Jef Huisman

Blooms Like It Hot

A link exists between global warming and the worldwide proliferation of harmful cyanobacterial blooms.

CHANGES IN TURBULENT MIXING SHIFT COMPETITION FOR LIGHT BETWEEN PHYTOPLANKTON SPECIES

The intriguing impact of physical mixing processes on species interactions has always fascinated ecologists. Here, we exploit recent advances in plankton models to develop competition theory that predicts how changes in turbulent mixing affect competition for light between buoyant and sinking phytoplankton species. We compared the model predictions with a lake experiment, in which the turbulence structure of the entire lake was manipulated using artificial mixing. Vertical eddy diffusivities were calculated from the measured temperature microstructure in the lake. Changes in turbulent mixing of the lake caused a dramatic shift in phytoplankton species composition, consistent with the predictions of the competition model. The buoyant and potentially toxic cyanobacterium Microcystis dominated at low turbulent diffusivity, whereas sinking diatoms and green algae dominated at high turbulent diffusivity. These findings warn that changes in the turbulence structure of natural waters, for instance driven by climate change, may induce major shifts in the species composition of phytoplankton communities.

A HIERARCHICAL SET OF MODELS FOR SPECIES RESPONSE ANALYSIS

Variation in the abundance of species in space and/ or time can be caused by a wide range of underlying processes. Before such causes can be analysed we need simple math- ematical models which can describe the observed response patterns. For this purpose a hierarchical set of models is presented. These models are applicable to positive data with an upper bound, like relative frequencies and percentages. The models are fitted to the observations by means of logistic and non-linear regression techniques. Working with models of increasing complexity allows us to choose for the simplest possible model which sufficiently explains the observed pat- tern. The models are particularly suited for description of responses in time or over major environmental gradients. Deviations from these temporal or spatial trends may be statistically ascribed to, for example, climatic fluctuations or small-scale spatial heterogeneity. The applicability of this approach is illustrated by examples from recent research. A combination of simple, descriptive models like those pre- sented in this paper and causal models as developed by several others, is advocated as a powerful tool towards a fuller under- standing of the dynamics and patterns of vegetational change.

Global biodiversity patterns of marine phytoplankton and zooplankton

Although the oceans cover 70% of the Earths surface, our knowledge of biodiversity patterns in marine phytoplankton and zooplankton is very limited compared to that of the biodiversity of plants and herbivores in the terrestrial world. Here, we present biodiversity data for marine plankton assemblages from different areas of the world ocean. Similar to terrestrial vegetation, marine phytoplankton diversity is a unimodal function of phytoplankton biomass, with maximum diversity at intermediate levels of phytoplankton biomass and minimum diversity during massive blooms. Contrary to expectation, we did not find a relation between phytoplankton diversity and zooplankton diversity. Zooplankton diversity is a unimodal function of zooplankton biomass. Most strikingly, these marine biodiversity patterns show a worldwide consistency, despite obvious differences in environmental conditions of the various oceanographic regions. These findings may serve as a new benchmark in the search for global biodiversity patterns of plants and herbivores.

Chaos in a long-term experiment with a plankton community

Mathematical models predict that species interactions such as competition and predation can generate chaos. However, experimental demonstrations of chaos in ecology are scarce, and have been limited to simple laboratory systems with a short duration and artificial species combinations. Here, we present the first experimental demonstration of chaos in a long-term experiment with a complex food web. Our food web was isolated from the Baltic Sea, and consisted of bacteria, several phytoplankton species, herbivorous and predatory zooplankton species, and detritivores. The food web was cultured in a laboratory mesocosm, and sampled twice a week for more than 2,300 days. Despite constant external conditions, the species abundances showed striking fluctuations over several orders of magnitude. These fluctuations displayed a variety of different periodicities, which could be attributed to different species interactions in the food web. The population dynamics were characterized by positive Lyapunov exponents of similar magnitude for each species. Predictability was limited to a time horizon of 15–30 days, only slightly longer than the local weather forecast. Hence, our results demonstrate that species interactions in food webs can generate chaos. This implies that stability is not required for the persistence of complex food webs, and that the long-term prediction of species abundances can be fundamentally impossible.

Adaptive divergence in pigment composition promotes phytoplankton biodiversity

The dazzling diversity of the phytoplankton has puzzled biologists for decades. The puzzle has been enlarged rather than solved by the progressive discovery of new phototrophic microorganisms in the oceans, including picocyanobacteria, pico-eukaryotes, and bacteriochlorophyll-based and rhodopsin-based phototrophic bacteria. Physiological and genomic studies suggest that natural selection promotes niche differentiation among these phototrophic microorganisms, particularly with respect to their photosynthetic characteristics. We have analysed competition for light between two closely related picocyanobacteria of the Synechococcus group that we isolated from the Baltic Sea. One of these two has a red colour because it contains the pigment phycoerythrin, whereas the other is blue-green because it contains high contents of the pigment phycocyanin. Here we report theory and competition experiments that reveal stable coexistence of the two picocyanobacteria, owing to partitioning of the light spectrum. Further competition experiments with a third marine cyanobacterium, capable of adapting its pigment composition, show that this species persists by investing in the pigment that absorbs the colour not used by its competitors. These results demonstrate the adaptive significance of divergence in pigment composition of phototrophic microorganisms, which allows an efficient utilization of light energy and favours species coexistence.

