Population regulation and adaptive dynamics of cross-feeding.

The particular importance of evolutionary studies in microbial experimental systems is that starting from the level of the metabolism of individual cells, the adaptive dynamics can be followed step by step by biochemical, genetic, and population dynamical tools. Moreover, the coincidence of evolutionary and ecological time scales helps to clarify the mutual role of ecological and evolutionary principles in predicting adaptive dynamics in general. Ecological principles define the ecological conditions under which adaptive branching can occur. This paper overviews and interprets the results of empirical and modeling studies of the evolution of metabolic cross-feeding in glucose-limited E.coli chemostats and batch cultures in the context of theories of robust coexistence and adaptive dynamics. Empirical results consistently demonstrate that the interactions between cells are mediated by the changing metabolite concentrations in the cultures and modeling confirms that these changes may control the adaptive dynamics of the clones. In consequence, the potential results of evolution can be predicted at the functional level by evolutionary flux balance analysis (evoFBA), while the genetic changes are more contingent. evoFBA follows the scheme of adaptive dynamics theory by calculating the feedback environment that changes during the evolutionary process and provides a promising tool to further investigate adaptive divergence in small microbial communities. Three general conclusions close the paper.

Competitive Exclusion and Evolution: Convergence Almost Never Produces Ecologically Equivalent Species: (A Comment on McPeek, "Limiting Similarity? The Ecological Dynamics of Natural Selection among Resources and Consumers Caused by Both Apparent and Resource Competition").

In a recent modeling study ("Limiting Similarity? The Ecological Dynamics of Natural Selection among Resources and Consumers Caused by Both Apparent and Resource Competition") that appeared in the April 2019 issue of The American Naturalist, Mark A. McPeek argued that ecologically equivalent species may emerge via competition-induced trait convergence, in conflict with naive expectations based on the limiting similarity principle. Although the emphasis on the possibility of the convergence of competitors is very timely, here we show that the proposed mechanism will only lead to actual coexistence in the converged state for specially chosen fine-tuned parameter settings. It is therefore not a robust mechanism for the evolution of ecologically equivalent species. We conclude that invoking trait convergence as an explanation for the co-occurrence of seemingly fully equivalent species in nature would be premature.

Sensitivity analysis of coexistence in ecological communities: theory and application.

Sensitivity analysis, the study of how ecological variables of interest respond to changes in external conditions, is a theoretically well-developed and widely applied approach in population ecology. Though the application of sensitivity analysis to predicting the response of species-rich communities to disturbances also has a long history, derivation of a mathematical framework for understanding the factors leading to robust coexistence has only been a recent undertaking. Here we suggest that this development opens up a new perspective, providing advances ranging from the applied to the theoretical. First, it yields a framework to be applied in specific cases for assessing the extinction risk of community modules in the face of environmental change. Second, it can be used to determine trait combinations allowing for coexistence that is robust to environmental variation, and limits to diversity in the presence of environmental variation, for specific community types. Third, it offers general insights into the nature of communities that are robust to environmental variation. We apply recent community-level extensions of mathematical sensitivity analysis to example models for illustration. We discuss the advantages and limitations of the method, and some of the empirical questions the theoretical framework could help answer.

Shared developmental roles and transcriptional control of autophagy and apoptosis in Caenorhabditis elegans.

Autophagy is a lysosome-mediated self-degradation process of eukaryotic cells that, depending on the cellular milieu, can either promote survival or act as an alternative mechanism of programmed cell death (PCD) in terminally differentiated cells. Despite the important developmental and medical implications of autophagy and the main form of PCD, apoptosis, orchestration of their regulation remains poorly understood. Here, we show in the nematode Caenorhabditis elegans, that various genetic and pharmacological interventions causing embryonic lethality trigger a massive cell death response that has both autophagic and apoptotic features. The two degradation processes are also redundantly required for normal development and viability in this organism. Furthermore, the CES-2-like basic region leucine-zipper (bZip) transcription factor ATF-2, an upstream modulator of the core apoptotic cell death pathway, is able to directly regulate the expression of at least two key autophagy-related genes, bec-1/ATG6 and lgg-1/ATG8. Thus, the two cell death mechanisms share a common method of transcriptional regulation. Together, these results imply that under certain physiological and pathological conditions, autophagy and apoptosis are co-regulated to ensure the proper morphogenesis and survival of the developing organism. The identification of apoptosis and autophagy as compensatory cellular pathways in C. elegans might help us to understand how dysregulated PCD in humans can lead to diverse pathologies, including cancer, neurodegeneration and diabetes.

Competitive exclusion and limiting similarity: a unified theory.

Robustness of coexistence against changes of parameters is investigated in a model-independent manner by analyzing the feedback loop of population regulation. We define coexistence as a fixed point of the community dynamics with no population having zero size. It is demonstrated that the parameter range allowing coexistence shrinks and disappears when the Jacobian of the dynamics decreases to zero. A general notion of regulating factors/variables is introduced. For each population, its impact and sensitivity niches are defined as the differential impact on, and the differential sensitivity towards, the regulating variables, respectively. Either the similarity of the impact niches or the similarity of the sensitivity niches results in a small Jacobian and in a reduced likelihood of coexistence. For the case of a resource continuum, this result reduces to the usual "limited niche overlap" picture for both kinds of niche. As an extension of these ideas to the coexistence of infinitely many species, we demonstrate that Roughgarden's example for coexistence of a continuum of populations is structurally unstable.