arXiv subjects

Simone Pigolotti

Publications and source records attributed to Simone Pigolotti.

At least 37 records · Page 2Linked to original sources

arXiv2019

Assembly of heteropolymers via a network of reaction coordinates

In biochemistry, heteropolymers encoding biological information are assembled out of equilibrium by sequentially incorporating available monomers found in the environment. Current models of polymerization treat monomer incorporation as a sequence of discrete chemical reactions between intermediate meta-stable states. In this paper, we use ideas from reaction rate theory and describe non-equilibrium assembly of a heteropolymer via a continuous reaction coordinate. Our approach allows to estimate the copy error and incorporation speed from the Gibbs free energy landscape of the process. We apply our theory to several examples, from a simple reaction characterized by a free energy barrier to more complex cases incorporating error correction mechanisms such as kinetic proofreading.

cond-mat.stat-mech↗

arXiv2019

Integral Fluctuation Relations for Entropy Production at Stopping Times

A stopping time $T$ is the first time when a trajectory of a stochastic process satisfies a specific criterion. In this paper, we use martingale theory to derive the integral fluctuation relation $\langle e^{-S_{\rm tot}(T)}\rangle=1$ for the stochastic entropy production $S_{\rm tot}$ in a stationary physical system at stochastic stopping times $T$. This fluctuation relation implies the law $\langle S_{\rm tot}(T)\rangle\geq 0$, which states that it is not possible to reduce entropy on average, even by stopping a stochastic process at a stopping time, and which we call the second law of thermodynamics at stopping times. This law implies bounds on the average amount of heat and work a system can extract from its environment when stopped at a random time. Furthermore, the integral fluctuation relation implies that certain fluctuations of entropy production are universal or are bounded by universal functions. These universal properties descend from the integral fluctuation relation by selecting appropriate stopping times: for example, when $T$ is a first-passage time for entropy production, then we obtain a bound on the statistics of negative records of entropy production. We illustrate these results on simple models of nonequilibrium systems described by Langevin equations and reveal two interesting phenomena. First, we demonstrate that isothermal mesoscopic systems can extract on average heat from their environment when stopped at a cleverly chosen moment and the second law at stopping times provides a bound on the average extracted heat. Second, we demonstrate that the average efficiency at stopping times of an autonomous stochastic heat engines, such as Feymann's ratchet, can be larger than the Carnot efficiency and the second law of thermodynamics at stopping times provides a bound on the average efficiency at stopping times.

cond-mat.stat-mech↗

arXiv2019

Error-speed correlations in biopolymer synthesis

Synthesis of biopolymers such as DNA, RNA, and proteins are biophysical processes aided by enzymes. Performance of these enzymes is usually characterized in terms of their average error rate and speed. However, because of thermal fluctuations in these single-molecule processes, both error and speed are inherently stochastic quantities. In this paper, we study fluctuations of error and speed in biopolymer synthesis and show that they are in general correlated. This means that, under equal conditions, polymers that are synthesized faster due to a fluctuation tend to have either better or worse errors than the average. The error-correction mechanism implemented by the enzyme determines which of the two cases holds. For example, discrimination in the forward reaction rates tends to grant smaller errors to polymers with faster synthesis. The opposite occurs for discrimination in monomer rejection rates. Our results provide an experimentally feasible way to identify error-correction mechanisms by measuring the error-speed correlations.

q-bio.SC↗

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

Mapping of uncertainty relations between continuous and discrete time

Lower bounds on fluctuations of thermodynamic currents depend on the nature of time: discrete or continuous. To understand the physical reason, we compare current fluctuations in discrete-time Markov chains and continuous-time master equations. We prove that current fluctuations in the master equations are always more likely, due to random timings of transitions. This comparison leads to a mapping of the moments of a current between discrete and continuous time. We exploit this mapping to obtain new uncertainty bounds. Our results reduce the quests for uncertainty bounds in discrete and continuous time to a single problem.

cond-mat.stat-mech↗

arXiv2018

Arcsine Laws in Stochastic Thermodynamics

We show that the fraction of time a thermodynamic current spends above its average value follows the arcsine law, a prominent result obtained by Lévy for Brownian motion. Stochastic currents with long streaks above or below their average are much more likely than those that spend similar fractions of time above and below their average. Our result is confirmed with experimental data from a Brownian Carnot engine. We also conjecture that two other random times associated with currents obey the arcsine law: the time a current reaches its maximum value and the last time a current crosses its average value. These results apply to, inter alia, molecular motors, quantum dots and colloidal systems.

