arXiv subjects

Miguel A. Muñoz

Publications and source records attributed to Miguel A. Muñoz.

At least 55 records · Page 3Linked to original sources

arXiv2018

Bet-hedging strategies in expanding populations

In ecology, species can mitigate their extinction risks in uncertain environments by diversifying individual phenotypes. This observation is quantified by the theory of bet-hedging, which provides a reason for the degree of phenotypic diversity observed even in clonal populations. The theory of bet-hedging in well-mixed populations is rather well developed. However, many species underwent range expansions during their evolutionary history, and the importance of phenotypic diversity in such scenarios still needs to be understood. In this paper, we develop a theory of bet-hedging for populations colonizing new, unknown environments that fluctuate either in space or time. In this case, we find that bet-hedging is a more favorable strategy than in well-mixed populations. For slow rates of variation, temporal and spatial fluctuations lead to different outcomes. In spatially fluctuating environments, bet-hedging is favored compared to temporally fluctuating environments. In the limit of frequent environmental variation, no opportunity for bet-hedging exists, regardless of the nature of the environmental fluctuations. For the same model, bet-hedging is never an advantageous strategy in the well-mixed case, supporting the view that range expansions strongly promote diversification. These conclusions are robust against stochasticity induced by finite population sizes. Our findings shed light on the importance of phenotypic heterogeneity in range expansions, paving the way to novel approaches to understand how biodiversity emerges and is maintained.

q-bio.PE↗

arXiv2018

Non-normality, reactivity, and intrinsic stochasticity in neural dynamics: a non-equilibrium potential approach

Intrinsic stochasticity can induce highly non-trivial effects on dynamical systems, including stochastic and coherence resonance, noise induced bistability, noise-induced oscillations, to name but a few. In this paper we revisit a mechanism first investigated in the context of neuroscience by which relatively small demographic (intrinsic) fluctuations can lead to the emergence of avalanching behavior in systems that are deterministically characterized by a single stable fixed point (up state). The anomalously large response of such systems to stochasticity stems (or is strongly associated with) the existence of a "non-normal" stability matrix at the deterministic fixed point, which may induce the system to be "reactive". Here, we further investigate this mechanism by exploring the interplay between non-normality and intrinsic (demographic) stochasticity, by employing a number of analytical and computational approaches. We establish, in particular, that the resulting dynamics in this type of systems cannot be simply derived from a scalar potential but, additionally, one needs to consider a curl flux which describes the essential non-equilibrium nature of this type of noisy non-normal systems. Moreover, we shed further light on the origin of the phenomenon, introduce the novel concept of "non-linear reactivity", and rationalize of the observed the value of the emerging avalanche exponents.

cond-mat.dis-nn↗

arXiv2018

Limited role of spatial self-structuring in emergent trade-offs during pathogen evolution

Pathogen transmission and virulence are main evolutionary variables broadly assumed to be linked through trade-offs. In well-mixed populations, these trade-offs are often ascribed to physiological restrictions, while populations with spatial self-structuring might evolve emergent trade-offs. Here, we reexamine a model of the latter kind proposed by Ballegooijen and Boerlijst with the aim of characterising the mechanisms causing the emergence of the trade-off and its structural robustness. Using invadability criteria, we establish the conditions under which an evolutionary feedback between transmission and virulence mediated by pattern formation can poise the system to a critical boundary separating a disordered state (without emergent trade-off) from a self-structured phase (where the trade-off emerges), and analytically calculate the functional shape of the boundary in a certain approximation. Beyond evolutionary parameters, the success of an invasion depends on the size and spatial structure of the invading and invaded populations. Spatial self-structuring is often destroyed when hosts are mobile, changing the evolutionary dynamics to those of a well-mixed population. In a metapopulation scenario, the systematic extinction of the pathogen in the disordered phase may counteract the disruptive effect of host mobility, favour pattern formation and therefore recover the emergent trade-off.

