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Trait-based approaches have broadened our understanding of how the composition of ecological communities responds to environmental drivers. This research has mainly focussed on abiotic factors and competition determining the community trait distribution, while effects of trophic interactions on trait dynamics, if considered at all, have been studied for two trophic levels at maximum. However, natural food webs are typically at least tritrophic. This enables indirect interactions of traits and biomasses among multiple trophic levels leading to underexplored effects on food web dynamics. Here, we demonstrate the occurrence of mutual trait adjustment among three trophic levels in a natural plankton food web (Lake Constance) and in a corresponding mathematical model. We found highly recurrent seasonal biomass and trait dynamics, where herbivorous zooplankton increased its size, and thus its ability to counter phytoplankton defense, before phytoplankton defense actually increased. This is contrary to predictions from bitrophic systems where counter-defense of the consumer is a reaction to prey defense. In contrast, counter-defense of carnivores by size adjustment followed the defense of herbivores as expected. By combining observations and model simulations, we show how the reversed trait dynamics at the two lower trophic levels result from a "trophic biomass-trait cascade" driven by the carnivores. Trait adjustment between two trophic levels can therefore be altered by biomass or trait changes of adjacent trophic levels. Hence, analyses of only pairwise trait adjustment can be misleading in natural food webs, while multitrophic trait-based approaches capture indirect biomass-trait interactions among multiple trophic levels.
Functional traits determine biomass dynamics, coexistence and energetics in plankton food webs
(2022)
Plankton food webs are the basis of marine and limnetic ecosystems. Especially aquatic ecosystems of high biodiversity provide important ecosystem services for humankind as providers of food, coastal protection, climate regulation, and tourism. Understanding the dynamics of biomass and coexistence in these food webs is a first step to understanding the ecosystems. It also lays the foundation for the development of management strategies for the maintenance of the marine and freshwater biodiversity despite anthropogenic influences.
Natural food webs are highly complex, and thus often equally complex methods are needed to analyse and understand them well. Models can help to do so as they depict simplified parts of reality. In the attempt to get a broader understanding of the complex food webs, diverse methods are used to investigate different questions.
In my first project, we compared the energetics of a food chain in two versions of an allometric trophic network model. In particular, we solved the problem of unrealistically high trophic transfer efficiencies (up to 70%) by accounting for both basal respiration and activity respiration, which decreased the trophic transfer efficiency to realistic values of ≤30%. Next in my second project I turned to plankton food webs and especially phytoplankton traits. Investigating a long-term data set from Lake Constance we found evidence for a trade-off between defence and growth rate in this natural phytoplankton community. I continued working with this data set in my third project focusing on ciliates, the main grazer of phytoplankton in spring. Boosted regression trees revealed that temperature and predators have the highest influence on net growth rates of ciliates. We finally investigated in my fourth project a food web model inspired by ciliates to explore the coexistence of plastic competitors and to study the new concept of maladaptive switching, which revealed some drawbacks of plasticity: faster adaptation led to higher maladaptive switching towards undefended phenotypes which reduced autotroph biomass and coexistence and increased consumer biomass.
It became obvious that even well-established models should be critically questioned as it is important not to forget reality on the way to a simplistic model. The results showed furthermore that long-term data sets are necessary as they can help to disentangle complex natural processes. Last, one should keep in mind that the interplay between models and experiments/ field data can deliver fruitful insights about our complex world.
Phytoplankton growth depends not only on the mean intensity but also on the dynamics of the light supply. The nonlinear light-dependency of growth is characterized by a small number of basic parameters: the compensation light intensity PARcompμ, where production and losses are balanced, the growth efficiency at sub-saturating light αµ, and the maximum growth rate at saturating light µmax. In surface mixed layers, phytoplankton may rapidly move between high light intensities and almost darkness. Because of the different frequency distribution of light and/or acclimation processes, the light-dependency of growth may differ between constant and fluctuating light. Very few studies measured growth under fluctuating light at a sufficient number of mean light intensities to estimate the parameters of the growth-irradiance relationship. Hence, the influence of light dynamics on µmax, αµ and PARcompμ are still largely unknown. By extension, accurate modelling predictions of phytoplankton development under fluctuating light exposure remain difficult to make. This PhD thesis does not intend to directly extrapolate few experimental results to aquatic systems – but rather improving the mechanistic understanding of the variation of the light-dependency of growth under light fluctuations and effects on phytoplankton development.
