Aquatic ecologists face challenges in identifying the general rules of the functioning of ecosystems. A common framework, including freshwater, marine, benthic, and pelagic ecologists is needed to bridge communication gaps and foster knowledge sharing. This framework should transcend local specificities and taxonomy in order to provide a common ground and shareable tools to address common scientific challenges. Here, we advocate the use of functional trait-based approaches (FTBAs) for aquatic ecologists, and propose concrete paths to go forward. Firstly, we propose to unify existing definitions in FTBAs to adopt a common language. Secondly, we list the numerous databases referencing functional traits for aquatic organisms. Thirdly, we present a synthesis on traditional as well as recent promising methods for the study of aquatic functional traits, including imaging and genomics. Finally, we conclude with a highlight on scientific challenges and promising venues for which FTBAs should foster opportunities for future research. By offering practical tools, our framework provides a clear path forward to the adoption of trait-based approaches in aquatic ecology.

/pdf/functional-trait-based-approaches-as-a-common-framework-for-2zpszi1sxq.pdf

Functional trait-based approaches as a common framework for aquatic ecologists

Phytoplankton is confronted with a variable assemblage of zooplankton grazers that create a strong selection pressure for traits that reduce mortality. Phytoplankton is, however, also challenged to remain suspended and to acquire sufficient resources for growth. Consequently, phytoplanktic organisms have evolved a variety of strategies to survive in a variable environment. An overview is presented of the various phytoplankton defense strategies, and costs and benefits of phytoplankton defenses with a zooming in on grazer-induced colony formation. The trade-off between phytoplankton competitive abilities and defenses against grazing favor adaptive trait changes—rapid evolution and phenotypic plasticity—that have the potential to influence population and community dynamics, as exemplified by controlled chemostat experiments. An interspecific defense–growth trade-off could explain seasonal shifts in the species composition of an in situ phytoplankton community yielding defense and growth rate as key traits of the phytoplankton. The importance of grazing and protection against grazing in shaping the phytoplankton community structure should not be underestimated. The trade-offs between nutrient acquisition, remaining suspended, and grazing resistance generate the dynamic phytoplankton community composition.

/pdf/grazing-resistance-in-phytoplankton-2pbutzqjxg.pdf

Grazing resistance in phytoplankton

Mixotrophy, the combination of autotrophic and heterotrophic nutrition, is a common trophic strategy among unicellular eukaryotes in the ocean. There are a number of hypotheses about the conditions that select for mixotrophy, and field studies have documented the prevalence of mixotrophy in a range of environments. However, there is currently little evidence for how mixotrophy varies across environmental gradients, and whether empirical patterns support theoretical predictions. Here I synthesize experiments that have quantified the abundance of phototrophic, mixotrophic, and heterotrophic nanoflagellates, to ask whether there are broad patterns in the prevalence of mixotrophy (relative to pure autotrophy and heterotrophy), and to ask whether observed patterns are consistent with a trait-based model of trophic strategies. The data suggest that mixotrophs increase in abundance at lower latitudes, while autotrophs and heterotrophs do not, and that this may be driven by increased light availability. Both mixotrophs and autotrophs increase greatly in productive coastal environments, while heterotrophs increase only slightly. These patterns are consistent with a model of resource competition in which nutrients and carbon can both limit growth and mixotrophs experience a trade-off in allocating biomass to phagotrophy vs. autotrophic functions. Importantly, mixotrophy is selected for under a range of conditions even when mixotrophs experience a penalty for using a generalist trophic strategy, due to the synergy between photosynthetically derived carbon and prey-derived nutrients. For this reason mixotrophy is favored relative to specialist strategies by increased irradiance, while at the same time increased nutrient supply increases the competitive ability of mixotrophs against heterotrophs.

https://www.pnas.org/content/pnas/116/13/6211.full.pdf

Mixotrophy in nanoflagellates across environmental gradients in the ocean.

Trade-offs constrain the improvement of performance of multiple traits simultaneously. Such trade-offs define Pareto fronts, which represent a set of optimal individuals that cannot be improved in any one trait without reducing performance in another. Surprisingly, experimental evolution often yields genotypes with improved performance in all measured traits, perhaps indicating an absence of trade-offs at least in the short term. Here we densely sample adaptive mutations in Saccharomyces cerevisiae to ask whether first-step adaptive mutations result in trade-offs during the growth cycle. We isolated thousands of adaptive clones evolved under carefully chosen conditions and quantified their performances in each part of the growth cycle. We too find that some first-step adaptive mutations can improve all traits to a modest extent. However, our dense sampling allowed us to identify trade-offs and establish the existence of Pareto fronts between fermentation and respiration, and between respiration and stationary phases. Moreover, we establish that no single mutation in the ancestral genome can circumvent the detected trade-offs. Finally, we sequenced hundreds of these adaptive clones, revealing new targets of adaptation and defining the genetic basis of the identified trade-offs.

