A fundamental goal of microbial ecology is to understand what determines the diversity, stability, and structure of microbial ecosystems. The microbial context poses special conceptual challenges because of the strong mutual influences between the microbes and their chemical environment through the consumption and production of metabolites. By analyzing a generalized consumer resource model that explicitly includes cross-feeding, stochastic colonization, and thermodynamics, we show that complex microbial communities generically exhibit a transition as a function of available energy fluxes from a "resource-limited" regime where community structure and stability is shaped by energetic and metabolic considerations to a diverse regime where the dominant force shaping microbial communities is the overlap between species' consumption preferences. These two regimes have distinct species abundance patterns, different functional profiles, and respond differently to environmental perturbations. Our model reproduces large-scale ecological patterns observed across multiple experimental settings such as nestedness and differential beta diversity patterns along energy gradients. We discuss the experimental implications of our results and possible connections with disorder-induced phase transitions in statistical physics.

/pdf/available-energy-fluxes-drive-a-transition-in-the-diversity-uedh8s8eqd.pdf

Available energy fluxes drive a transition in the diversity, stability, and functional structure of microbial communities.

Treatment with antibiotics is one of the most extreme perturbations to the human microbiome. Even standard courses of antibiotics dramatically reduce the microbiome’s diversity and can cause transitions to dysbiotic states. Conceptually, this is often described as a ‘stability landscape’: the microbiome sits in a landscape with multiple stable equilibria, and sufficiently strong perturbations can shift the microbiome from its normal equilibrium to another state. However, this picture is only qualitative and has not been incorporated in previous mathematical models of the effects of antibiotics. Here, we outline a simple quantitative model based on the stability landscape concept and demonstrate its success on real data. Our analytical impulse-response model has minimal assumptions with three parameters. We fit this model in a Bayesian framework to data from a previous study of the year-long effects of short courses of four common antibiotics on the gut and oral microbiomes, allowing us to compare parameters between antibiotics and microbiomes, and further validate our model using data from another study looking at the impact of a combination of last-resort antibiotics on the gut microbiome. Using Bayesian model selection we find support for a long-term transition to an alternative microbiome state after courses of certain antibiotics in both the gut and oral microbiomes. Quantitative stability landscape frameworks are an exciting avenue for future microbiome modelling.

/pdf/modelling-microbiome-recovery-after-antibiotics-using-a-3e56ithe6z.pdf

Modelling microbiome recovery after antibiotics using a stability landscape framework.

The biodiversity of food webs is composed of horizontal (i.e. within trophic levels) and vertical diversity (i.e. the number of trophic levels). Understanding their joint effect on stability is a key challenge. Theory mostly considers their individual effects and focuses on small perturbations near equilibrium in hypothetical food webs. Here, we study the joint effects of horizontal and vertical diversity on the stability of hypothetical (modelled) and empirical food webs. In modelled food webs, horizontal and vertical diversity increased and decreased stability, respectively, with a stronger positive effect of producer diversity on stability at higher consumer diversity. Experiments with an empirical plankton food web, where we manipulated horizontal and vertical diversity and measured stability from species interactions and from resilience against large perturbations, confirmed these predictions. Taken together, our findings highlight the need to conserve horizontal biodiversity at different trophic levels to ensure stability.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/ele.13282

Horizontal and vertical diversity jointly shape food web stability against small and large perturbations.

Sparse non-Hermitian random matrices arise in the study of disordered physical systems with asymmetric local interactions, and have applications ranging from neural networks to ecosystem dynamics. The spectral characteristics of these matrices provide crucial information on system stability and susceptibility, however, their study is greatly complicated by the twin challenges of a lack of symmetry and a sparse interaction structure. In this review we provide a concise and systematic introduction to the main tools and results in this field. We show how the spectra of sparse non-Hermitian matrices can be computed via an analogy with infinite dimensional operators obeying certain recursion relations. With reference to three illustrative examples --- adjacency matrices of regular oriented graphs, adjacency matrices of oriented Erdős-Renyi graphs, and adjacency matrices of weighted oriented Erdős-Renyi graphs --- we demonstrate the use of these methods to obtain both analytic and numerical results for the spectrum, the spectral distribution, the location of outlier eigenvalues, and the statistical properties of eigenvectors.

Spectral Theory of Sparse Non-Hermitian Random Matrices

We use dynamical generating functionals to study the stability and size of communities evolving in Lotka-Volterra systems with random interaction coefficients. The size of the eco-system is not set from the beginning. Instead, we start from a set of possible species, which may undergo extinction. How many species survive depends on the properties of the interaction matrix; the size of the resulting food web at stationarity is a property of the system itself in our model, and not a control parameter as in most studies based on random matrix theory. We find that prey-predator relations enhance stability, and that variability of species interactions promotes instability. Complexity of inter-species couplings leads to reduced sizes of ecological communities. Dynamically evolved community size and stability are hence positively correlated.

Dynamically evolved community size and stability of random Lotka-Volterra ecosystems

Random matrix theory successfully connects the structure of interactions of large ecological communities to their ability to respond to perturbations. One of the most debated aspects of this approach is that so far studies have neglected the role of population abundances on stability. While species abundances are well studied and empirically accessible, studies on stability have so far failed to incorporate this information. Here we tackle this question by explicitly including population abundances in a random matrix framework. We derive an analytical formula that describes the spectrum of a large community matrix for arbitrary feasible species abundance distributions. The emerging picture is remarkably simple: while population abundances affect the rate to return to equilibrium after a perturbation, the stability of large ecosystems is uniquely determined by the interaction matrix. We confirm this result by showing that the likelihood of having a feasible and unstable solution in the Lotka-Volterra system of equations decreases exponentially with the number of species for stable interaction matrices.

/pdf/effect-of-population-abundances-on-the-stability-of-large-t8o7wjl5re.pdf

Theo Gibbs

Papers

Effect of population abundances on the stability of large random ecosystems.