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arXiv2018

Landau-Ginzburg theory of cortex dynamics: Scale-free avalanches emerge at the edge of synchronization

Understanding the origin, nature, and functional significance of complex patterns of neural activity, as recorded by diverse electrophysiological and neuroimaging techniques, is a central challenge in neuroscience. Such patterns include collective oscillations emerging out of neural synchronization as well as highly heterogeneous outbursts of activity interspersed by periods of quiescence, called "neuronal avalanches." Much debate has been generated about the possible scale invariance or criticality of such avalanches and its relevance for brain function. Aimed at shedding light onto this, here we analyze the large-scale collective properties of the cortex by using a mesoscopic approach following the principle of parsimony of Landau-Ginzburg. Our model is similar to that of Wilson-Cowan for neural dynamics but crucially, includes stochasticity and space; synaptic plasticity and inhibition are considered as possible regulatory mechanisms. Detailed analyses uncover a phase diagram including down-state, synchronous, asynchronous, and up-state phases and reveal that empirical findings for neuronal avalanches are consistently reproduced by tuning our model to the edge of synchronization. This reveals that the putative criticality of cortical dynamics does not correspond to a quiescent-to-active phase transition as usually assumed in theoretical approaches but to a synchronization phase transition, at which incipient oscillations and scale-free avalanches coexist. Furthermore, our model also accounts for up and down states as they occur (e.g., during deep sleep). This approach constitutes a framework to rationalize the possible collective phases and phase transitions of cortical networks in simple terms, thus helping to shed light on basic aspects of brain functioning from a very broad perspective.

q-bio.NC↗

arXiv2017

Stochastic spatial models in ecology: a statistical physics approach

Ecosystems display a complex spatial organization. Ecologists have long tried to characterize them by looking at how different measures of biodiversity change across spatial scales. Ecological neutral theory has provided simple predictions accounting for general empirical patterns in communities of competing species. However, while neutral theory in well-mixed ecosystems is mathematically well understood, spatial models still present several open problems, limiting the quantitative understanding of spatial biodiversity. In this review, we discuss the state of the art in spatial neutral theory. We emphasize the connection between spatial ecological models and the physics of non-equilibrium phase transitions and how concepts developed in statistical physics translate in population dynamics, and vice versa. We focus on non-trivial scaling laws arising at the critical dimension D = 2 of spatial neutral models, and their relevance for biological populations inhabiting two-dimensional environments. We conclude by discussing models incorporating non-neutral effects in the form of spatial and temporal disorder, and analyze how their predictions deviate from those of purely neutral theories.

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arXiv2017

Impact of environmental colored noise in single-species population dynamics

Variability on external conditions has important consequences for the dynamics and the organization of biological systems. In many cases, the characteristic timescale of environmental changes as well as their correlations play a fundamental role in the way living systems adapt and respond to it. A proper mathematical approach to understand population dynamics, thus, requires approaches more refined than, e.g., simple white-noise approximations. To shed further light onto this problem, in this paper we propose a unifying framework based on different analytical and numerical tools available to deal with "colored" environmental noise. In particular, we employ a "unified colored noise approximation" to map the original problem into an effective one with white noise, and then we apply a standard path integral approach to gain analytical understanding. For the sake of specificity, we present our approach using as a guideline a variation of the contact process--which can also be seen as a birth-death process of the Malthus-Verhulst class--where the propagation or birth rate varies stochastically in time. Our approach allows us to tackle in a systematic manner some of the relevant questions concerning population dynamics under environmental variability, such as determining the stationary population density, establishing the conditions under which a population may become extinct, and estimating extinction times. We focus on the emerging phase diagram and its possible phase transitions, underlying how these are affected by the presence of environmental noise time-correlations.

