arXiv subjects

Irene Giardina

Publications and source records attributed to Irene Giardina.

At least 19 recordsLinked to original sources

arXiv2026

Inheritance Entropy: A Model-Independent Method to Probe the Hereditary Structure of Cell Lineage Trees

Human bone marrow stromal cells (BMSC) include skeletal stem cells with ground-breaking therapeutic potential. However, BMSC colonies have very heterogeneous in vivo behaviour, due to their different potency; this unpredictability is the greatest hurdle to the development of skeletal regeneration therapies. Colony-level heterogeneity urges a fundamental question: how is it possible that one colony as a collective unit behaves differently from another one? If cell-to-cell variability were just an uncorrelated random process, a million cells in a transplant-bound colony would be enough to yield statistical homogeneity, hence washing out any colony-level traits. A possible answer is that the differences between two originating cells are transmitted to their progenies and collectively persist through an hereditary mechanism. But non-genetic inheritance remains an elusive notion, both at the experimental and at the theoretical level. Here, we prove that heterogeneity in the lineage topology of BMSC clonal colonies is determined by heritable traits that regulate cell-cycle exit. The cornerstone of this result is the definition of a novel entropy of the colony, which measures the hereditary ramifications in the distribution of inactive cells across different branches of the proliferation tree. We measure the entropy in 32 clonal colonies, obtained from single-cell lineage tracing experiments, and show that in the greatest majority of clones this entropy is decisively smaller than that of the corresponding non-hereditary lineage. This result indicates that hereditary epigenetic factors play a major role in determining cycle exit of bone marrow stromal cells.

physics.bio-ph↗

arXiv2025

Lineage topology, replication kinetics and cell cycle synchronization reveal regulated growth dynamics in human bone marrow stromal cell colonies

Bone marrow stromal cells (BMSC) -- which include skeletal stem cells -- are a promising tool in regenerative medicine. However, their heterogeneous and unpredictable in vivo behaviour remains a critical barrier preventing the development of standardized therapeutic approaches for skeletal tissue regeneration. Several studies have attempted to identify in vitro features that could correlate with the in vivo differentiation properties, yet the mechanisms ruling BMSC heterogeneity remain poorly understood. Here, using time-lapse imaging, we lineage-trace 32 single-cell-derived BMSC colonies through seven generations. We observe significant inter-colony and intra-colony heterogeneity in lineage topology (determined by the number of senescent or apoptotic cells) and in replicative kinetics (measured from proliferating cells only). Interestingly, topology and kinetics result strongly correlated, suggesting the existence of regulatory factors linking the non-dividing/apoptotic subpopulations with proliferating cells. Furthermore, BMSCs display highly synchronized cell cycles during early generations, indicating stage-specific regulatory mechanisms through which cells influence each other. By employing a non-interacting population growth model, we demonstrate that the observed synchronisation cannot be explained by an uncorrelated branching process; instead, cell-to-cell correlation of division times must exist. Our findings reveal fundamental mechanisms governing BMSC heterogeneity and growth dynamics that may inform strategies to control their regenerative potential.

q-bio.CB↗

arXiv2025

Spin-Waves without Spin-Waves: A Case for Soliton Propagation in Starling Flocks

Collective turns in starling flocks propagate linearly with negligible attenuation, indicating the existence of an underdamped sector in the dispersion relation. Beside granting linear propagation of the phase perturbations, the real part of the frequency should also yield a spin-wave form of the unperturbed correlation function. However, new high-resolution experiments on real flocks show that underdamped traveling waves coexist with an overdamped Lorentzian correlation. Theory and experiments are reconciled once we add to the dynamics a Fermi-Pasta-Ulam-Tsingou term.

cond-mat.stat-mech↗

arXiv2024

The 2024 Motile Active Matter Roadmap

Activity and autonomous motion are fundamental aspects of many living and engineering systems. Here, the scale of biological agents covers a wide range, from nanomotors, cytoskeleton, and cells, to insects, fish, birds, and people. Inspired by biological active systems, various types of autonomous synthetic nano- and micromachines have been designed, which provide the basis for multifunctional, highly responsive, intelligent active materials. A major challenge for understanding and designing active matter is their inherent non-equilibrium nature due to persistent energy consumption, which invalidates equilibrium concepts such as free energy, detailed balance, and time-reversal symmetry. Furthermore, interactions in ensembles of active agents are often non-additive and non-reciprocal. An important aspect of biological agents is their ability to sense the environment, process this information, and adjust their motion accordingly. It is an important goal for the engineering of micro-robotic systems to achieve similar functionality. With many fundamental properties of motile active matter now reasonably well understood and under control, the ground is prepared for the study of physical aspects and mechanisms of motion in complex environments, of the behavior of systems with new physical features like chirality, of the development of novel micromachines and microbots, of the emergent collective behavior and swarming of intelligent self-propelled particles, and of particular features of microbial systems. The vast complexity of phenomena and mechanisms involved in the self-organization and dynamics of motile active matter poses major challenges, which can only be addressed by a truly interdisciplinary effort involving scientists from biology, chemistry, ecology, engineering, mathematics, and physics.

