arXiv subjects

Robert Marsland III

Publications and source records attributed to Robert Marsland III.

At least 19 recordsLinked to original sources

arXiv2024

Les Houches Lectures on Community Ecology: From Niche Theory to Statistical Mechanics

Ecosystems are among the most interesting and well-studied examples of self-organized complex systems. Community ecology, the study of how species interact with each other and the environment, has a rich tradition. Over the last few years, there has been a growing theoretical and experimental interest in these problems from the physics and quantitative biology communities. Here, we give an overview of community ecology, highlighting the deep connections between ecology and statistical physics. We start by introducing the two classes of mathematical models that have served as the workhorses of community ecology: Consumer Resource Models (CRM) and the generalized Lotka-Volterra models (GLV). We place a special emphasis on graphical methods and general principles. We then review recent works showing a deep and surprising connection between ecological dynamics and constrained optimization. We then shift our focus by analyzing these same models in "high-dimensions" (i.e. in the limit where the number of species and resources in the ecosystem becomes large) and discuss how such complex ecosystems can be analyzed using methods from the statistical physics of disordered systems such as the cavity method and Random Matrix Theory.

q-bio.PE↗

arXiv2023

Emergent competition shapes the ecological properties of multi-trophic ecosystems

Ecosystems are commonly organized into trophic levels -- organisms that occupy the same level in a food chain (e.g., plants, herbivores, carnivores). A fundamental question in theoretical ecology is how the interplay between trophic structure, diversity, and competition shapes the properties of ecosystems. To address this problem, we analyze a generalized Consumer Resource Model with three trophic levels using the zero-temperature cavity method and numerical simulations. We find that intra-trophic diversity gives rise to ``emergent competition'' between species within a trophic level due to feedbacks mediated by other trophic levels. This emergent competition gives rise to a crossover from a regime of top-down control (populations are limited by predators) to a regime of bottom-up control (populations are limited by primary producers) and is captured by a simple order parameter related to the ratio of surviving species in different trophic levels. We show that our theoretical results agree with empirical observations, suggesting that the theoretical approach outlined here can be used to understand complex ecosystems with multiple trophic levels.

q-bio.PE↗

arXiv2021

Cross-feeding shapes both competition and cooperation in microbial ecosystems

Recent work suggests that cross-feeding -- the secretion and consumption of metabolic biproducts by microbes -- is essential for understanding microbial ecology. Yet how cross-feeding and competition combine to give rise to ecosystem-level properties remains poorly understood. To address this question, we analytically analyze the Microbial Consumer Resource Model (MiCRM), a prominent ecological model commonly used to study microbial communities. Our mean-field solution exploits the fact that unlike replicas, the cavity method does not require the existence of a Lyapunov function. We use our solution to derive new species-packing bounds for diverse ecosystems in the presence of cross-feeding, as well as simple expressions for species richness and the abundance of secreted resources as a function of cross-feeding (metabolic leakage) and competition. Our results show how a complex interplay between competition for resources and cooperation resulting from metabolic exchange combine to shape the properties of microbial ecosystems.

q-bio.PE↗

arXiv2021

Diverse communities behave like typical random ecosystems

In 1972, Robert May triggered a worldwide research program studying ecological communities using random matrix theory. Yet, it remains unclear if and when we can treat real communities as random ecosystems. Here, we draw on recent progress in random matrix theory and statistical physics to extend May's approach to generalized consumer-resource models. We show that in diverse ecosystems adding even modest amounts of noise to consumer preferences results in a transition to "typicality" where macroscopic ecological properties of communities are indistinguishable from those of random ecosystems, even when resource preferences have prominent designed structures. We test these ideas using numerical simulations on a wide variety of ecological models. Our work offers an explanation for the success of random consumer-resource models in reproducing experimentally observed ecological patterns in microbial communities and highlights the difficulty of scaling up bottom-up approaches in synthetic ecology to diverse communities.

q-bio.PE↗

arXiv2020

Tregs self-organize into a "computing ecosystem" and implement a sophisticated optimization algorithm for mediating immune response