How Do Sinking Phytoplankton Species Manage to Persist

Phytoplankton require light for photosynthesis. Yet, most phytoplankton species are heavier than water and therefore sink. How can these sinking species persist? Somehow, the answer should lie in the turbulent motion that redisperses sinking phytoplankton over the vertical water column. Here, we show, using a reaction‐advection‐diffusion equation of light‐limited phytoplankton, that there is a turbulence window sustaining sinking phytoplankton species in deep waters. If turbulent diffusion is too high, phytoplankton are mixed to great depths, and the depth‐averaged light conditions are too low to allow net positive population growth. Conversely, if turbulent diffusion is too low, sinking phytoplankton populations end up at the ocean floor and succumb in the dark. At intermediate levels of turbulent diffusion, however, phytoplankton populations can outgrow both mixing rates and sinking rates. In this way, the reproducing population as a whole can maintain a position in the well‐lit zone near the top of the water column, even if all individuals within the population have a tendency to sink. This theory unites earlier classic results by Sverdrup and Riley as well as our own recent findings and provides a new conceptual framework for the understanding of phytoplankton dynamics under the influence of mixing processes.

Fundamental unpredictability in multispecies competition

One of the central goals of ecology is to predict the distribution and abundance of organisms. Here, we show that, in ecosystems of high biodiversity, the outcome of multispecies competition can be fundamentally unpredictable. We consider a competition model widely applied in phytoplankton ecology and plant ecology in which multiple species compete for three resources. We show that this competition model may have several alternative outcomes, that the dynamics leading to these alternative outcomes may exhibit transient chaos, and that the basins of attraction of these alternative outcomes may have an intermingled fractal geometry. As a consequence of this fractal geometry, it is impossible to predict the winners of multispecies competition in advance.

Species Dynamics in Phytoplankton Blooms: Incomplete Mixing and Competition for Light

With the eutrophication of many freshwaters and coastal environments, phytoplankton blooms have become a common phenomenon. This article uses a reaction‐diffusion model to investigate the implications of mixing processes for the dynamics and species composition of phytoplankton blooms. The model identifies four key parameters for bloom development: incident light intensity, background turbidity, water column depth, and turbulent mixing rates. The model predicts that the turbulent mixing rate is a major determinant of the species composition of phytoplankton blooms. In well‐mixed environments, the species with lowest “critical light intensity” should become dominant. But at low mixing rates, the species with lowest critical light intensity is displaced if other species obtain a better position in the light gradient. Instead of a gradual change in species composition, the model predicts steep transitions between the dominance regions of the various species. The model predicts a low species diversity: phytoplankton blooms in eutrophic environments should be dominated by one or a few species only. The model predictions are consistent with laboratory competition experiments and many existing field data. We recommend examining competition in phytoplankton blooms under well‐controlled laboratory conditions, and we derive scaling rules that facilitate translation from the laboratory to the field.

Competition for light between toxic and nontoxic strains of the harmful cyanobacterium Microcystis

ABSTRACT The cyanobacterium Microcystis can produce microcystins, a family of toxins that are of major concern in water management. In several lakes, the average microcystin content per cell gradually declines from high levels at the onset of Microcystis blooms to low levels at the height of the bloom. Such seasonal dynamics might result from a succession of toxic to nontoxic strains. To investigate this hypothesis, we ran competition experiments with two toxic and two nontoxic Microcystis strains using light-limited chemostats. The population dynamics of these closely related strains were monitored by means of characteristic changes in light absorbance spectra and by PCR amplification of the rRNA internal transcribed spacer region in combination with denaturing gradient gel electrophoresis, which allowed identification and semiquantification of the competing strains. In all experiments, the toxic strains lost competition for light from nontoxic strains. As a consequence, the total microcystin concentrations in the competition experiments gradually declined. We did not find evidence for allelopathic interactions, as nontoxic strains became dominant even when toxic strains were given a major initial advantage. These findings show that, in our experiments, nontoxic strains of Microcystis were better competitors for light than toxic strains. The generality of this finding deserves further investigation with other Microcystis strains. The competitive replacement of toxic by nontoxic strains offers a plausible explanation for the gradual decrease in average toxicity per cell during the development of dense Microcystis blooms.