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arXiv2017

Topological dimension tunes activity patterns in hierarchical modular network models

Connectivity patterns of relevance in neuroscience and systems biology can be encoded in hierarchical modular networks (HMNs). Moreover, recent studies highlight the role of hierarchical modular organization in shaping brain activity patterns, providing an excellent substrate to promote both the segregation and integration of neural information. Here we propose an extensive numerical analysis of the critical spreading rate (or "epidemic" threshold) --separating a phase with endemic persistent activity from one in which activity ceases-- on diverse HMNs. By employing analytical and computational techniques we determine the nature of such a threshold and scrutinize how it depends on general structural features of the underlying HMN. We critically discuss the extent to which current graph-spectral methods can be applied to predict the onset of spreading in HMNs, and we propose the network topological dimension as a relevant and unifying structural parameter, controlling the epidemic threshold.

cond-mat.dis-nn↗

arXiv2017

Neutral theory and scale-free neural dynamics

Avalanches of electrochemical activity in brain networks have been empirically reported to obey scale-invariant behavior --characterized by power-law distributions up to some upper cut-off-- both in vitro and in vivo. Elucidating whether such scaling laws stem from the underlying neural dynamics operating at the edge of a phase transition is a fascinating possibility, as systems poised at criticality have been argued to exhibit a number of important functional advantages. Here we employ a well-known model for neural dynamics with synaptic plasticity, to elucidate an alternative scenario in which neuronal avalanches can coexist, overlapping in time, but still remaining scale-free. Remarkably their scale-invariance does not stem from underlying criticality nor self-organization at the edge of a continuous phase transition. Instead, it emerges from the fact that perturbations to the system exhibit a neutral drift --guided by demographic fluctuations-- with respect to endogenous spontaneous activity. Such a neutral dynamics --similar to the one in neutral theories of population genetics-- implies marginal propagation of activity, characterized by power-law distributed causal avalanches. Importantly, our results underline the importance of considering causal information --on which neuron triggers the firing of which-- to properly estimate the statistics of avalanches of neural activity. We discuss the implications of these findings both in modeling and to elucidate experimental observations, as well as its possible consequences for actual neural dynamics and information processing in actual neural networks.

q-bio.NC↗

arXiv2017

A simple unified view of branching process statistics: random walks in balanced logarithmic potentials

We revisit the problem of deriving the mean-field values of avalanche critical exponents in systems with absorbing states. These are well-known to coincide with those of an un-biased branching process. Here, we show that for at least 4 different universality classes (directed percolation, dynamical percolation, the voter model or compact directed percolation class, and the Manna class of stochastic sandpiles) this common result can be obtained by mapping the corresponding Langevin equations describing each of these classes into a random walker confined close to the origin by a logarithmic potential. Many of the results derived here appear in the literature as independently derived for individual universality classes or for the branching process. However, the emergence of non-universal continuously-varying exponent values --which, as shown here, stems fro the presence of small external driving, that might induce avalanche merging-- has not been noticed (or emphasized) in the past. We believe that a simple an unified perspective as the one presented here can (i) help to clarify the overall picture, (ii) underline the super-universality of the behavior as well as the dependence on external driving, and (iii) help avoiding the common existing confusion between un-biased branching processes (equivalent to a random walker in a balanced logarithmic potential) and standard (un-confined) random walkers.

cond-mat.dis-nn↗

arXiv2016

Self-Organized Bistability Associated With First-Order Phase Transitions

Self-organized criticality elucidates the conditions under which physical and biological systems tune themselves to the edge of a second-order phase transition, with scale invariance. Motivated by the empirical observation of bimodal distributions of activity in neuroscience and other fields, we propose and analyze a theory for the self-organization to the point of phase-coexistence in systems exhibiting a first-order phase transition. It explains the emergence of regular avalanches with attributes of scale-invariance which coexist with huge anomalous ones, with realizations in many fields.

cond-mat.stat-mech↗

arXiv2016

Intrinsic noise and deviations from criticality in Boolean gene-regulatory networks

Gene regulatory networks can be successfully modeled as Boolean networks. A much discussed hypothesis says that such model networks reproduce empirical findings the best if they are tuned to operate at criticality, i.e. at the borderline between their ordered and disordered phases. Critical networks have been argued to lead to a number of functional advantages such as maximal dynamical range, maximal sensitivity to environmental changes, as well as to an excellent trade off between stability and flexibility. Here, we study the effect of noise within the context of Boolean networks trained to learn complex tasks under supervision. We verify that quasi-critical networks are the ones learning in the fastest possible way --even for asynchronous updating rules-- and that the larger the task complexity the smaller the distance to criticality. On the other hand, when additional sources of intrinsic noise in the network states and/or in its wiring pattern are introduced, the optimally performing networks become clearly subcritical. These results suggest that in order to compensate for inherent stochasticity, regulatory and other type of biological networks might become subcritical rather than being critical, all the most if the task to be performed has limited complexity.

