Spatiotemporal organization of ant foraging from a complex systems perspective.

We use complex systems science to explore the emergent behavioral patterns that typify eusocial species, using collective ant foraging as a paradigmatic example. Our particular aim is to provide a methodology to quantify how the collective orchestration of foraging provides functional advantages to ant colonies. For this, we combine (i) a purpose-built experimental arena replicating ant foraging across realistic spatial and temporal scales, and (ii) a set of analytical tools, grounded in information theory and spin-glass approaches, to explore the resulting data. This combined approach yields computational replicas of the colonies; these are high-dimensional models that store the experimental foraging patterns through a training process, and are then able to generate statistically similar patterns, in an analogous way to machine learning tools. These in silico models are then used to explore the colony performance under different resource availability scenarios. Our findings highlight how replicas of the colonies trained under constant and predictable experimental food conditions exhibit heightened foraging efficiencies, manifested in reduced times for food discovery and gathering, and accelerated transmission of information under similar conditions. However, these same replicas demonstrate a lack of resilience when faced with new foraging conditions. Conversely, replicas of colonies trained under fluctuating and uncertain food conditions reveal lower efficiencies at specific environments but increased resilience to shifts in food location.

Intermittent random walks under stochastic resetting.

We analyze a one-dimensional intermittent random walk on an unbounded domain in the presence of stochastic resetting. In this process, the walker alternates between local intensive search, diffusion, and rapid ballistic relocations in which it does not react to the target. We demonstrate that Poissonian resetting leads to the existence of a non-equilibrium steady state. We calculate the distribution of the first arrival time to a target along with its mean and show the existence of an optimal reset rate. In particular, we prove that the initial condition of the walker, i.e., either starting diffusely or relocating, can significantly affect the long-time properties of the search process. Moreover, we demonstrate the presence of distinct parameter regimes for the global optimization of the mean first arrival time when ballistic and diffusive movements are in direct competition.

Random Walks on Comb-like Structures under Stochastic Resetting.

We study the long-time dynamics of the mean squared displacement of a random walker moving on a comb structure under the effect of stochastic resetting. We consider that the walker's motion along the backbone is diffusive and it performs short jumps separated by random resting periods along fingers. We take into account two different types of resetting acting separately: global resetting from any point in the comb to the initial position and resetting from a finger to the corresponding backbone. We analyze the interplay between the waiting process and Markovian and non-Markovian resetting processes on the overall mean squared displacement. The Markovian resetting from the fingers is found to induce normal diffusion, thereby minimizing the trapping effect of fingers. In contrast, for non-Markovian local resetting, an interesting crossover with three different regimes emerges, with two of them subdiffusive and one of them diffusive. Thus, an interesting interplay between the exponents characterizing the waiting time distributions of the subdiffusive random walk and resetting takes place. As for global resetting, its effect is even more drastic as it precludes normal diffusion. Specifically, such a resetting can induce a constant asymptotic mean squared displacement in the Markovian case or two distinct regimes of subdiffusive motion in the non-Markovian case.

Informational Entropy Threshold as a Physical Mechanism for Explaining Tree-like Decision Making in Humans.

While approaches based on physical grounds (such as the drift-diffusion model-DDM) have been exhaustively used in psychology and neuroscience to describe perceptual decision making in humans, similar approaches to complex situations, such as sequential (tree-like) decisions, are still scarce. For such scenarios that involve a reflective prospection of future options, we offer a plausible mechanism based on the idea that subjects can carry out an internal computation of the uncertainty about the different options available, which is computed through the corresponding Shannon entropy. When the amount of information gathered through sensory evidence is enough to reach a given threshold in the entropy, this will trigger the decision. Experimental evidence in favor of this entropy-based mechanism was provided by exploring human performance during navigation through a maze on a computer screen monitored with the help of eye trackers. In particular, our analysis allows us to prove that (i) prospection is effectively used by humans during such navigation tasks, and an indirect quantification of the level of prospection used is attainable; in addition, (ii) the distribution of decision times during the task exhibits power-law tails, a feature that our entropy-based mechanism is able to explain, unlike traditional (DDM-like) frameworks.

Conditioned backward and forward times of diffusion with stochastic resetting: A renewal theory approach.

Stochastic resetting can be naturally understood as a renewal process governing the evolution of an underlying stochastic process. In this work, we formally derive well-known results of diffusion with resets from a renewal theory perspective. Parallel to the concepts from renewal theory, we introduce the conditioned backward B and forward F times being the times since the last and until the next reset, respectively, given that the current state of the system X(t) is known. These magnitudes are introduced with the paradigmatic case of diffusion under resetting, for which the backward and forward times are conditioned to the position of the walker. We find analytical expressions for the conditioned backward and forward time probability density functions (PDFs), and we compare them with numerical simulations. The general expressions allow us to study particular scenarios. For instance, for power-law reset time PDFs such that φ(t)∼t^{-1-α}, significant changes in the properties of the conditioned backward and forward times happen at half-integer values of α due to the composition between the long-time scaling of diffusion P(x,t)∼1/sqrt[t] and the reset time PDF.

