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Tuesday 29 September

Biological Physics · Cross-listed from physics.soc-ph

Statistical Physics of Fish Collective Motion

We review recent theoretical and empirical advances that recast fish schooling within a statistical-physics framework, emphasizing how heterogeneous and time-dependent interactions govern collective motion. Building on extensions of Vicsek-type models to complex and weighted social networks, we show that topology and link strength qualitatively alter flocking stability and critical thresholds, with empirical weights typically reducing global alignment. High-resolution trajectory inference reveals selective, nonreciprocal responses: individuals preferentially attend to faster neighbors, producing transient, speed-induced leadership rather than fixed hierarchies. At the mesoscopic scale, schools exhibit avalanche-like turning cascades with scale-free size and duration statistics, aftershock clustering, and an Omori-type temporal decay with a short memory. Together, these findings support a unified picture in which collective order and critical-like fluctuations coexist: alignment provides stability while near-critical variability preserves responsiveness. This synthesis highlights fish schools as a tractable experimental model for how living collectives organize, transmit and process information across scales.
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Biological Physics · Cross-listed from q-bio.NC

An adaptive fractional state links circuit mechanisms to cortical dynamics across the visual hierarchy

Cortical circuits must respond flexibly to new inputs while integrating information about the past, yet the way in which neural activity reconciles these competing demands remains unclear. Combining Neuropixels recordings from six mouse visual areas with mechanistic circuit modeling, we identify a dynamical regime in which heavy-tailed superdiffusive fluctuations coexist with long-range temporal dependence and oscillations. We formalize this regime as the adaptive fractional (AF) state, using an effective mean-field theory in which spatial and temporal fractional derivatives capture heavy-tailed excursions and long-range memory, respectively. We find that dynamical exponents characterizing the AF state vary systematically across the visual hierarchy: higher visual areas exhibit weaker superdiffusion and stronger temporal memory. In the circuit model, this hierarchical shift emerges from a progressive weakening of effective inhibition. These findings extend the classical hierarchy of timescales to a hierarchy of dynamical regimes and suggest that the AF state allows cortical areas to jointly express rapid responses and long-term temporal integration.
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Biological Physics · Cross-listed from physics.plasm-ph

Rectification of plasma-powered active matter into macroscopic work

Plasmas sustain strong energy and momentum flux, yet this activity is typically treated as a dissipative loss channel rather than a usable resource, and its conversion into macroscopic work remains challenging. Here we demonstrate a plasma-powered active engine by coupling self-propelled micromotors to ratchet rectification. Dielectric microspheres in the plasma sheath spontaneously develop asymmetric surface charging and undergo a Quincke rotational instability, forming fast micromotors that extract energy directly from the plasma. Their stochastic impacts are rectified by a sawtooth rotor into a directed angular-momentum flux that drives steady rotation, while an outer asymmetric gear organizes the active bath into coherent circulation to amplify torque transfer. Operating in an inertia-relevant regime, the engine achieves orders-of-magnitude enhancements in both power output and end-to-end efficiency compared with liquid-phase active engines, and remains functional down to the single-micromotor limit. More broadly, our results establish a general route to rectify nonequilibrium plasma activity for macroscopic work.
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All authors Ting-yu Yao, Shuo Wang, Shao-peng Li, Ming-hang Zhao, Bao-quan Ai, Ya-feng He

Biological Physics · Cross-listed from physics.plasm-ph

Cold atmospheric plasma decontaminates Arabidopsis thaliana seeds and remodels seedling fungal microbiota

Seed-associated microorganisms influence seed quality, seedling establishment, and plant health and also constitute a major source of seed-borne pathogens. Cold atmospheric plasma (CAP) has emerged as a promising alternative to chemical seed treatments because of its antimicrobial activity, although its effects on fungal communities associated with developing seedlings remain poorly understood. Here, Arabidopsis thaliana seeds from two ecotypes (Columbia and Landsberg erecta) were exposed to CAP for 5 or 15 min. Seed decontamination efficiency was assessed by culturing on malt extract agar and nephelometric analyses, while fungal communities associated with seedlings derived from treated and untreated seeds were characterized by ITS1 amplicon sequencing. CAP efficiently reduced fungal contamination without affecting seed germination. CAP altered the composition of fungal communities associated with developing seedlings, but the magnitude of these changes depended on seed batch and ecotype. Dominant taxa markedly declined after treatment, whereas several low-abundance taxa increased in relative abundance. These findings demonstrate that CAP is an effective pesticide-free technology for seed decontamination and can also reshape fungal communities associated with developing seedlings, highlighting broader ecological consequences of plasma-based seed treatments.
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All authors Lena Taras, Nicole Chaumont, Caroline Kunz, Thierry Dufour, Christophe Bailly

