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Bernat Corominas-Murtra

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Published work

18 published item(s)

preprint2026arXiv

A hyperbolic cell cycle law for early embryonic developmental timing

Across metazoans, early embryos exhibit a strikingly conserved slowing down of their cell duplication speed, despite widely varying developmental paces and underlying molecular mechanisms. Here we show that this common behavior arises because early development unfolds along a biochemical rather than a chronological timescale, resulting from the coupling of finite maternal resource consumption to the Michaelis-Menten-like kinetics governing the rates of the biochemical reactions involved in cell duplication. This leads to a hyperbolic growth of the Cell Cycle Length (CCL), approaching a mathematical singularity, which would correspond to developmental arrest. Data from a wide range of organisms -- cnidarians, nematodes, arthropods, molluscs, echinoderms, tunicates, amphibians, and fish -- collapse on a single curve, quantitatively capturing not only a universal CCL dynamical behaviour, but also key hallmarks of early metazoan development, including cell-number temporal evolution, the dependency of CCL on cell size, and, remarkably, gastrulation timing at the predicted singularity. Crucially, experimental modulation of resource availability and consumption rates validate the model and further demonstrate that a source of heterochrony in early development is an altered biochemical timescale of resource depletion. Overall, this work reveals resource consumption rates as a fundamental mechanism driving developmental timing in early embryogenesis across species.

preprint2022arXiv

The typical set and entropy in stochastic systems with arbitrary phase space growth

The existence of the {\em typical set} is key for data compression strategies and for the emergence of robust statistical observables in macroscopic physical systems. Standard approaches derive its existence from a restricted set of dynamical constraints. However, given the enormous consequences for the understanding of the system's dynamics, and its role underlying the presence of stable, almost deterministic statistical patterns, a question arises whether typical sets exist in much more general scenarios. We demonstrate here that the typical set can be defined and characterized from general forms of entropy for a much wider class of stochastic processes than it was previously thought. This includes processes showing arbitrary path dependence, long range correlations or dynamic sampling spaces; suggesting that typicality is a generic property of stochastic processes, regardless of their complexity. Our results impact directly in the understanding of the stability of complex systems, open the door to new data compression strategies and points to the existence of statistical mechanics-like approaches to systems arbitrarily away from equilibrium with dynamic phase spaces. We argue that the potential emergence of robust properties in complex stochastic systems provided by the existence of typical sets has special relevance to biological systems.

preprint2020arXiv

Stem cell lineage survival as a noisy competition for niche access

Understanding to what extent stem cell potential is a cell-intrinsic property, or an emergent behavior coming from global tissue dynamics and geometry, is a key outstanding question of systems and stem cell biology. Here, we propose a theory of stem cell dynamics as a stochastic competition for access to a spatially-localized niche, giving rise to a stochastic conveyor-belt model. Cell divisions produce a steady cellular stream which advects cells away from the niche, while random rearrangements enable cells away from the niche to be favourably repositioned. Importantly, even when assuming that all cells in a tissue are molecularly equivalent, we predict a common ("universal") functional dependence of the long-term clonal survival probability on distance from the niche, as well as the emergence of a well-defined number of functional stem cells, dependent only on the rate of random movements vs. mitosis-driven advection. We test the predictions of this theory on datasets on pubertal mammary gland tips, embryonic kidney tips as well homeostatic intestinal crypt. Importantly, we find good agreement for the predicted functional dependency of the competition as a function of position, and thus functional stem cell number in each organ. This argues for a key role of positional fluctuations in dictating stem cell number and dynamics, and we discuss the applicability of this theory to other settings.

preprint2016arXiv

Analytical computation of frequency distributions of path-dependent processes by means of a non-multinomial maximum entropy approach

Path-dependent stochastic processes are often non-ergodic and observables can no longer be computed within the ensemble picture. The resulting mathematical difficulties pose severe limits to the analytical understanding of path-dependent processes. Their statistics is typically non-multinomial in the sense that the multiplicities of the occurrence of states is not a multinomial factor. The maximum entropy principle is tightly related to multinomial processes, non-interacting systems, and to the ensemble picture; It loses its meaning for path-dependent processes. Here we show that an equivalent to the ensemble picture exists for path-dependent processes, such that the non-multinomial statistics of the underlying dynamical process, by construction, is captured correctly in a functional that plays the role of a relative entropy. We demonstrate this for self-reinforcing Pólya urn processes, which explicitly generalise multinomial statistics. We demonstrate the adequacy of this constructive approach towards non-multinomial pendants of entropy by computing frequency and rank distributions of Pólya urn processes. We show how microscopic update rules of a path-dependent process allow us to explicitly construct a non-multinomial entropy functional, that, when maximized, predicts the time-dependent distribution function.