Nonstandard diffusion under Markovian resetting in bounded domains.

We consider a walker moving in a one-dimensional interval with absorbing boundaries under the effect of Markovian resettings to the initial position. The walker's motion follows a random walk characterized by a general waiting time distribution between consecutive short jumps. We investigate the existence of an optimal reset rate, which minimizes the mean exit passage time, in terms of the statistical properties of the waiting time probability. Generalizing previous results, we find that when the waiting time probability has first and second finite moments, resetting can be either (i) never beneficial, (ii) beneficial depending on the distance of the reset point to the boundary, or (iii) always beneficial. Instead, when the waiting time probability has the first or the two first moments diverging we find that resetting is always beneficial. Finally, we have also found that the optimal strategy to exit the domain depends on the reset rate. For low reset rates, walkers with exponential waiting times are found to be optimal, while for high reset rate, anomalous waiting times optimize the search process.

Phase Transition in a Non-Markovian Animal Exploration Model with Preferential Returns.

We study a non-Markovian and nonstationary model of animal mobility incorporating both exploration and memory in the form of preferential returns. Exact results for the probability of visiting a given number of sites are derived and a practical WKB approximation to treat the nonstationary problem is developed. A mean-field version of this model, first suggested by Song et al., [Modelling the scaling properties of human mobility, Nat. Phys. 6, 818 (2010)NPAHAX1745-247310.1038/nphys1760] was shown to well describe human movement data. We show that our generalized model adequately describes empirical movement data of Egyptian fruit bats (Rousettus aegyptiacus) when accounting for interindividual variation in the population. We also study the probability of visiting any site a given number of times and derive a mean-field equation. Our analysis yields a remarkable phase transition occurring at preferential returns which scale linearly with past visits. Following empirical evidence, we suggest that this phase transition reflects a trade-off between extensive and intensive foraging modes.

Optimal escape-and-feeding dynamics of random walkers: Rethinking the convenience of ballistic strategies.

Excited random walks represent a convenient model to study food intake in a media which is progressively depleted by the walker. Trajectories in the model alternate between (i) feeding and (ii) escape (when food is missed and so it must be found again) periods, each governed by different movement rules. Here, we explore the case where the escape dynamics is adaptive, so at short times an area-restricted search is carried out, and a switch to extensive or ballistic motion occurs later if necessary. We derive for this case explicit analytical expressions of the mean escape time and the asymptotic growth of the depleted region in one dimension. These, together with numerical results in two dimensions, provide surprising evidence that ballistic searches are detrimental in such scenarios, a result which could explain why ballistic movement is barely observed in animal searches at microscopic and millimetric scales, therefore providing significant implications for biological foraging.

Continuous time random walks under Markovian resetting.

We investigate the effects of Markovian resetting events on continuous time random walks where the waiting times and the jump lengths are random variables distributed according to power-law probability density functions. We prove the existence of a nonequilibrium stationary state and finite mean first arrival time. However, the existence of an optimum reset rate is conditioned to a specific relationship between the exponents of both power-law tails. We also investigate the search efficiency by finding the optimal random walk which minimizes the mean first arrival time in terms of the reset rate, the distance of the initial position to the target, and the characteristic transport exponents.

Occupancy patterns in superorganisms: a spin-glass approach to ant exploration.

Emergence of collective, as well as superorganism-like, behaviour in biological populations requires the existence of rules of communication, either direct or indirect, between organisms. Because reaching an understanding of such rules at the individual level can be often difficult, approaches carried out at higher, or effective, levels of description can represent a useful alternative. In the present work, we show how a spin-glass approach characteristic of statistical physics can be used as a tool to characterize the properties of the spatial occupancy patterns of a biological population. We exploit the presence of pairwise interactions in spin-glass models for detecting correlations between occupancies at different sites in the media. Such correlations, we claim, represent a proxy to the existence of planned and/or social strategies in the spatial organization of the population. Our spin-glass approach does not only identify those correlations but produces a statistical replica of the system (at the level of occupancy patterns) that can be subsequently used for testing alternative conditions/hypothesis. Here, this methodology is presented and illustrated for a particular case of study: we analyse occupancy patterns of Aphaenogaster senilis ants during foraging through a simplified environment consisting of a discrete (tree-like) artificial lattice. Our spin-glass approach consistently reproduces the experimental occupancy patterns across time, and besides, an intuitive biological interpretation of the parameters is attainable. Likewise, we prove that pairwise correlations are important for reproducing these dynamics by showing how a null model, where such correlations are neglected, would perform much worse; this provides a solid evidence to the existence of superorganism-like strategies in the colony.