Regulatory T cells (Tregs) play a crucial role in mediating immune response. Yet an algorithmic understanding of the role of Tregs in adaptive immunity remains lacking. Here, we present a biophysically realistic model of Treg mediated self-tolerance in which Tregs bind to self-antigens and locally inhibit the proliferation of nearby activated T cells. By exploiting a duality between ecological dynamics and constrained optimization, we show that Tregs tile the potential antigen space while simultaneously minimizing the overlap between Treg activation profiles. We find that for sufficiently high Treg diversity, Treg mediated self-tolerance is robust to fluctuations in self-antigen concentrations but lowering the Treg diversity results in a sharp transition -- related to the Gardner transition in perceptrons -- to a regime where changes in self-antigen concentrations can result in an auto-immune response. We propose a novel experimental test of this transition in immune-deficient mice and discuss potential implications for autoimmune diseases.

q-bio.PE↗

arXiv2020

Effect of resource dynamics on species packing in diverse ecosystems

The competitive exclusion principle asserts that coexisting species must occupy distinct ecological niches (i.e. the number of surviving species can not exceed the number of resources). An open question is to understand if and how different resource dynamics affect this bound. Here, we analyze a generalized consumer resource model with externally supplied resources and show that -- in contrast to self-renewing resources -- species can occupy only half of all available environmental niches. This motivates us to construct a new schema for classifying ecosystems based on species packing properties.

q-bio.PE↗

arXiv2020

Niche theory for mutualism: A graphical approach to plant-pollinator network dynamics

Contemporary Niche Theory is a useful framework for understanding how organisms interact with each other and with their shared environment. Its graphical representation, popularized by Tilman's Resource Ratio Hypothesis, facilitates the analysis of the equilibrium structure of complex dynamical models including species coexistence. This theory has been applied primarily to resource competition since its early beginnings. Here, we integrate mutualism into niche theory by expanding Tilman's graphical representation to the analysis of consumer-resource dynamics of plant-pollinator networks. We graphically explain the qualitative phenomena previously found by numerical simulations, including the effects on community dynamics of nestedness, adaptive foraging, and pollinator invasions. Our graphical approach promotes the unification of niche and network theories, and deepens the synthesis of different types of interactions within a consumer-resource framework.

q-bio.PE↗

arXiv2020

Data-driven modeling reveals a universal dynamic underlying the COVID-19 pandemic under social distancing

We show that the COVID-19 pandemic under social distancing exhibits universal dynamics. The cumulative numbers of both infections and deaths quickly cross over from exponential growth at early times to a longer period of power law growth, before eventually slowing. In agreement with a recent statistical forecasting model by the IHME, we show that this dynamics is well described by the erf function. Using this functional form, we perform a data collapse across countries and US states with very different population characteristics and social distancing policies, confirming the universal behavior of the COVID-19 outbreak. We show that the predictive power of statistical models is limited until a few days before curves flatten, forecast deaths and infections assuming current policies continue and compare our predictions to the IHME models. We present simulations showing this universal dynamics is consistent with disease transmission on scale-free networks and random networks with non-Markovian transmission dynamics.

q-bio.PE↗

arXiv2020

The Community Simulator: A Python package for microbial ecology

Natural microbial communities contain hundreds to thousands of interacting species. For this reason, computational simulations are playing an increasingly important role in microbial ecology. In this manuscript, we present a new open-source, freely available Python package called Community Simulator for simulating microbial population dynamics in a reproducible, transparent and scalable way. The Community Simulator includes five major elements: tools for preparing the initial states and environmental conditions for a set of samples, automatic generation of dynamical equations based on a dictionary of modeling assumptions, random parameter sampling with tunable levels of metabolic and taxonomic structure, parallel integration of the dynamical equations, and support for metacommunity dynamics with migration between samples. To significantly speed up simulations using Community Simulator, our Python package implements a new Expectation-Maximization (EM) algorithm for finding equilibrium states of community dynamics that exploits a recently discovered duality between ecological dynamics and convex optimization. We present data showing that this EM algorithm improves performance by between one and two orders compared to direct numerical integration of the corresponding ordinary differential equations. We conclude by listing several recent applications of the Community Simulator to problems in microbial ecology, and discussing possible extensions of the package for directly analyzing microbiome compositional data.