Single nucleotide mapping of trait space reveals Pareto fronts that constrain adaptation

Both intra‐ and interspecific differences in traits may modulate interactions between plants. Two mechanisms are hypothesized to regulate these effects: competitive hierarchies and trait dissimilarities, but it is unclear how the prevalence of each might depend on environmental conditions and on intra and interspecific differences. We sowed six replicates of all possible pairwise combinations across eight annual species (including conspecific competition and individuals without competitors), in pots under two levels of fertilization. We measured above‐ and below‐ground traits and total biomass of the focal individuals. We expressed competition as the decrease in biomass of focal individuals relative to biomass without competition, and modelled competition using directional (hierarchy) or non‐directional (dissimilarity) trait differences, with or without intraspecific variability (ITV). We found evidence of different trait hierarchies operating under different fertilization conditions but little support for trait dissimilarities. The negative effect of competition on focal plants increased with the height of the competitors in both of fertilization levels. Further, in unfertilized conditions, plants with lower specific leaf area (SLA) and larger root systems experienced less competition. Including ITV in trait hierarchies substantially improved our ability to predict the intensity of competition. This was partly due to ITV reducing competitive hierarchies. Synthesis. Our results underscore the importance of traits in driving interactions among plants. Competitive relationships between species depend on complex interactions between trait intra and interspecific differences and resource availability. ITV appears to be a mechanism capable of reducing trait hierarchies, and hence the intensity of competition between coexisting plants.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1111/1365-2745.13248

Trait hierarchies and intraspecific variability drive competitive interactions in Mediterranean annual plants

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.

Trait-fitness relationships determine how trade-off shapes affect species coexistence.

Theory predicts that trade-offs, quantifying costs of functional trait adjustments, crucially affect community trait adaptation to altered environmental conditions, but empirical verification is scarce. We evaluated trait dynamics (antipredator defense, maximum growth rate, and phosphate affinity) of a lake phytoplankton community in a seasonally changing environment, using literature trait data and 21 years of species-resolved high-frequency biomass measurements. The trait data indicated a concave defense-growth trade-off, promoting fast-growing species with intermediate defense. With seasonally increasing grazing pressure, the community shifted toward higher defense levels at the cost of lower growth rates along the trade-off curve, while phosphate affinity explained some deviations from it. We discuss how low fitness differences of species, inferred from model simulations, in concert with stabilizing mechanisms, e.g., arising from further trait dimensions, may lead to the observed phytoplankton diversity. In conclusion, quantifying trade-offs is key for predictions of community trait adaptation and biodiversity under environmental change.

/pdf/the-shape-of-a-defense-growth-trade-off-governs-seasonal-um2apr3z06.pdf

The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton.

Functional trait compositions of communities can adapt to altered environmental conditions ensuring community persistence. Theory predicts that the shape of trade-offs between traits crucially affects these trait dynamics, but its empirical verification from the field is missing. Here, we show how the shape of a defense-growth trade-off governs seasonal trait dynamics of a natural community, using high-frequency, long-term measurements of phytoplankton from Lake Constance. As expected from the lab-derived concave trade-off curve, we observed an alternating dominance of several fast-growing species with intermediate defense levels and gradual changes of the biomass-trait distribution due to seasonally changing grazing pressure. By combining data and modelling, we obtain mechanistic insights on the underlying fitness landscape, and show that low fitness differences can maintain trait variation along the trade-off curve. We provide firm evidence for a frequently assumed trade-off and conclude that quantifying its shape allows to understand environmentally driven trait changes within communities.

https://www.biorxiv.org/content/biorxiv/early/2018/11/05/462622.full.pdf

The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton

https://aslopubs.onlinelibrary.wiley.com/doi/pdf/10.1002/lno.11466

Coupled changes in traits and biomasses cascading through a tritrophic plankton food web

It is well-known that prey species often face trade-offs between defense against predation and competitiveness, enabling predator-mediated coexistence. However, we lack an understanding of how the large variety of different defense traits with different competition costs affects coexistence and population dynamics. Our study focusses on two general defense mechanisms, that is, pre-attack (e.g., camouflage) and post-attack defenses (e.g., weaponry) that act at different phases of the predator—prey interaction. We consider a food web model with one predator, two prey types and one resource. One prey type is undefended, while the other one is pre- or post-attack defended paying costs either by a higher half-saturation constant for resource uptake or a lower maximum growth rate. We show that post-attack defenses promote prey coexistence and stabilize the population dynamics more strongly than pre-attack defenses by interfering with the predator's functional response: Because the predator spends time handling “noncrackable” prey, the undefended prey is indirectly facilitated. A high half-saturation constant as defense costs promotes coexistence more and stabilizes the dynamics less than a low maximum growth rate. The former imposes high costs at low resource concentrations but allows for temporally high growth rates at predator-induced resource peaks preventing the extinction of the defended prey. We evaluate the effects of the different defense mechanisms and costs on coexistence under different enrichment levels in order to vary the importance of bottom-up and top-down control of the prey community.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1002/ece3.4145

Not attackable or not crackable-How pre- and post-attack defenses with different competition costs affect prey coexistence and population dynamics.

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