Biological Physics · Cross-listed from cond-mat.stat-mech

Thermodynamics of the space of trajectories governed by a combination of two additive boundary functionals

This paper extends the formalism of stochastic path-space thermodynamics by systematically expanding the space of thermodynamic variables with a spectrum of boundary and relative functionals of random processes. Generalizing the approaches introduced in preprints arXiv:2607.24078 and arXiv:2609.08477, we investigate the fluctuation statistics of a one-dimensional Ornstein-Uhlenbeck process with an asymmetric linear drift and a step penalty potential at the origin, which models the waiting phase of a molecular Brownian motor. Utilizing the Feynman-Kac formalism and the parabolic cylinder theory, a numerical scheme based on matching logarithmic derivatives via the secant method is constructed. A high-precision eigenvalues of the effective Hamiltonian is obtained, which define the cumulant generating function. We formulate a modified path-space fluctuation theorem of the Gallavotti-Cohen type for the conjugate drift current at a fixed integral occupation time of the system in the dissipative half-plane. The physical and thermodynamic significance of the new variables is elucidated. A combination of "residence time" and "integral current" functionals is applied to describe an ion channel sensitive to mechanical or electrical stimuli.
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Monday 28 September

Biological Physics · Cross-listed from cond-mat.soft

Quorum sensing with density-enhanced motility and size regulation

Motivated by biological systems and my previous studies of quorum sensing with density-enhanced motility, I study a model of density-enhanced size and motility, in which both particle size and motility increase when the local density (over a distance larger than particle diameter) exceeds the global density. The emergent structures along different size ratios are spots, holes, bands and labyrinths controlled by several distinct observations. First, the characteristic pattern length scale is controlled by the sensing radius and remains approximately independent of the size ratio. Second, the total area occupied by the particles decreases with increasing size ratio. Third, the active fraction barely depends on the size ratio. These results demonstrate how both particle size and motility can couple microscopic regulation to large-scale pattern formation. I conclude by discussing possible extensions of the model.
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Biological Physics

DNA geometry around highly transcribed bacterial genes

Recent high-resolution (500 bp) Hi-C experiments reported contact maps with unusual patterns around highly expressed bacterial DNA loci. These patterns, described as arched stripes and bundled domains, extend over nearly 100 kbp. We performed Brownian Dynamics simulations with a specially designed coarse-grained model to rationalize these findings. The main feature of the model is that it takes explicitly into account the waves of positive (respectively, negative) supercoiling generated by the translocating polymerase downstream (respectively, upstream) of its position. Contact maps computed from the simulations also display the arched stripe and bundled domain patterns for above-threshold values of the rate of twist injection. Computed DNA conformations indicate that these patterns reflect an extraordinarily entangled DNA geometry, which involves both plectonemic and toroidal supercoiling, with some DNA segments experiencing both of them simultaneously. Moreover, DNA segments located on each side of the polymerase systematically wind around tracts located on the other side, which is a purely out-of-equilibrium effect driven by the continuous injection of twist. The present work therefore reveals that repeated transcription of a DNA locus systematically brings into contact DNA segments separated by several tens of kbp, which may eventually contribute to allosteric modulation and long-range communication.
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Biological Physics · Cross-listed from cond-mat.soft

Cell division sets a universal flow geometry in cell layers

Collective flows in epithelial tissues contain a geometric backbone of vortical interfaces whose statistics exhibit hallmarks of critical percolation and conformal invariance. Yet how fundamental cellular processes govern the breakdown of such symmetry-rich flow geometry remains unclear. Here we show that cell division plays a central physical role in regulating this universal flow organization by controlling both the geometric and mechanical flexibility of the cell-cell network. Using pharmacological perturbations, we find that when proliferation is suppressed through two independent interventions, coherent flows persist but neighbor exchanges decline and conformally invariant geometry is lost. Blocking apoptosis does not affect universality, isolating division as the key control. A vertex model with tunable division quantitatively reproduces these effects and restores conformal invariance when division is allowed. We further trace this effect to changes in both the geometric and mechanical organization of the cell layer: Divisions act as intermittent topological renewals that loosen constraints, preserving the network's flexibility and capacity to rearrange across scales. Thus, beyond its canonical role in growth, cell division acts as a structural and mechanical regulator of collective self-organization. These findings establish a direct connection between fundamental biological processes and emergent physical symmetries.
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All authors Tianxiang Ma, Lasse Bonn, Valeriia Grudtsyna, Nigar Abbasova, Martin Cramer Pedersen, Nuno A. M. Araujo, Amin Doostmohammadi