cond-mat.soft↗

arXiv2024

Discrete Laplacian thermostat for flocks and swarms: the fully conserved Inertial Spin Model

Experiments on bird flocks and midge swarms reveal that these natural systems are well described by an active theory in which conservation laws play a crucial role. By building a symplectic structure that couples the particles' velocities to the generator of their internal rotations (spin), the Inertial Spin Model (ISM) reinstates a second-order temporal dynamics that captures many phenomenological traits of flocks and swarms. The reversible structure of the ISM predicts that the total spin is a constant of motion, the central conservation law responsible for all the novel dynamical features of the model. However, fluctuations and dissipation introduced in the original model to make it relax, violate the spin conservation law, so that the ISM aligns with the biophysical phenomenology only within finite-size regimes, beyond which the overdamped dynamics characteristic of the Vicsek model takes over. Here, we introduce a novel version of the ISM, in which the irreversible terms needed to relax the dynamics strictly respect the conservation of the spin. We perform a numerical investigation of the fully conservative model, exploring both the fixed-network case, which belongs to the equilibrium class of Model G, and the active case, characterized by self-propulsion of the agents and an out-of-equilibrium reshuffling of the underlying interaction network. Our simulations not only capture the correct spin wave phenomenology of the ordered phase, but they also yield dynamical critical exponents in the near-ordering phase that agree very well with the theoretical predictions.

cond-mat.stat-mech↗

arXiv2024

Out of equilibrium response and fluctuation-dissipation violations across scales in flocking systems

Flocking systems are known to be strongly out of equilibrium. Energy input occurs at the individual level to ensure self-propulsion, and the individual motility in turn contributes to ordering, enhancing information propagation and strengthening collective motion. However, even beyond ordering, a crucial feature of natural aggregations is response. How, then, do off-equilibrium features affect the response of the system? In this work, we consider a minimal model of flocking and investigate response behavior under directional perturbations. We show that equilibrium dynamical fluctuation-dissipation relations between response and correlations are violated, both at the local and at the global level. The amount of violation peaks at the ordering transition, exactly as for the entropy production rate. Entropy is always produced locally and connected to the local fluctuation-dissipation violation via Harada-Sasa relationships. However, cooperative mechanisms close to the transition spread off-equilibrium effects to the whole system, producing an out of equilibrium response on the global scale. Our findings elucidate the role of activity and interactions in the cost repartition of collective behavior and explain what observed in experiments on natural living groups.

cond-mat.stat-mech↗

arXiv2024

From noise on the sites to noise on the links: discretizing the conserved Kardar-Parisi-Zhang equation in real space

Numerical analysis of conserved field dynamics has been generally performed with pseudo spectral methods. Finite differences integration, the common procedure for non-conserved field dynamics, indeed struggles to implement a conservative noise in the discrete spatial domain. In this work, we present a novel method to generate a conservative noise in the finite differences framework, which works for any discrete topology and boundary conditions. We apply it to numerically solve the conserved Kardar-Parisi-Zhang (cKPZ) equation, widely used to describe surface roughening when the number of particles is conserved. Our numerical simulations recover the correct scaling exponents $α$, $β$, and $z$ in $d=1$ and in $d=2$. To illustrate the potentiality of the method, we further consider the cKPZ equation on different kinds of non-standard lattices and on the random Euclidean graph. This is the first numerical study of conserved field dynamics on an irregular topology, paving the way to a broad spectrum of possible applications.

cond-mat.stat-mech↗

arXiv2023

Discrete Laplacian Thermostat for Spin Systems with Conserved Dynamics

A well-established numerical technique to study the dynamics of spin systems in which symmetries and conservation laws play an important role is to microcanonically integrate their reversible equations of motion, obtaining thermalization through initial conditions drawn with the canonical distribution. In order to achieve a more realistic relaxation of the magnetic energy, numerically expensive methods that explicitly couple the spins to the underlying lattice are normally employed. Here we introduce a stochastic conservative thermostat that relaxes the magnetic energy while preserving the constant of motions, thus turning microcanonical spin dynamics into a conservative canonical dynamics, without actually simulating the lattice. We test the thermostat on the Heisenberg antiferromagnet in d=3 and show that the method reproduces the exact values of the static and dynamic critical exponents, while in the low-temperature phase it yields the correct spin wave phenomenology. Finally, we demonstrate that the relaxation coefficient of the new thermostat is quantitatively connected to the microscopic parameters of the spin-lattice coupling.