q-bio.MN↗

arXiv2016

Complex synchronization patterns in the human connectome network

A major challenge in neuroscience is posed by the need for relating the emerging dynamical features of brain activity with the underlying modular structure of neural connections, hierarchically organized throughout several scales. The spontaneous emergence of coherence and synchronization across such scales is crucial to neural function, while its anomalies often relate to pathological conditions. Here we provide a numerical study of synchronization dynamics in the human connectome network. Our purpose is to provide a detailed characterization of the recently uncovered broad dynamic regime, interposed between order and disorder, which stems from the hierarchical modular organization of the human connectome. In this regime -similar in essence to a Griffiths phase- synchronization dynamics are trapped within metastable attractors of local coherence. Here we explore the role of noise, as an effective description of external perturbations, and discuss how its presence accounts for the ability of the system to escape intermittently from such attractors and explore complex dynamic repertoires of locally coherent states, in analogy with experimentally recorded patterns of cerebral activity.

q-bio.NC↗

arXiv2016

Intervality and coherence in complex networks

Food webs -- networks of predators and prey -- have long been known to exhibit "intervality": species can generally be ordered along a single axis in such a way that the prey of any given predator tend to lie on unbroken compact intervals. Although the meaning of this axis -- identified with a "niche" dimension -- has remained a mystery, it is assumed to lie at the basis of the highly non-trivial structure of food webs. With this in mind, most trophic network modelling has for decades been based on assigning species a niche value by hand. However, we argue here that intervality should not be considered the cause but rather a consequence of food-web structure. First, analysing a set of $46$ empirical food webs, we find that they also exhibit {\it predator} intervality: the predators of any given species are as likely to be contiguous as the prey are, but in a different ordering. Furthermore, this property is not exclusive of trophic networks: several networks of genes, neurons, metabolites, cellular machines, airports, and words are found to be approximately as interval as food webs. We go on to show that a simple model of food-web assembly which does not make use of a niche axis can nevertheless generate significant intervality. Therefore, the niche dimension (in the sense used for food-web modelling) could in fact be the consequence of other, more fundamental structural traits, such as trophic coherence. We conclude that a new approach to food-web modelling is required for a deeper understanding of ecosystem assembly, structure and function, and propose that certain topological features thought to be specific of food webs are in fact common to many complex networks.

q-bio.PE↗

arXiv2015

A novel brain partition highlights the modular skeleton shared by structure and function

Elucidating the intricate relationship between brain structure and function, both in healthy and pathological conditions, is a key challenge for modern neuroscience. Recent technical and methodological progress in neuroimaging has helped advance our understanding of this important issue, with diffusion weighted images providing information about structural connectivity (SC) and functional magnetic resonance imaging shedding light on resting state functional connectivity (rsFC). However, comparing these two distinct datasets, each of which can be encoded into a different complex network, is by no means trivial as pairwise link-to-link comparisons represent a relatively restricted perspective and provide only limited information. Thus, we have adopted a more integrative systems approach, exploiting theoretical graph analyses to study both SC and rsFC datasets gathered independently from healthy human subjects. The aim is to find the main architectural traits shared by the structural and functional networks by paying special attention to their common hierarchical modular organization. This approach allows us to identify a common skeleton from which a new, optimal, brain partition can be extracted, with modules sharing both structure and function. We describe these emerging common structure-function modules (SFMs) in detail. In addition, we compare SFMs with the classical Resting State Networks derived from independent component analysis of rs-fMRI functional activity, as well as with anatomical parcellations in the Automated Anatomical Labeling atlas and with the Broadmann partition, highlighting their similitude and differences. The unveiling of SFMs brings to light the strong correspondence between brain structure and resting-state dynamics.