Sunday 27 September

Biological Physics

Harmonic Theory of Behavior

Traditional models of collective behavior rely on prescribed interaction rules, leaving unresolved the question of how behavior arises from neural representations of space. Here, we develop a first-principles theory in which movement, decision-making, and collective organization emerge by coarse-graining fast neural dynamics on a topological representation of directional space. For a ring manifold encoding heading, this reduction yields a macroscopic theory of behavior: a decision landscape over directions that admits a harmonic decomposition. In this framework, behavior is governed by a spectral organization of directional information rather than ad hoc rules. We demonstrate that target-seeking, avoidance, choice, spatial decision-making, and diverse forms of collective motion arise as distinct organizations of the underlying harmonic landscape. The theory unifies neural representations, behavioral decisions, and collective dynamics in a single mathematical description.
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Biological Physics · Cross-listed from q-bio.NC

Pre-registered spectral and certified mixing analysis of the male Drosophila central nervous system connectome

We report a pre-registered test of five hypotheses about the synapse-flow random walk on the connectome of the male Drosophila central nervous system (166,700 neurons, 124 million synapses), in which a walker moves to a postsynaptic partner in proportion to synapse number. The hypotheses concern the spectral gap and its dependence on the neck connective, the localisation of the leading modes of an input-normalised signed map, the certified mixing depth measured by the Dobrushin coefficient, its robustness to per-synapse detection confidence, and the dependence of the signed spectrum on the neurotransmitter sign convention. After public registration and a code freeze, each was tested once. Two held: the two-neuron photoreceptor modes that lead the signed spectrum are an artefact of normalising by very small inputs, removed by an input floor at the median, and they do not depend on the sign convention. Three failed: at a threshold of five synapses per connection the spectral gap is an order of magnitude smaller than predicted ($|\lambda_2| = 0.9965$) and the chain is certifiably far from mixed after 128 steps, and at no confidence level tested was the mixing depth certifiably robust. Exploratory analyses trace the slow mode to eleven neurons of the cord, seven of them mesothoracic efferent neurons, that become nearly closed once weak connections are dropped, find a similar boundary set in the lamina of the FlyWire female brain, show that null models preserving cell-type wiring reproduce the slow modes, and show that exact chains built from the confidence data change the depth profile after 32 steps by at most 0.017 at the pre-registered confidence level, far less than the worst-case certificate allows. Random-walk summaries of connectomes are dominated by nearly closed sets at the edges of the reconstruction unless these are treated explicitly.
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Saturday 26 September

Biological Physics · Cross-listed from cond-mat.soft

Domain growth in lipid membranes and the budding instability

We investigate the growth of a circular domain in a phase-separating lipid membrane and its relation to the budding instability. We describe the in-plane dynamics with a conserved, time-dependent Ginzburg-Landau model under radial symmetry. Numerical solutions show a crossover of the domain size $D$ as a function of time $t$ from $D\sim t^{1/3}$ at early times to $D\sim t^{1/2}$ at later times, followed by saturation due to finite system size. The time-dependent line tension $\sigma(t)$, evaluated from the evolving concentration profile, rises rapidly before approaching the planar-interface value. This behavior distinguishes an early line-tension-dominated regime from a later domain-size-dominated regime. To estimate the budding instability, we combine $D(t)$ and $\sigma(t)$ with a quasistatic criterion for the loss of stability of a partially budded domain. The budding-instability time decreases with increasing membrane spontaneous curvature and increases near the critical point. Thus, the evolution of both domain size and line tension is essential for determining the time required for the budding instability.
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All authors Jean Wolff, Fabrice Thalmann, Carlos Marques, David Andelman, Haim Diamant, Shigeyuki Komura

Biological Physics · Cross-listed from cond-mat.soft

Active matter within flexible boundaries: a novel experimental approach with T. aceti nematodes

We experimentally explore the collective behavior of the nematode T. Aceti inside flexible boundaries showing the emergence of previously unreported states. These nematodes have been previously shown to be able to synchronize their body oscillations in a favorable condition of confined space. In this collective state, they are able to exert a strong pushing force which we exploit to study how their collective motion could deform a pliable boundary. Using a novel experimental technique, we were able to simulate a non-rigid border at a liquid-liquid-gas triple point. We discovered a state with periodic boundary deformations which are due to the synchronized oscillations of the nematodes. More interestingly, we found a novel state where the nematodes are able to form multiple protrusions in the border which oscillate, rotate, merge, and split. We propose a numerical model that is able to reproduce some of the observed parameters of this state as well as offer an analytical insight into protrusions dynamics. Our research opens a pathway for exploring the interaction between active matter and flexible interfaces with a new kinds of active matter agents forming dynamic non-chaotic border deformations.
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All authors Christina M. Ceballos, Matthew S. Mizuhara, Mykhailo Potomkin, Anton Peshkov