cond-mat.stat-mech↗

arXiv2023

Collective response to local perturbations: how to evade threats without losing coherence

Living groups move in complex environments and are constantly subject to external stimuli, predatory attacks and disturbances. An efficient response to such perturbations is vital to maintain the group's coherence and cohesion. Perturbations are often local, i.e. they are initially perceived only by few individuals in the group, but can elicit a global response. This is the case of starling flocks, that can turn very quickly to evade predators. In this paper, we investigate the conditions under which a global change of direction can occur upon local perturbations. Using minimal models of self-propelled particles, we show that a collective directional response occurs on timescales that grow with the system size and it is, therefore, a finite-size effect. The larger the group is, the longer it will take to turn. We also show that global coherent turns can only take place if i) the mechanism for information propagation is efficient enough to transmit the local reaction undamped through the whole group; and if ii) motility is not too strong, to avoid that the perturbed individual leaves the group before the turn is complete. No compliance with such conditions results in the group's fragmentation or in a non-efficient response.

cond-mat.stat-mech↗

arXiv2022

Natural Swarms in $\bf 3.99$ Dimensions

The dynamical critical exponent $z$ of natural swarms of insects is calculated using the renormalization group to order $ε= 4-d$. A novel fixed point emerges, where both activity and inertia are relevant. In three dimensions the critical exponent at the new fixed point is $z = 1.35$, in agreement with both experiments ($1.37 \pm 0.11$) and numerical simulations ($1.35 \pm 0.04$).

cond-mat.stat-mech↗

arXiv2022

Signatures of irreversibility in microscopic models of flocking

Flocking in $d=2$ is a genuine non-equilibrium phenomenon for which irreversibility is an essential ingredient. We study a class of minimal flocking models whose only source of irreversibility is self-propulsion and use the entropy production rate (EPR) to quantify the departure from equilibrium across their phase diagrams. The EPR is maximal in the vicinity of the order-disorder transition, where reshuffling of the interaction network is fast. We show that signatures of irreversibility come in the form of asymmetries in the steady state distribution of the flock's microstates. They occur as consequences of the time reversal symmetry breaking in the considered self-propelled systems, independently of the interaction details. In the case of metric pairwise forces, they reduce to local asymmetries in the distribution of pairs of particles. This study suggests a possible use of pair asymmetries both to quantify the departure from equilibrium and to learn relevant information about aligning interaction potentials from data.

cond-mat.stat-mech↗

arXiv2022

Marginal speed confinement resolves the conflict between correlation and control in natural flocks of birds

Speed fluctuations of individual birds in natural flocks are moderate, due to the aerodynamic and biomechanical constraints of flight. Yet the spatial correlations of such fluctuations are scale-free, namely they have a range as wide as the entire group, a property linked to the capacity of the system to collectively respond to external perturbations. Scale-free correlations and moderate fluctuations set conflicting constraints on the mechanism controlling the speed of each agent, as the factors boosting correlation amplify fluctuations, and vice versa. Here, using a statistical field theory approach, we suggest that a marginal speed confinement that ignores small deviations from the natural reference value while ferociously suppressing larger speed fluctuations, is able to reconcile scale-free correlations with biologically acceptable group's speed. We validate our theoretical predictions by comparing them with field experimental data on starling flocks with group sizes spanning an unprecedented interval of over two orders of magnitude.

cond-mat.stat-mech↗

arXiv2021

A Renormalization Group Approach to Connect Discrete- and Continuous-Time Descriptions of Gaussian Processes

Discretization of continuous stochastic processes is needed to numerically simulate them or to infer models from experimental time series. However, depending on the nature of the process, the same discretization scheme, if not accurate enough, may perform very differently for the two tasks. Exact discretizations, which work equally well at any scale, are characterized by the property of invariance under coarse-graining. Motivated by this observation, we build an explicit Renormalization Group approach for Gaussian time series generated by auto-regressive models. We show that the RG fixed points correspond to discretizations of linear SDEs, and only come in the form of first order Markov processes or non-Markovian ones. This fact provides an alternative explanation of why standard delay-vector embedding procedures fail in reconstructing partially observed noise-driven systems. We also suggest a possible effective Markovian discretization for the inference of partially observed underdamped equilibrium processes based on the exploitation of the Einstein relation.

stat.ML↗

arXiv2021

Dynamical renormalization group for mode-coupling field theories with solenoidal constraint