cond-mat.stat-mech↗

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.

q-bio.PE↗

arXiv2017

Energetic funnel facilitates facilitated diffusion

Transcription factors are able to associate to their binding sites on DNA faster than the physical limit posed by diffusion. Such high association rates can be achieved by alternating between three-dimensional diffusion and one-dimensional sliding along the DNA chain, a mechanism dubbed Facilitated Diffusion. By studying a collection of transcription factor binding sites of Escherichia coli from the RegulonDB database and of Bacillus subtilis from DBTBS, we reveal a funnel in the binding energy landscape around the target sequences. We show that such a funnel is linked to the presence of gradients of AT in the base composition in the DNA region around the binding sites. An extensive computational study of the stochastic sliding process along the energetic landscapes obtained from the database shows that the funnel can significantly enhance the probability of transcription factors to find their target sequences when sliding in their proximity. We demonstrate that this enhancement leads to a speed-up of the association process.

q-bio.BM↗

arXiv2017

Generic Properties of Stochastic Entropy Production

We derive an Ito stochastic differential equation for entropy production in nonequilibrium Langevin processes. Introducing a random-time transformation, entropy production obeys a one-dimensional drift-diffusion equation, independent of the underlying physical model. This transformation allows us to identify generic properties of entropy production. It also leads to an exact uncertainty equality relating the Fano factor of entropy production and the Fano factor of the random time, which we also generalize to non steady-state conditions.

cond-mat.stat-mech↗

arXiv2016

Competition between fast- and slow-diffusing species in non-homogeneous environments

We study an individual-based model in which two spatially-distributed species, characterized by different diffusivities, compete for resources. We consider three different ecological settings. In the first, diffusing faster has a cost in terms of reproduction rate. In the second case, resources are not uniformly distributed in space. In the third case, the two species are transported by a fluid flow. In all these cases, at varying the parameters, we observe a transition from a regime in which diffusing faster confers an effective selective advantage to one in which it constitutes a disadvantage. We analytically estimate the magnitude of this advantage (or disadvantage) and test it by measuring fixation probabilities in simulations of the individual-based model. Our results provide a framework to quantify evolutionary pressure for increased or decreased dispersal in a given environment.

q-bio.PE↗

arXiv2016

Protocols for copying and proofreading in template-assisted polymerization

We discuss how information encoded in a template polymer can be stochastically copied into a copy polymer. We consider four different stochastic copy protocols of increasing complexity, inspired by building blocks of the mRNA translation pathway. In the first protocol, monomer incorporation occurs in a single stochastic transition. We then move to a more elaborate protocol in which an intermediate step can be used for error correction. Finally, we discuss the operating regimes of two kinetic proofreading protocols: one in which proofreading acts from the final copying step, and one in which it acts from an intermediate step. We review known results for these models and, in some cases, extend them to analyze all possible combinations of energetic and kinetic discrimination. We show that, in each of these protocols, only a limited number of these combinations leads to an improvement of the overall copying accuracy.

q-bio.SC↗

arXiv2015

Thermodynamics of Error Correction

Information processing at the molecular scale is limited by thermal fluctuations. This can cause undesired consequences in copying information since thermal noise can lead to errors that can compromise the functionality of the copy. For example, a high error rate during DNA duplication can lead to cell death. Given the importance of accurate copying at the molecular scale, it is fundamental to understand its thermodynamic features. In this paper, we derive a universal expression for the copy error as a function of entropy production and {\cred work dissipated by the system during wrong incorporations}. Its derivation is based on the second law of thermodynamics, hence its validity is independent of the details of the molecular machinery, be it any polymerase or artificial copying device. Using this expression, we find that information can be copied in three different regimes. In two of them, work is dissipated to either increase or decrease the error. In the third regime, the protocol extracts work while correcting errors, reminiscent of a Maxwell demon. As a case study, we apply our framework to study a copy protocol assisted by kinetic proofreading, and show that it can operate in any of these three regimes. We finally show that, for any effective proofreading scheme, error reduction is limited by the chemical driving of the proofreading reaction.