preprint2015arXiv

Non-Equilibrium Thermodynamics of Self-Replicating Protocells

We provide a non-equilibrium thermodynamic description of the life-cycle of a droplet based, chemically feasible, system of protocells. By coupling the protocells metabolic kinetics with its thermodynamics, we demonstrate how the system can be driven out of equilibrium to ensure protocell growth and replication. This coupling allows us to derive the equations of evolution and to rigorously demonstrate how growth and replication life-cycle can be understood as a non-equilibrium thermodynamic cycle. The process does not appeal to genetic information or inheritance, and is based only on non-equilibrium physics considerations. Our non-equilibrium thermodynamic description of simple, yet realistic, processes of protocell growth and replication, represents an advance in our physical understanding of a central biological phenomenon both in connection to the origin of life and for modern biology.

preprint2015arXiv

Understanding scaling through history-dependent processes with collapsing sample space

History-dependent processes are ubiquitous in natural and social systems. Many such stochastic processes, especially those that are associated with complex systems, become more constrained as they unfold, meaning that their sample-space, or their set of possible outcomes, reduces as they age. We demonstrate that these sample-space reducing (SSR) processes necessarily lead to Zipf's law in the rank distributions of their outcomes. We show that by adding noise to SSR processes the corresponding rank distributions remain exact power-laws, $p(x)\sim x^{-λ}$, where the exponent directly corresponds to the mixing ratio of the SSR process and noise. This allows us to give a precise meaning to the scaling exponent in terms of the degree to how much a given process reduces its sample-space as it unfolds. Noisy SSR processes further allow us to explain a wide range of scaling exponents in frequency distributions ranging from $α= 2$ to $\infty$. We discuss several applications showing how SSR processes can be used to understand Zipf's law in word frequencies, and how they are related to diffusion processes in directed networks, or ageing processes such as in fragmentation processes. SSR processes provide a new alternative to understand the origin of scaling in complex systems without the recourse to multiplicative, preferential, or self-organised critical processes.

preprint2015arXiv

Understanding Zipf's law of word frequencies through sample-space collapse in sentence formation

The formation of sentences is a highly structured and history-dependent process. The probability of using a specific word in a sentence strongly depends on the 'history' of word-usage earlier in that sentence. We study a simple history-dependent model of text generation assuming that the sample-space of word usage reduces along sentence formation, on average. We first show that the model explains the approximate Zipf law found in word frequencies as a direct consequence of sample-space reduction. We then empirically quantify the amount of sample-space reduction in the sentences of ten famous English books, by analysis of corresponding word-transition tables that capture which words can follow any given word in a text. We find a highly nested structure in these transition tables and show that this `nestedness' is tightly related to the power law exponents of the observed word frequency distributions. With the proposed model it is possible to understand that the nestedness of a text can be the origin of the actual scaling exponent, and that deviations from the exact Zipf law can be understood by variations of the degree of nestedness on a book-by-book basis. On a theoretical level we are able to show that in case of weak nesting, Zipf's law breaks down in a fast transition. Unlike previous attempts to understand Zipf's law in language the sample-space reducing model is not based on assumptions of multiplicative, preferential, or self-organised critical mechanisms behind language formation, but simply used the empirically quantifiable parameter 'nestedness' to understand the statistics of word frequencies.

preprint2014arXiv

Detection of the elite structure in a virtual multiplex social system by means of a generalized $K$-core

Elites are subgroups of individuals within a society that have the ability and means to influence, lead, govern, and shape societies. Members of elites are often well connected individuals, which enables them to impose their influence to many and to quickly gather, process, and spread information. Here we argue that elites are not only composed of highly connected individuals, but also of intermediaries connecting hubs to form a cohesive and structured elite-subgroup at the core of a social network. For this purpose we present a generalization of the $K$-core algorithm that allows to identify a social core that is composed of well-connected hubs together with their `connectors'. We show the validity of the idea in the framework of a virtual world defined by a massive multiplayer online game, on which we have complete information of various social networks. Exploiting this multiplex structure, we find that the hubs of the generalized $K$-core identify those individuals that are high social performers in terms of a series of indicators that are available in the game. In addition, using a combined strategy which involves the generalized $K$-core and the recently introduced $M$-core, the elites of the different 'nations' present in the game are perfectly identified as modules of the generalized $K$-core. Interesting sudden shifts in the composition of the elite cores are observed at deep levels. We show that elite detection with the traditional $K$-core is not possible in a reliable way. The proposed method might be useful in a series of more general applications, such as community detection.

