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ABSTRACT Spatial processes, particularly scale‐dependent feedbacks, may play important and underappreciated roles in the dynamics of bistable ecosystems. For example, self‐organised spatial patterns can allow for stable coexistence of alternative states outside regions of bistability, a phenomenon known as a Busse balloon. We used partial differential equations to explore the potential for such dynamics in coral reefs, focusing on how herbivore behaviour and mobility affect the stability of coral‐ and macroalgal‐dominated states. Herbivore attraction to coral resulted in a Busse balloon that enhanced macroalgal resilience, with patterns persisting in regions of parameter space where nonspatial models predict uniform coral dominance. Thus, our work suggests herbivore association with coral (e.g., for shelter) can prevent reefs from reaching a fully coral‐dominated state. More broadly, this study illustrates how consumer space use can prevent ecosystems from undergoing wholesale state transitions, highlighting the importance of explicitly accounting for space when studying bistable systems. 

Mixotrophic protists combine photosynthesis with the ingestion of prey to thrive in resource-limited conditions in the ocean. Yet, how they fine-tune resource investments between their two different metabolic strategies remains unclear. Here, we present a modeling framework (Mixotroph Optimal Contributions to Heterotrophy and Autotrophy) that predicts the optimal (growth-maximizing) investments of carbon and nitrogen as a function of environmental conditions. Our model captures a full spectrum of trophic modes, in which the optimal investments reflect zero-waste solutions (i.e., growth is colimited by carbon and nitrogen) and accurately reproduces experimental results. By fitting the model to data forOchromonas, we were able to predict metabolic strategies at a global scale. We find that high phagotrophic investment is the dominant strategy across different oceanic biomes, used primarily for nitrogen acquisition. Our results therefore support empirical observations of the importance of mixotrophic grazers to upper ocean bacterivory. 

Protist plankton can be divided into three main groups: phytoplankton, zooplankton, and mixoplankton.In situmethods for studying phytoplankton and zooplankton are relatively straightforward since they generally target chlorophyll/photosynthesis or grazing activity, while the integration of both processes within a single cell makes mixoplankton inherently challenging to study. As a result, we understand less about mixoplankton physiology and their role in food webs, biogeochemical cycling, and ecosystems compared to phytoplankton and zooplankton. In this paper, we posit that by merging conventional techniques, such as microscopy and physiological data, with innovative methods likein situsingle-cell sorting and omics datasets, in conjunction with a diverse array of modeling approaches ranging from single-cell modeling to comprehensive Earth system models, we can propel mixoplankton research into the forefront of aquatic ecology. We present eight crucial research questions pertaining to mixoplankton and mixotrophy, and briefly outline a combination of existing methods and models that can be used to address each question. Our intent is to encourage more interdisciplinary research on mixoplankton, thereby expanding the scope of data acquisition and knowledge accumulation for this understudied yet critical component of aquatic ecosystems. 

Abstract Background Climate change is expected to lead to warming in ocean surface temperatures which will have unequal effects on the rates of photosynthesis and heterotrophy. As a result of this changing metabolic landscape, directional phenotypic evolution will occur, with implications that cascade up to the ecosystem level. While mixotrophic phytoplankton, organisms that combine photosynthesis and heterotrophy to meet their energetic and nutritional needs, are expected to become more heterotrophic with warmer temperatures due to heterotrophy increasing at a faster rate than photosynthesis, it is unclear how evolution will influence how these organisms respond to warmer temperatures. In this study, we used adaptive dynamics to model the consequences of temperature-mediated increases in metabolic rates for the evolution of mixotrophic phytoplankton, focusing specifically on phagotrophic mixotrophs. Results We find that mixotrophs tend to evolve to become more reliant on phagotrophy as temperatures rise, leading to reduced prey abundance through higher grazing rates. However, if prey abundance becomes too low, evolution favors greater reliance on photosynthesis. These responses depend upon the trade-off that mixotrophs experience between investing in photosynthesis and phagotrophy. Mixotrophs with a convex trade-off maintain mixotrophy over the greatest range of temperatures; evolution in these “generalist” mixotrophs was found to exacerbate carbon cycle impacts, with evolving mixotrophs exhibiting increased sensitivity to rising temperature. Conclusions Our results show that mixotrophs may respond more strongly to climate change than predicted by phenotypic plasticity alone due to evolutionary shifts in metabolic investment. However, the type of metabolic trade-off experienced by mixotrophs as well as ecological feedback on prey abundance may ultimately limit the extent of evolutionary change along the heterotrophy-phototrophy spectrum.