In Lake TaiHu and at the Three Gorges Reservoir (China), we incubated phytoplankton communities in bottles placed either at fixed depths or moved vertically through the water column to mimic vertical mixing. Phytoplankton at fixed depths received only the diurnal changes in light (defined as constant light regime), while phytoplankton received rapidly fluctuating light by superimposing the vertical light gradient on the natural sinusoidal diurnal sunlight. The vertically moved samples followed a circular movement with 20 min per revolution, replicating to some extent the full overturn of typical Langmuir cells. Growth, photosynthesis, oxygen production and respiration of communities (at Lake TaiHu) were
measured. To complete these investigations, a physiological experiment was performed in the laboratory on a toxic strain of Microcystis aeruginosa (FACBH 1322) incubated under 20 min period fluctuating light. Here, we measured electron transport rates and net oxygen production at a much higher time resolution (single minute timescale).
The present PhD thesis provides evidence for substantial effects of fluctuating light on the eco-physiology of phytoplankton. Both experiments performed under semi-natural conditions in Lake TaiHu and at the Three Gorges Reservoir gave similar results. The significant decline in community growth efficiencies αµ under fluctuating light was caused for a great share by different frequency distribution of light intensities that shortened the effective daylength for production. The remaining gap in community αµ was attributed to species-specific photoacclimation mechanisms and to light-dependent respiratory losses. In contrast, community maximal growth rates µmax were similar between incubations at constant and fluctuating light. At daily growth saturating light supply, differences in losses for biosynthesis between the two light regimes were observed. Phytoplankton experiencing constant light suffered photo-inhibition - leading to photosynthesis foregone and additional respiratory costs for photosystems repair. On the contrary, intermittent exposure to low and high light intensities prevented photo-inhibition of mixed algae but forced them to develop alternative light strategy. They better harvested and exploited surface irradiance by enhancing their photosynthesis. In the laboratory, we showed that Microcystis aeruginosa increased its oxygen consumption by dark respiration in the light few minutes only after exposure to increasing light intensities. More, we proved that within a simulated Langmuir cell, the net production at saturating light and the compensation light intensity for production at limiting light are positively related. These results are best explained by an accumulation of photosynthetic products at increasing irradiance and mobilization of these fresh resources by rapid enhancement of dark respiration for maintenance and biosynthesis at decreasing irradiance. At the daily timescale, we showed that the enhancement of photosynthesis at high irradiance for biosynthesis of species increased their maintenance respiratory costs at limiting light. Species-specific growth at saturating light µmax and compensation light intensity for growth PARcompμ of species incubated in Lake TaiHu were positively related. Because of this species-specific physiological tradeoff, species displayed different light affinities to limiting and saturating light - thereby exhibiting a gleaner-opportunist tradeoff. In Lake TaiHu, we showed that inter-specific differences in light acquisition traits (µmax and PARcompμ) allowed coexis¬tence of species on a gradient of constant
light while avoiding competitive exclusion. More interestingly we demonstrated for the first time that vertical mixing (inducing fluctuating light supply for phytoplankton) may alter or even reverse the light utilization strategies of species within couple of days. The intra-specific variation in traits under fluctuating light increased the niche space for acclimated species, precluding competitive exclusion.
Overall, this PhD thesis contributes to a better understanding of phytoplankton eco-physiology under fluctuating light supply. This work could enhance the quality of predictions of phytoplankton development under certain weather conditions or climate change scenarios.