preprint2014arXiv

The weak core and the structure of elites in social multiplex networks

Recent approaches on elite identification highlighted the important role of {\em intermediaries}, by means of a new definition of the core of a multiplex network, the {\em generalised} $K$-core. This newly introduced core subgraph crucially incorporates those individuals who, in spite of not being very connected, maintain the cohesiveness and plasticity of the core. Interestingly, it has been shown that the performance on elite identification of the generalised $K$-core is sensibly better that the standard $K$-core. Here we go further: Over a multiplex social system, we isolate the community structure of the generalised $K$-core and we identify the weakly connected regions acting as bridges between core communities, ensuring the cohesiveness and connectivity of the core region. This gluing region is the {\em Weak core} of the multiplex system. We test the suitability of our method on data from the society of 420.000 players of the Massive Multiplayer Online Game {\em Pardus}. Results show that the generalised $K$-core displays a clearly identifiable community structure and that the weak core gluing the core communities shows very low connectivity and clustering. Nonetheless, despite its low connectivity, the weak core forms a unique, cohesive structure. In addition, we find that members populating the weak core have the best scores on social performance, when compared to the other elements of the generalised $K$-core. The weak core provides a new angle on understanding the social structure of elites, highlighting those subgroups of individuals whose role is to glue different communities in the core.

preprint2013arXiv

On the origin of ambiguity in efficient communication

This article studies the emergence of ambiguity in communication through the concept of logical irreversibility and within the framework of Shannon's information theory. This leads us to a precise and general expression of the intuition behind Zipf's vocabulary balance in terms of a symmetry equation between the complexities of the coding and the decoding processes that imposes an unavoidable amount of logical uncertainty in natural communication. Accordingly, the emergence of irreversible computations is required if the complexities of the coding and the decoding processes are balanced in a symmetric scenario, which means that the emergence of ambiguous codes is a necessary condition for natural communication to succeed.

preprint2013arXiv

On the Origins of Hierarchy in Complex Networks

Hierarchy seems to pervade complexity in both living and artificial systems. Despite its relevance, no general theory that captures all features of hierarchy and its origins has been proposed yet. Here we present a formal approach resulting from the convergence of theoretical morphology and network theory that allows constructing a 3D morphospace of hierarchies and hence comparing the hierarchical organization of ecological, cellular, technological and social networks. Embedded within large voids in the morphospace of all possible hierarchies, four major groups are identified. Two of them match the expected from random networks with similar connectivity, thus suggesting that non-adaptive factors are at work. Ecological and gene networks define the other two, indicating that their topological order is the result of functional constraints. These results are consistent with an exploration of the morphospace using in silico evolved networks.

preprint2011arXiv

Emergence of Zipf's Law in the Evolution of Communication

Zipf's law seems to be ubiquitous in human languages and appears to be a universal property of complex communicating systems. Following the early proposal made by Zipf concerning the presence of a tension between the efforts of speaker and hearer in a communication system, we introduce evolution by means of a variational approach to the problem based on Kullback's Minimum Discrimination of Information Principle. Therefore, using a formalism fully embedded in the framework of information theory, we demonstrate that Zipf's law is the only expected outcome of an evolving, communicative system under a rigorous definition of the communicative tension described by Zipf.

preprint2011arXiv

Measuring the Hierarchy of Feedforward Networks

In this paper we explore the concept of hierarchy as a quantifiable descriptor of ordered structures, departing from the definition of three conditions to be satisfied for a hierarchical structure: {\em order}, {\em predictability} and {\em pyramidal structure}. According to these principles we define a hierarchical index taking concepts from graph and information theory. This estimator allows to quantify the hierarchical character of any system susceptible to be abstracted in a feedforward causal graph, i.e., a directed acyclic graph defined in a single connected structure. Our hierarchical index is a balance between this predictability and pyramidal condition by the definition of two entropies: one attending the onward flow and other for the backward reversion. We show how this index allows to identify hierarchical, anti-hierarchical and non hierarchical structures. Our formalism reveals that departing from the defined conditions for a hierarchical structure, feedforward trees and the inverted tree graphs emerge as the only causal structures of maximal hierarchical and anti-hierarchical systems, respectively. Conversely, null values of the hierarchical index are attributed to a number of different configuration networks; from linear chains, due to their lack of pyramid structure, to full-connected feedforward graphs where the diversity of onward pathways is canceled by the uncertainty (lack of predictability) when going backwards. Some illustrative examples are provided for the distinction among these three types of hierarchical causal graphs.