General scaling in bidirectional flows of self-avoiding agents.

The analysis of the classical radial distribution function of a system provides a possible procedure for uncovering interaction rules between individuals out of collective movement patterns. A formal extension of this approach has revealed recently the existence of a universal scaling in the collective spatial patterns of pedestrians, characterized by an effective potential of interaction [Formula: see text] conveniently defined in the space of the times-to-collision [Formula: see text] between the individuals. Here we significantly extend and clarify this idea by exploring numerically the emergence of that scaling for different scenarios. In particular, we compare the results of bidirectional flows when completely different rules of self-avoidance between individuals are assumed (from physical-like repulsive potentials to standard heuristic rules commonly used to reproduce pedestrians dynamics). We prove that all the situations lead to a common scaling in the t-space both in the disordered phase ([Formula: see text]) and in the lane-formation regime ([Formula: see text]), independent of the nature of the interactions considered. Our results thus suggest that these scalings cannot be interpreted as a proxy for how interactions between pedestrians actually occur, but they rather represent a common feature for bidirectional flows of self-avoiding agents.

Transport properties of random walks under stochastic noninstantaneous resetting.

Random walks with stochastic resetting provides a treatable framework to study interesting features about central-place motion. In this work, we introduce noninstantaneous resetting as a two-state model being a combination of an exploring state where the walker moves randomly according to a propagator and a returning state where the walker performs a ballistic motion with constant velocity towards the origin. We study the emerging transport properties for two types of reset time probability density functions (PDFs): exponential and Pareto. In the first case, we find the stationary distribution and a general expression for the stationary mean-square displacement (MSD) in terms of the propagator. We find that the stationary MSD may increase, decrease or remain constant with the returning velocity. This depends on the moments of the propagator. Regarding the Pareto resetting PDF we also study the stationary distribution and the asymptotic scaling of the MSD for diffusive motion. In this case, we see that the resetting modifies the transport regime, making the overall transport subdiffusive and even reaching a stationary MSD, i.e., a stochastic localization. This phenomena is also observed in diffusion under instantaneous Pareto resetting. We check the main results with stochastic simulations of the process.

Recurrence time correlations in random walks with preferential relocation to visited places.

Random walks with memory usually involve rules where a preference for either revisiting or avoiding those sites visited in the past are introduced somehow. Such effects have a direct consequence on the transport properties as well as on the statistics of first-passage and subsequent recurrence times through a site. A preference for revisiting sites is thus expected to result in a positive correlation between consecutive recurrence times. Here we derive a continuous-time generalization of the random walk model with preferential relocation to visited sites proposed in Phys. Rev. Lett. 112, 240601 (2014)PRLTAO0031-900710.1103/PhysRevLett.112.240601 to explore this effect, together with the main transport properties induced by the long-range memory. Despite the long-range memory effects governing the process, our analytical treatment allows us to (i) observe the existence of an asymptotic logarithmic (ultraslow) growth for the mean square displacement, in accordance to the results found for the original discrete-time model, and (ii) confirm the existence of positive correlations between first-passage and subsequent recurrence times. This analysis is completed with a comprehensive numerical study which reveals, among other results, that these correlations between first-passage and recurrence times also exhibit clear signatures of this ultraslow relaxation dynamics.

Demographic stochasticity and extinction in populations with Allee effect.

We study simple stochastic scenarios, based on birth-and-death Markovian processes, that describe populations with the Allee effect, to account for the role of demographic stochasticity. In the mean-field deterministic limit we recover well-known deterministic evolution equations widely employed in population ecology. The mean time to extinction is in general obtained by the Wentzel-Kramers-Brillouin (WKB) approximation for populations with the strong and weak Allee effects. An exact solution for the mean time to extinction can be found via a recursive equation for special cases of the stochastic dynamics. We study the conditions for the validity of the WKB solution and analyze the boundary between the weak and strong Allee effect by comparing exact solutions with numerical simulations.

Transport properties and first-arrival statistics of random motion with stochastic reset times.

Stochastic resets have lately emerged as a mechanism able to generate finite equilibrium mean-square displacement (MSD) when they are applied to diffusive motion. Furthermore, walkers with an infinite mean first-arrival time (MFAT) to a given position x may reach it in a finite time when they reset their position. In this work we study these emerging phenomena from a unified perspective. On one hand, we study the existence of a finite equilibrium MSD when resets are applied to random motion with 〈x^{2}(t)〉_{m}∼t^{p} for 0

Stochastic foundations in nonlinear density-regulation growth.

