. A peak abundance of Cladocera (mainly Bosmina longirostris) during May or June caused only a short-term reduction in chlorophyll concentrations that was more pronounced in 1997 than in 1996. Measured light attenuation during 1999 was closely correlated with light attenuation calculated from the amount of suspended solids, chlorophyll concentrations, and water colour. Detritus contributed most to the total amount of suspended solids, while chlorophyll was the main contributor to light attenuation. A long-term decrease of the ratios between chlorophyll and total phosphorus suggests that phytoplankton in the clear water state is limited by factors other than total phosphorus concentrations. Increased sedimentation rate, carbon limitation, allelopathy and a lower bioavailable fraction of the total amount of phosphorus are possible explanations, while nitrogen limitation and grazing from zooplankton probably are of minor importance. Possible reasons for the "instability" of the clear water state during 1997 and 1999 are discussed. Unusually high water level as well as cold and windy weather during the spring of 1996-1999 may have caused a slow and late growth of the plants and thus a temporary instability. However, a tendency for an increase in total phosphorus concentrations and sediment accumulation along the wind-protected shores during the clear water state indicate the possibility of a long-term. destabilization which contradicts the alternative stable states model. (C) 2002 Elsevier Science B.V. All rights reserved

The role of different reproductive barriers during phenotypic divergence of isopod ecotypes.

The question of how diverging populations become separate species by restraining gene flow is a central issue in evolutionary biology. Assortative mating might emerge early during adaptive divergence, but the role of other types of reproductive barriers such as migration modification have recently received increased attention. We demonstrate that two recently diverged ecotypes of a freshwater isopod (Asellus aquaticus) have rapidly developed premating isolation, and this isolation barrier has emerged independently and in parallel in two south Swedish lakes. This is consistent with ecological speciation theory, which predicts that reproductive isolation arises as a byproduct of ecological divergence. We also find that in one of these lakes, habitat choice acts as the main barrier to gene flow. These observations and experimental results suggest that migration modification might be as important as assortative mating in the early stages of ecological speciation. Simulations suggest that the joint action of these two isolating barriers is likely to greatly facilitate adaptive divergence, compared to if each barrier was acting alone

Habitat-specific pigmentation in a freshwater isopod: Adaptive evolution over a small spatiotemporal scale

Pigmentation in the freshwater isopod Asellus aquaticus (Crustacea) differed between habitats in two Swedish lakes. In both lakes, isopods had lighter pigmentation in stands of submerged vegetation, consisting of stoneworts (Chara spp.), than in nearby stands of reed (Phragmites australis). Experimental crossings of light and dark isopods in a common environment showed that pigmentation had a genetic basis and that genetic variance was additive. Environmental effects of diet or chromatophore adjustment to the background had minor influence on pigmentation, as shown by laboratory rearing of isopods on stonewort or reed substrates, as well as analyses of stable isotope ratios for isopods collected in the field. In both study lakes, the average phenotype became lighter with time (across generations) in recently established stonewort stands. Taken together, these results indicate that altered phenotype pigmentation result from evolutionary responses to local differences in natural selection. Based on the assumption of two generations per year, the evolutionary rate of change in pigmentation was 0.08 standard deviations per generation (haldanes) over 20 generations in one lake and 0.22 haldanes over two generations in the other lake. This genetic change occurred during an episode of population growth in a novel habitat, a situation known to promote adaptive evolution. In addition, stonewort stands constitute large and persistent patches, characteristics that tend to preserve local adaptations produced by natural selection. Results from studies on selective forces behind the adaptive divergence suggest that selective predation from visually oriented predators is a possible selective agent. We found no indications of phenotype-specific movements between habitats. Mating within stonewort stands was random with respect to pigmentation, but on a whole-lake scale it is likely that mating is assortative, as a result of local differences in phenotype distribution