preprint2010arXiv

Diversity, competition, extinction: the ecophysics of language change

As early indicated by Charles Darwin, languages behave and change very much like living species. They display high diversity, differentiate in space and time, emerge and disappear. A large body of literature has explored the role of information exchanges and communicative constraints in groups of agents under selective scenarios. These models have been very helpful in providing a rationale on how complex forms of communication emerge under evolutionary pressures. However, other patterns of large-scale organization can be described using mathematical methods ignoring communicative traits. These approaches consider shorter time scales and have been developed by exploiting both theoretical ecology and statistical physics methods. The models are reviewed here and include extinction, invasion, origination, spatial organization, coexistence and diversity as key concepts and are very simple in their defining rules. Such simplicity is used in order to catch the most fundamental laws of organization and those universal ingredients responsible for qualitative traits. The similarities between observed and predicted patterns indicate that an ecological theory of language is emerging, supporting (on a quantitative basis) its ecological nature, although key differences are also present. Here we critically review some recent advances lying and outline their implications and limitations as well as open problems for future research.

preprint2010arXiv

Exploring the randomness of Directed Acyclic Networks

The feed-forward relationship naturally observed in time-dependent processes and in a diverse number of real systems -such as some food-webs and electronic and neural wiring- can be described in terms of so-called directed acyclic graphs (DAGs). An important ingredient of the analysis of such networks is a proper comparison of their observed architecture against an ensemble of randomized graphs, thereby quantifying the {\em randomness} of the real systems with respect to suitable null models. This approximation is particularly relevant when the finite size and/or large connectivity of real systems make inadequate a comparison with the predictions obtained from the so-called {\em configuration model}. In this paper we analyze four methods of DAG randomization as defined by the desired combination of topological invariants (directed and undirected degree sequence and component distributions) aimed to be preserved. A highly ordered DAG, called \textit{snake}-graph and a Erd\:os-Rényi DAG were used to validate the performance of the algorithms. Finally, three real case studies, namely, the \textit{C. elegans} cell lineage network, a PhD student-advisor network and the Milgram's citation network were analyzed using each randomization method. Results show how the interpretation of degree-degree relations in DAGs respect to their randomized ensembles depend on the topological invariants imposed. In general, real DAGs provide disordered values, lower than the expected by chance when the directedness of the links is not preserved in the randomization process. Conversely, if the direction of the links is conserved throughout the randomization process, disorder indicators are close to the obtained from the null-model ensemble, although some deviations are observed.

preprint2010arXiv

Topological reversibility and causality in feed-forward networks

Systems whose organization displays causal asymmetry constraints, from evolutionary trees to river basins or transport networks, can be often described in terms of directed paths (causal flows) on a discrete state space. Such a set of paths defines a feed-forward, acyclic network. A key problem associated with these systems involves characterizing their intrinsic degree of path reversibility: given an end node in the graph, what is the uncertainty of recovering the process backwards until the origin? Here we propose a novel concept, \textit{topological reversibility}, which rigorously weigths such uncertainty in path dependency quantified as the minimum amount of information required to successfully revert a causal path. Within the proposed framework we also analytically characterize limit cases for both topologically reversible and maximally entropic structures. The relevance of these measures within the context of evolutionary dynamics is highlighted.

preprint2010arXiv

Universality in percolation of arbitrary Uncorrelated Nested Subgraphs

The study of percolation in so-called {\em nested subgraphs} implies a generalization of the concept of percolation since the results are not linked to specific graph process. Here the behavior of such graphs at criticallity is studied for the case where the nesting operation is performed in an uncorrelated way. Specifically, I provide an analyitic derivation for the percolation inequality showing that the cluster size distribution under a generalized process of uncorrelated nesting at criticality follows a power law with universal exponent $γ=3/2$. The relevance of the result comes from the wide variety of processes responsible for the emergence of the giant component that fall within the category of nesting operations, whose outcome is a family of nested subgraphs.

preprint2009arXiv

Switcher-random-walks: a cognitive-inspired mechanism for network exploration

Semantic memory is the subsystem of human memory that stores knowledge of concepts or meanings, as opposed to life specific experiences. The organization of concepts within semantic memory can be understood as a semantic network, where the concepts (nodes) are associated (linked) to others depending on perceptions, similarities, etc. Lexical access is the complementary part of this system and allows the retrieval of such organized knowledge. While conceptual information is stored under certain underlying organization (and thus gives rise to a specific topology), it is crucial to have an accurate access to any of the information units, e.g. the concepts, for efficiently retrieving semantic information for real-time needings. An example of an information retrieval process occurs in verbal fluency tasks, and it is known to involve two different mechanisms: -clustering-, or generating words within a subcategory, and, when a subcategory is exhausted, -switching- to a new subcategory. We extended this approach to random-walking on a network (clustering) in combination to jumping (switching) to any node with certain probability and derived its analytical expression based on Markov chains. Results show that this dual mechanism contributes to optimize the exploration of different network models in terms of the mean first passage time. Additionally, this cognitive inspired dual mechanism opens a new framework to better understand and evaluate exploration, propagation and transport phenomena in other complex systems where switching-like phenomena are feasible.