q-bio.SC↗

arXiv2015

Numerical and experimental study of the effects of noise on the permutation entropy

We analyze the effects of noise on the permutation entropy of dynamical systems. We take as numerical examples the logistic map and the Rössler system. Upon varying the noise strengthfaster, we find a transition from an almost-deterministic regime, where the permutation entropy grows slower than linearly with the pattern dimension, to a noise-dominated regime, where the permutation entropy grows faster than linearly with the pattern dimension. We perform the same analysis on experimental time-series by considering the stochastic spiking output of a semiconductor laser with optical feedback. Because of the experimental conditions, the dynamics is found to be always in the noise-dominated regime. Nevertheless, the analysis allows to detect regularities of the underlying dynamics. By comparing the results of these three different examples, we discuss the possibility of determining from a time series whether the underlying dynamics is dominated by noise or not.

physics.data-an↗

arXiv2015

Stochasticity enhances the gaining of bet-hedging strategies in contact-process-like dynamics

In biology and ecology, individuals or communities of individuals living in unpredictable environments often alternate between different evolutionary strategies to spread and reduce risks. Such behavior is commonly referred to as "bet-hedging." Long-term survival probabilities and population sizes can be much enhanced by exploiting such hybrid strategies. Here, we study the simplest possible birth-death stochastic model in which individuals can choose among a poor but safe strategy, a better but risky alternative, or a combination of both. We show analytically and computationally that the benefits derived from bet-hedging strategies are much stronger for higher environmental variabilities (large external noise) and/or for small spatial dimensions (large intrinsic noise). These circumstances are typically encountered by living systems, thus providing us with a possible justification for the ubiquitousness of bet-hedging in nature.

q-bio.PE↗

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↗

arXiv2014

Selective advantage of diffusing faster

We study a stochastic spatial model of biological competition in which two species have the same birth and death rates, but different diffusion constants. In the absence of this difference, the model can be considered as an off-lattice version of the Voter model and presents similar coarsening properties. We show that even a relative difference in diffusivity on the order of a few percent may lead to a strong bias in the coarsening process favoring the more agile species. We theoretically quantify this selective advantage and present analytical formulas for the average growth of the fastest species and its fixation probability.

q-bio.PE↗

arXiv2014

Analysis of self-overlap reveals trade-offs in plankton swimming trajectories

Movement is a fundamental behaviour of organisms that brings about beneficial encounters with resources and mates, but at the same time exposes the organism to dangerous encounters with predators. The movement patterns adopted by organisms should reflect a balance between these contrasting processes. This trade-off can be hypothesized as being evident in the behaviour of plankton, which inhabit a dilute 3D environment with few refuges or orienting landmarks. We present an analysis of the swimming path geometries based on a volumetric Monte Carlo sampling approach, which is particularly adept at revealing such trade-offs by measuring the self-overlap of the trajectories. Application of this method to experimentally measured trajectories reveals that swimming patterns in copepods are shaped to efficiently explore volumes at small scales, while achieving a large overlap at larger scales. Regularities in the observed trajectories make the transition between these two regimes always sharper than in randomized trajectories or as predicted by random walk theory. Thus real trajectories present a stronger separation between exploration for food and exposure to predators. The specific scale and features of this transition depend on species, gender, and local environmental conditions, pointing at adaptation to state and stage dependent evolutionary trade-offs.

q-bio.PE↗

arXiv2013

Species abundances and lifetimes: from neutral to niche-stabilized communities

We study a stochastic community model able to interpolate from a neutral regime to a niche partitioned regime upon varying a single parameter tuning the intensity of niche stabilization, namely the difference between intraspecific and interspecific competition. By means of a self-consistent approach, we obtain an analytical expression for the species abundance distribution, in excellent agreement with stochastic simulations of the model. In the neutral limit, the Fisher log-series is recovered, while upon increasing the stabilization strength the species abundance distribution develops a maximum for species at intermediate abundances, corresponding to the emergence of a carrying capacity. Numerical studies of species extinction-time distribution show that niche-stabilization strongly affects also the dynamical properties of the system by increasing the average species lifetimes, while suppressing their fluctuations. The results are discussed in view of the niche-neutral debate and of their potential relevance to field data.

q-bio.PE↗