q-bio.PE↗

arXiv2020

A minimal model for microbial biodiversity can reproduce experimentally observed ecological patterns

Surveys of microbial biodiversity such as the Earth Microbiome Project (EMP) and the Human Microbiome Project (HMP) have revealed robust ecological patterns across different environments. A major goal in ecology is to leverage these patterns to identify the ecological processes shaping microbial ecosystems. One promising approach is to use minimal models that can relate mechanistic assumptions at the microbe scale to community-level patterns. Here, we demonstrate the utility of this approach by showing that the Microbial Consumer Resource Model (MiCRM) -- a minimal model for microbial communities with resource competition, metabolic crossfeeding and stochastic colonization -- can qualitatively reproduce patterns found in survey data including compositional gradients, dissimilarity/overlap correlations, richness/harshness correlations, and nestedness of community composition. By using the MiCRM to generate synthetic data with different environmental and taxonomical structure, we show that large scale patterns in the EMP can be reproduced by considering the energetic cost of surviving in harsh environments and HMP patterns may reflect the importance of environmental filtering in shaping competition. We also show that recently discovered dissimilarity-overlap correlations in the HMP likely arise from communities that share similar environments rather than reflecting universal dynamics. We identify ecologically meaningful changes in parameters that alter or destroy each one of these patterns, suggesting new mechanistic hypotheses for further investigation. These findings highlight the promise of minimal models for microbial ecology.

q-bio.PE↗

arXiv2019

The Minimum Environmental Perturbation Principle: A New Perspective on Niche Theory

Fifty years ago, Robert MacArthur showed that stable equilibria optimize quadratic functions of the population sizes in several important ecological models. Here, we generalize this finding to a broader class of systems within the framework of contemporary niche theory, and precisely state the conditions under which an optimization principle (not necessarily quadratic) can be obtained. We show that conducting the optimization in the space of environmental states instead of population sizes leads to a universal and transparent physical interpretation of the objective function. Specifically, the equilibrium state minimizes the perturbation of the environment induced by the presence of the competing species, subject to the constraint that no species has a positive net growth rate. We use this "minimum environmental perturbation principle" to make new predictions for eco-evolution and community assembly, and describe a simple experimental setting where its conditions of validity have been empirically tested.

q-bio.PE↗

arXiv2019

Machine Learning as Ecology

Machine learning methods have had spectacular success on numerous problems. Here we show that a prominent class of learning algorithms - including Support Vector Machines (SVMs) -- have a natural interpretation in terms of ecological dynamics. We use these ideas to design new online SVM algorithms that exploit ecological invasions, and benchmark performance using the MNIST dataset. Our work provides a new ecological lens through which we can view statistical learning and opens the possibility of designing ecosystems for machine learning. Supplemental code is found at https://github.com/owenhowell20/EcoSVM.

cs.LG↗

arXiv2019

Active regeneration unites high- and low-temperature features in cooperative self-assembly

Cytoskeletal filaments are capable of self-assembly in the absence of externally supplied chemical energy, but the rapid turnover rates essential for their biological function require a constant flux of ATP or GTP hydrolysis. The same is true for two-dimensional protein assemblies employed in the formation of vesicles from cellular membranes, which rely on ATP-hydrolyzing enzymes to rapidly disassemble upon completion of the process. Recent observations suggest that the nucleolus, p granules and other three-dimensional membraneless organelles may also demand dissipation of chemical energy to maintain their fluidity. Cooperative binding plays a crucial role in the dynamics of these higher-dimensional structures, but is absent from classic models of 1-dimensional cytoskeletal assembly. In this Letter, we present a thermodynamically consistent model of actively regeneration with cooperative assembly, and compute the maximum turnover rate and minimum disassembly time as a function of the chemical driving force and the binding energy. We find that these driven structures resemble different equilibrium states above and below the nucleation barrier. In particular, we show that the maximal acceleration under large binding energies unites infinite-temperature local fluctuations with low-temperature nucleation kinetics.