Trait-based approaches have become increasingly successful in community ecology. They assume that the distribution of functional traits within communities responds in a predictable way to alterations in environmental forcing and that strong forcing may accelerate such trait changes. We used high frequency measurements of phytoplankton to test these assumptions. We analyzed the seasonal and long-term dynamics of the community trait mean within a multi-dimensional trait space under alternating multifactorial environmental conditions. The community trait mean exhibited a distinct recurrent annual pattern that reflected minor changes in climate, herbivory and nutrients. Independent of early spring conditions, the community trait mean was repeatedly driven into a narrow confined area in the trait space under pronounced herbivory during the clear water phase. The speed of movement was highest at the onset and the relaxation of such strong unidirectional forcing. Thus, our data support the conceptual framework of trait-based ecology that alterations in environmental conditions are systematically tracked by adjustments in the dominant functional trait values and that the speed of trait changes depends on the kind and intensity of the selection pressure. Our approach provides a sensitive tool to detect small functional differences in the community related to subtle differences in forcing.
1. During the last couple of decades, invasive species have become a worldwide problem in many freshwater systems. Besides higher plants and animals, microbes, in particular the potentially toxic cyanobacterium Cylindrospermopsis raciborskii, has attracted increasing attention, due to its spread towards temperate zones of the northern and southern hemisphere. A number of advantageous functional traits and a high intraspecific plasticity have been suggested to explain its invasion success. 2. The aim of this study was to examine intraspecific functional trait variability in 12 different isolates of C.raciborskii originating from different lakes in an invaded region in Northeast Germany. We measured growth rate, C:N:P ratios, chlorophyll-a content and the abundance of heterocysts under nutrient-replete and phosphorus-limited conditions. Moreover, the isolate-specific morphology and grazing losses by an herbivorous rotifer, as a top-down force, were studied. 3. DNA fingerprinting revealed that all isolates were genetically different. C.raciborskii exhibited a large variability in all measured traits among isolates. The C:P, N:P and Chl-a:C ratios differed by a factor of two or more. The trait variability among isolates was higher under nutrient-replete conditions, except for the C:P ratio, which varied most during phosphorus limitation. The susceptibility to grazing, calculated as maximum ingestion rates of the rotifer Brachionus calyciflorus on C.raciborskii, varied most among isolates, but was not related to any of the measured physiological or morphological traits, i.e. no trade-off was found. 4. Ecological and genetic clustering did not match, indicating that the genetic relationship based on DNA fingerprinting did not cover ecological differences. 5. Our results show a high trait variability within locally occurring and partly co-occurring C.raciborskii isolates. No overall trade-offs between the measured functional traits were found. This demonstrates the ecological relevance of linking multiple traits, e.g. competitive and consumptive. Furthermore, this study emphasises the importance of analysing more than one strain of a species, as different strains show different trait values potentially relevant for their invasibility and the field of general trait-based ecology.
Trade-offs between functional traits are ubiquitous in nature and can promote species coexistence depending on their shape. Classic theory predicts that convex trade-offs facilitate coexistence of specialized species with extreme trait values (extreme species) while concave trade-offs promote species with intermediate trait values (intermediate species). We show here that this prediction becomes insufficient when the traits translate non-linearly into fitness which frequently occurs in nature, e.g., an increasing length of spines reduces grazing losses only up to a certain threshold resulting in a saturating or sigmoid trait-fitness function. We present a novel, general approach to evaluate the effect of different trade-off shapes on species coexistence. We compare the trade-off curve to the invasion boundary of an intermediate species invading the two extreme species. At this boundary, the invasion fitness is zero. Thus, it separates trait combinations where invasion is or is not possible. The invasion boundary is calculated based on measurable trait-fitness relationships. If at least one of these relationships is not linear, the invasion boundary becomes non-linear, implying that convex and concave trade-offs not necessarily lead to different coexistence patterns. Therefore, we suggest a new ecological classification of trade-offs into extreme-favoring and intermediate-favoring which differs from a purely mathematical description of their shape. We apply our approach to a well-established model of an empirical predator-prey system with competing prey types facing a trade-off between edibility and half-saturation constant for nutrient uptake. We show that the survival of the intermediate prey depends on the convexity of the trade-off. Overall, our approach provides a general tool to make a priori predictions on the outcome of competition among species facing a common trade-off in dependence of the shape of the trade-off and the shape of the trait-fitness relationships.