The recent inflow of empirical data about the collective behaviour of strongly correlated biological systems has brought field theory and the renormalization group into the biophysical arena. Experiments on bird flocks and insect swarms show that social forces act on the particles' velocity through the generator of its rotations, namely the spin, indicating that mode-coupling field theories are necessary to reproduce the correct dynamical behaviour. Unfortunately, a theory for three coupled fields - density, velocity and spin - has a prohibitive degree of intricacy. A simplifying path consists in getting rid of density fluctuations by studying incompressible systems. This requires imposing a solenoidal constraint on the primary field, an unsolved problem even for equilibrium mode-coupling theories. Here, we perform an equilibrium dynamic renormalization group analysis of a mode-coupling field theory subject to a solenoidal constraint; using the classification of Halperin and Hohenberg, we can dub this case as a solenoidal Model G. We demonstrate that the constraint produces a new vertex that mixes static and dynamical coupling constants, and that this vertex is essential to grant the closure of the renormalization group structure and the consistency of dynamics with statics. Interestingly, although the solenoidal constraint leads to a modification of the static universality class, we find that it does not change the dynamical universality class, a result that seems to represent an exception to the general rule that dynamical universality classes are narrower than static ones. Our results constitute a solid stepping stone in the admittedly large chasm towards developing an off-equilibrium mode-coupling theory of biological groups.

cond-mat.stat-mech↗

arXiv2020

Equilibrium to off-equilibrium crossover in homogeneous active matter

We study the crossover between equilibrium and off-equilibrium dynamical universality classes in the Vicsek model near its ordering transition. Starting from the incompressible hydrodynamic theory of Chen et al \cite{chen2015critical}, we show that increasing the activity leads to a renormalization group (RG) crossover between the equilibrium ferromagnetic fixed point, with dynamical critical exponent $z = 2$, and the off-equilibrium active fixed point, with $z = 1.7$ (in $d=3$). We run simulations of the classic Vicsek model in the near-ordering regime and find that critical slowing down indeed changes with activity, displaying two exponents that are in remarkable agreement with the RG prediction. The equilibrium-to-off-equilibrium crossover is ruled by a characteristic length scale beyond which active dynamics takes over. Such length scale is smaller the larger the activity, suggesting the existence of a general trade-off between activity and system's size in determining the dynamical universality class of active matter.

cond-mat.stat-mech↗

arXiv2020

Building general Langevin models from discrete data sets

Many living and complex systems exhibit second order emergent dynamics. Limited experimental access to the configurational degrees of freedom results in data that appears to be generated by a non-Markovian process. This poses a challenge in the quantitative reconstruction of the model from experimental data, even in the simple case of equilibrium Langevin dynamics of Hamiltonian systems. We develop a novel Bayesian inference approach to learn the parameters of such stochastic effective models from discrete finite length trajectories. We first discuss the failure of naive inference approaches based on the estimation of derivatives through finite differences, regardless of the time resolution and the length of the sampled trajectories. We then derive, adopting higher order discretization schemes, maximum likelihood estimators for the model parameters that provide excellent results even with moderately long trajectories. We apply our method to second order models of collective motion and show that our results also hold in the presence of interactions.

q-bio.QM↗

arXiv2019

Comment on "Velocity and Speed Correlations in Hamiltonian Flocks" by M. Casiulis et al. [arXiv:1911.06042]

In arXiv:1911.06042v1 M. Casiulis et al. study a Hamiltonian model in which rigid rotations of moving clusters give rise to scale-free correlations of velocity and speed. M. Casiulis et al. compare correlations in their model to those observed in real flocks of birds and claim that rigid-body rotations provide an explanation that stands in contrast with, and it is simpler than, previously proposed explanations of correlations in bird flocks, namely Goldstone modes in the velocity orientations and marginal (or near-critical) modes in the speed. Here, we show that the rigid rotation scenario is completely inconsistent with a large body of well-established experimental evidence on real flocks of birds, and it therefore does not provide an appropriate explanation for the observed phenomenology.

physics.bio-ph↗

arXiv2019

Low-temperature marginal ferromagnetism explains anomalous scale-free correlations in natural flocks

We introduce a new ferromagnetic model capable of reproducing one of the most intriguing properties of collective behaviour in starling flocks, namely the fact that strong collective order of the system coexists with scale-free correlations of the modulus of the microscopic degrees of freedom, that is the birds' speeds. The key idea of the new theory is that the single-particle potential needed to bound the modulus of the microscopic degrees of freedom around a finite value, is marginal, that is has zero curvature. We study the model by using mean-field approximation and Monte Carlo simulations in three dimensions, complemented by finite-size scaling analysis. While at the standard critical temperature, $T_c$, the properties of the marginal model are exactly the same as a normal ferromagnet with continuous symmetry-breaking, our results show that a novel zero-temperature critical point emerges, so that in its deeply ordered phase the marginal model develops divergent susceptibility and correlation length of the modulus of the microscopic degrees of freedom, in complete analogy with experimental data on natural flocks of starlings.

cond-mat.stat-mech↗