In this work we construct individual-based models that give rise to the generalized logistic model at the mean-field deterministic level and that allow us to interpret the parameters of these models in terms of individual interactions. We also study the effect of internal fluctuations on the long-time dynamics for the different models that have been widely used in the literature, such as the theta-logistic and Savageau models. In particular, we determine the conditions for population extinction and calculate the mean time to extinction. If the population does not become extinct, we obtain analytical expressions for the population abundance distribution. Our theoretical results are based on WKB theory and the probability generating function formalism and are verified by numerical simulations.

Nonstationary dynamics of encounters: Mean valuable territory covered by a random searcher.

Inspired by recent experiments on the organism Caenorhabditis elegans we present a stochastic problem to capture the adaptive dynamics of search in living beings, which involves the exploration-exploitation dilemma between remaining in a previously preferred area and relocating to new places. We assess the question of search efficiency by introducing a new magnitude, the mean valuable territory covered by a Browinan searcher, for the case where each site in the domain becomes valuable only after a random time controlled by a nonhomogeneous rate which expands from the origin outwards. We explore analytically this magnitude for domains of dimensions 1, 2, and 3 and discuss the theoretical and applied (biological) interest of our approach. As the main results here, we (i) report the existence of some universal scaling properties for the mean valuable territory covered as a function of time and (ii) reveal the emergence of an optimal diffusivity which appears only for domains in two and higher dimensions.

Variability in individual activity bursts improves ant foraging success.

Using experimental and computational methods, we study the role of behavioural variability in activity bursts (or temporal activity patterns) for individual and collective regulation of foraging in A. senilis ants. First, foraging experiments were carried out under special conditions (low densities of ants and food and absence of external cues or stimuli) where individual-based strategies are most prevalent. By using marked individuals and recording all foraging trajectories, we were then able to precisely quantify behavioural variability among individuals. Our main conclusions are that (i) variability of ant trajectories (turning angles, speed, etc.) is low compared with variability of temporal activity profiles, and (ii) this variability seems to be driven by plasticity of individual behaviour through time, rather than the presence of fixed behavioural stereotypes or specialists within the group. The statistical measures obtained from these experimental foraging patterns are then used to build a general agent-based model (ABM) which includes the most relevant properties of ant foraging under natural conditions, including recruitment through pheromone communication. Using the ABM, we are able to provide computational evidence that the characteristics of individual variability observed in our experiments can provide a functional advantage (in terms of foraging success) to the group; thus, we propose the biological basis underpinning our observations. Altogether, our study reveals the potential utility of experiments under simplified (laboratory) conditions for understanding information-gathering in biological systems.

Foraging success under uncertainty: search tradeoffs and optimal space use.

Understanding the structural complexity and the main drivers of animal search behaviour is pivotal to foraging ecology. Yet, the role of uncertainty as a generative mechanism of movement patterns is poorly understood. Novel insights from search theory suggest that organisms should collect and assess new information from the environment by producing complex exploratory strategies. Based on an extension of the first passage time theory, and using simple equations and simulations, we unveil the elementary heuristics behind search behaviour. In particular, we show that normal diffusion is not enough for determining optimal exploratory behaviour but anomalous diffusion is required. Searching organisms go through two critical sequential phases (approach and detection) and experience fundamental search tradeoffs that may limit their encounter rates. Using experimental data, we show that biological search includes elements not fully considered in contemporary physical search theory. In particular, the need to consider search movement as a non-stationary process that brings the organism from one informational state to another. For example, the transition from remaining in an area to departing from it may occur through an exploratory state where cognitive search is challenged. Therefore, a more comprehensive view of foraging ecology requires including current perspectives about movement under uncertainty.

Characterization of stationary states in random walks with stochastic resetting.

It is known that introducing a stochastic resetting in a random-walk process can lead to the emergence of a stationary state. Here we study this point from a general perspective through the derivation and analysis of mesoscopic (continuous-time random walk) equations for both jump and velocity models with stochastic resetting. In the case of jump models it is shown that stationary states emerge for any shape of the waiting-time and jump length distributions. The existence of such state entails the saturation of the mean square displacement to an universal value that depends on the second moment of the jump distribution and the resetting probability. The transient dynamics towards the stationary state depends on how the waiting time probability density function decays with time. If the moments of the jump distribution are finite then the tail of the stationary distributions is universally exponential, but for Lévy flights these tails decay as a power law whose exponent coincides with that from the jump distribution. For velocity models we observe that the stationary state emerges only if the distribution of flight durations has finite moments of lower order; otherwise, as occurs for Lévy walks, the stationary state does not exist, and the mean square displacement grows ballistically or superdiffusively, depending on the specific shape of the distribution of movement durations.