physics.bio-ph↗

arXiv2019

The Thermodynamic Uncertainty Relation in Biochemical Oscillations

Living systems regulate many aspects of their behavior through periodic oscillations of molecular concentrations, which function as `biochemical clocks.' These clocks are intrinsically subject to thermal fluctuations, so that the duration of a full oscillation cycle is random. Their success in carrying out their biological function is thought to depend on the degree to which these fluctuations in the cycle period can be suppressed. Biochemical oscillators also require a constant supply of free energy in order to break detailed balance and maintain their cyclic dynamics. For a given free energy budget, the recently discovered `thermodynamic uncertainty relation' yields the magnitude of period fluctuations in the most precise conceivable free-running clock. In this paper, we show that computational models of real biochemical clocks severely underperform this optimum, with fluctuations several orders of magnitude larger than the theoretical minimum. We argue that this suboptimal performance is due to the small number of internal states per molecule in these models, combined with the high level of thermodynamic force required to maintain the system in the oscillatory phase. We introduce a new model with a tunable number of internal states per molecule, and confirm that it approaches the optimal precision as this number increases.

physics.bio-ph↗

arXiv2019

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

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.

physics.bio-ph↗

arXiv2018

Constrained optimization as ecological dynamics with applications to random quadratic programming in high dimensions

Quadratic programming (QP) is a common and important constrained optimization problem. Here, we derive a surprising duality between constrained optimization with inequality constraints -- of which QP is a special case -- and consumer resource models describing ecological dynamics. Combining this duality with a recent `cavity solution', we analyze high-dimensional, random QP where the optimization function and constraints are drawn randomly. Our theory shows remarkable agreement with numerics and points to a deep connection between optimization, dynamical systems, and ecology.

cond-mat.stat-mech↗

arXiv2017

Limits of Predictions in Thermodynamic Systems: A Review

The past twenty years have seen a resurgence of interest in nonequilibrium thermodynamics, thanks to advances in the theory of stochastic processes and in their thermodynamic interpretation. Fluctuation theorems provide fundamental constraints on the dynamics of systems arbitrarily far from thermal equilibrium. Thermodynamic uncertainty relations bound the dissipative cost of precision in a wide variety of processes. Concepts of excess work and excess heat provide the basis for a complete thermodynamics of nonequilibrium steady states, including generalized Clausius relations and thermodynamic potentials. But these general results carry their own limitations: fluctuation theorems involve exponential averages that can depend sensitively on unobservably rare trajectories; steady-state thermodynamics makes use of a dual dynamics that lacks any direct physical interpretation. This review aims to present these central results of contemporary nonequilibrium thermodynamics in such a way that the power of each claim for making physical predictions can be clearly assessed, using examples from current topics in soft matter and biophysics.

cond-mat.stat-mech↗

arXiv2015

Far-from-Equilibrium Distribution from Near-Steady-State Work Fluctuations

A longstanding goal of nonequilibrium statistical mechanics has been to extend the conceptual power of the Boltzmann distribution to driven systems. We report some new progress towards this goal. Instead of writing the nonequilibrium steady-state distribution in terms of perturbations around thermal equilibrium, we start from the linearized driven dynamics of observables about their stable fixed point, and expand in the strength of the nonlinearities encountered during typical fluctuations away from the fixed point. The first terms in this expansion retain the simplicity of known expansions about equilibrium, but can correctly describe the statistics of a certain class of systems even under strong driving. We illustrate this approach by comparison with a numerical simulation of a sheared Brownian colloid, where we find that the first two terms in our expansion are sufficient to account for the shear thinning behavior at high shear rates.

cond-mat.stat-mech↗