q-bio.NC↗

arXiv2015

Inherent directionality explains the lack of feedback loops in empirical networks

We explore the hypothesis that the relative abundance of feedback loops in many empirical complex networks is severely reduced owing to the presence of an inherent global directionality. Aimed at quantifying this idea, we propose a simple probabilistic model in which a free parameter $γ$ controls the degree of inherent directionality. Upon strengthening such directionality, the model predicts a drastic reduction in the fraction of loops which are also feedback loops. To test this prediction, we extensively enumerated loops and feedback loops in many empirical biological, ecological and socio- technological directed networks. We show that, in almost all cases, empirical networks have a much smaller fraction of feedback loops than network randomizations. Quite remarkably, this empirical finding is quantitatively reproduced, for all loop lengths, by our model by fitting its only parameter $γ$. Moreover, the fitted value of $γ$ correlates quite well with another direct measurement of network directionality, performed by means of a novel algorithm. We conclude that the existence of an inherent network directionality provides a parsimonious quantitative explanation for the observed lack of feedback loops in empirical networks.

q-bio.MN↗

arXiv2015

Ranking species in mutualistic networks

Understanding the architectural subtleties of ecological networks, believed to confer them enhanced stability and robustness, is a subject of outmost relevance. Mutualistic interactions have been profusely studied and their corresponding bipartite networks, such as plant-pollinator networks, have been reported to exhibit a characteristic "nested" structure. Assessing the importance of any given species in mutualistic networks is a key task when evaluating extinction risks and possible cascade effects. Inspired in a recently introduced algorithm --similar in spirit to Google's PageRank but with a built-in non-linearity-- here we propose a method which --by exploiting their nested architecture-- allows us to derive a sound ranking of species importance in mutualistic networks. This method clearly outperforms other existing ranking schemes and can become very useful for ecosystem management and biodiversity preservation, where decisions on what aspects of ecosystems to explicitly protect need to be made.

q-bio.PE↗

arXiv2014

Coexistence in neutral theories: interplay of criticality and mild local preferences

Neutral theories have played a crucial and revolutionary role in fields such as population genetics and biogeography. These theories are critical by definition, in the sense that the overall growth rate of each single allele/species/type vanishes. Thus each species in a neutral model sits at the edge between invasion and extinction, allowing for the coexistence of symmetric/neutral types. However, in finite systems, mono-dominated states are ineludibly reached in relatively short times owing to demographic fluctuations, thus leaving us with an unsatisfactory framework to rationalize empirically-observed long-term coexistence. Here, we scrutinize the effect of heterogeneity in quasi-neutral theories, in which there can be a local mild preference for some of the competing species at some sites, even if the overall species symmetry is maintained. As we show here, mild biases at a small fraction of locations suffice to induce overall robust and durable species coexistence, even in regions arbitrarily far apart from the biased locations. This result stems from the long-range nature of the underlying critical bulk dynamics and has a number of implications, for example, in conservation ecology as it suggests that constructing local specific "sanctuaries" for different competing species can result in global enhancement of biodiversity, even in regions arbitrarily distant from the protected refuges.

cond-mat.stat-mech↗

arXiv2014

Trophic coherence determines food-web stability

Why are large, complex ecosystems stable? Both theory and simulations of current models predict the onset of instability with growing size and complexity, so for decades it has been conjectured that ecosystems must have some unidentified structural property exempting them from this outcome. We show that 'trophic coherence' -- a hitherto ignored feature of food webs which current structural models fail to reproduce -- is a better statistical predictor of linear stability than size or complexity. Furthermore, we prove that a maximally coherent network with constant interaction strengths will always be linearly stable. We also propose a simple model which, by correctly capturing the trophic coherence of food webs, accurately reproduces their stability and other basic structural features. Most remarkably, our model shows that stability can increase with size and complexity. This suggests a key to May's Paradox, and a range of opportunities and concerns for biodiversity conservation.

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