The importance of intraspecific trait variability for community dynamics and ecosystem functioning has been underappreciated. There are theoretical reasons for predicting that species that differ in intraspecific trait variability will also differ in their effects on ecosystem functioning, particularly in variable environments. We discuss whether species with greater trait variability are likely to exhibit greater temporal stability in their population dynamics, and under which conditions this might lead to stability in ecosystem functioning. Resolving this requires us to consider several questions. First, are species with high levels of variation for one trait equally variable in others? In particular, is variability in response and effects traits typically correlated? Second, what is the relative contribution of local adaptation and phenotypic plasticity to trait variability? If local adaptation dominates, then stability in function requires one of two conditions: (i) individuals of appropriate phenotypes present in the environment at high enough frequencies to allow for populations to respond rapidly to the changing environment, and (ii) high levels of dispersal and gene flow. While we currently lack sufficient information on the causes and distribution of variability in functional traits, filling in these key data gaps should increase our ability to predict how changing biodiversity will alter ecosystem functioning.
In most biodiversity studies, taxonomic diversity is the measure for the multiplicity of species and is often considered to represent functional diversity. However, trends in taxonomic diversity and functional diversity may differ, for example, when many functionally similar but taxonomically different species co-occur in a community. The differences between these diversity measures are of particular interest in diversity research for understanding diversity patterns and their underlying mechanisms. We analysed a temporally highly resolved 20-year time series of lake phytoplankton to determine whether taxonomic diversity and functional diversity exhibit similar or contrasting seasonal patterns. We also calculated the functional mean of the community in n-dimensional trait space for each sampling day to gain further insights into the seasonal dynamics of the functional properties of the community. We found an overall weak positive relationship between taxonomic diversity and functional diversity with a distinct seasonal pattern. The two diversity measures showed synchronous behaviour from early spring to mid-summer and a more complex and diverging relationship from autumn to late winter. The functional mean of the community exhibited a recurrent annual pattern with the most prominent changes before and after the clear-water phase. From late autumn to winter, the functional mean of the community and functional diversity were relatively constant while taxonomic diversity declined, suggesting competitive exclusion during this period. A further decline in taxonomic diversity concomitant with increasing functional diversity in late winter to early spring is seen as a result of niche diversification together with competitive exclusion. Under these conditions, several different sets of traits are suitable to thrive, but within one set of functional traits only one, or very few, morphotypes can persist. Taxonomic diversity alone is a weak descriptor of trait diversity in phytoplankton. However, the combined analysis of taxonomic diversity and functional diversity, along with the functional mean of the community, allows for deeper insights into temporal patterns of community assembly and niche diversification.
In ecology, biodiversity-ecosystem functioning (BEE) research has seen a shift in perspective from taxonomy to function in the last two decades, with successful application of trait-based approaches. This shift offers opportunities for a deeper mechanistic understanding of the role of biodiversity in maintaining multiple ecosystem processes and services. In this paper, we highlight studies that have focused on BEE of microbial communities with an emphasis on integrating trait-based approaches to microbial ecology. In doing so, we explore some of the inherent challenges and opportunities of understanding BEE using microbial systems. For example, microbial biologists characterize communities using gene phylogenies that are often unable to resolve functional traits. Additionally, experimental designs of existing microbial BEE studies are often inadequate to unravel BEE relationships. We argue that combining eco-physiological studies with contemporary molecular tools in a trait-based framework can reinforce our ability to link microbial diversity to ecosystem processes. We conclude that such trait-based approaches are a promising framework to increase the understanding of microbial BEE relationships and thus generating systematic principles in microbial ecology and more generally ecology.
1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored.
2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate).
3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR.
4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect.
5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.