Biological Physics · Cross-listed from q-bio.QM

Structured Population Models for Follicular Development

Ovarian follicle development has been modeled either by compartmental models, where developmental stages are represented by discrete compartments, or by physiologically structured population models (PSPMs), formulated as transport PDEs describing follicles along a continuous maturation variable. Compartmental models reproduce full-cycle hormonal and follicular dynamics but cannot describe cellular features. PSPMs capture multiscale behavior, yet lack a direct link to compartmental descriptions, making hormonal feedback difficult to incorporate. Here, we introduce a class of ODE systems derived as finite-dimensional reductions of PSPMs via a perturbative moment closure. These systems provide backbone approximations of the structured dynamics, retaining key nonlinear mechanisms governing recruitment, selection for dominance, and atresia while projecting maturation onto macroscopic variables. Zero- and first-order moments, derived from phenomenological assumptions regarding follicular maturation, allow us to realistically model follicle sizes and blood estradiol concentrations over one follicular wave. This application is supported by bovine data and likely generalizable to other mammalian species.
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Thursday 24 September

Biological Physics · Cross-listed from q-bio.PE

Consistent determination of stability regimes in natural ecological communities from abundance time series

The stability of an ecological community is conventionally defined through species interactions, which quantify how species affect one another. Interaction strengths are notoriously difficult to measure in species-rich assemblages. Long-term monitoring of species-rich communities, however, provides species abundance time series, increasingly available across habitats and taxa but not yet connected to the stability properties of the communities they describe. Here we develop a statistical framework that infers stability regimes in large ecological communities directly from abundance time series. We study stochastic Generalized Lotka–Volterra dynamics with random interactions and environmental fluctuations using dynamical mean-field theory, reducing the multispecies system to an effective stochastic process for a representative species. The theory predicts three stability regimes (stable coexistence, intermittent dynamics close to extinction, and unbounded growth) separated by analytical boundaries, and shows that environmental stochasticity systematically destabilizes coexistence by promoting intermittent low-abundance dynamics. The resulting steady-state species abundance distribution is a Gamma law that ties the dynamical phases to the variability-based stability metrics used in empirical studies. Recasting the effective dynamics as a multivariate regression model, we infer interaction statistics, environmental variability and the characteristic timescale of the dynamics from community data, without reconstructing the interaction network. Applied to natural communities spanning a broad range of habitats and taxa, the method resolves contrasting stability regimes and yields quantitative estimates of species extinction risk.
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All authors Alberto Megías, Vicente J. Ontiveros, David Alonso, José A. Capitán

Biological Physics · Cross-listed from nlin.AO

Higher-Order Competition as a Minimal Mechanism for Spatial Pattern Diversity in Population Dynamics

We show that negative feedback alone generates the diversity of self-organized shapes usually attributed to scale-dependent activation-inhibition. For a broad class of kernels, pairwise competition models only generate hexagonal spot arrays. Higher-order terms eliminate this restriction and promote stripes and gaps. Combining individual-based simulations and nonlinear analysis, we derive the pattern-selection thresholds and construct the full state diagram, including an unusual spots-stripes-spots sequence. Spot patterns are thus not a reliable indicator of proximity to a tipping point.
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Wednesday 23 September

Biological Physics · Cross-listed from math.AP

The Naito–Okuda–Ouyang Biconcave Red Blood Cell Profile as a Finite-Energy Unbranched Weak Immersion with Point-Force Residues

The axisymmetric profile \(\sin\psi(\rho)=c_0\rho\log(\rho/\rho_B)\) introduced by Naito, Okuda, and Ouyang is an exact solution of the unconstrained Helfrich shape equation for every \(\rho>0\), but its mean curvature diverges logarithmically at the dimple center. A covariant local analysis shows that in Cartesian graph coordinates the immersion is nondegenerate and belongs to \(W^{2,p}\) for every finite \(p\); it is \(C^{1,\alpha}\) for every \(\alpha<1\), but neither \(C^{1,1}\) nor \(C^2\). The conformal factor has a finite nonzero limit, so the point has branch multiplicity one and is not a branch point in the sense used in weak-immersion compactness theory. The Helfrich energy is finite, whereas the complete Euler–Lagrange operator carries the distributional residue \(\E_{\CH}=4\pi c_0\delta_p\) under the convention specified below. Hence the profile is a classical solution on the punctured surface and a finite-energy weak immersion, but it is not a free weak critical point of the unforced Canham–Helfrich functional. Stationarity is restored after adding the corresponding point-force potential or under a pinned-point constraint. Area and volume multipliers cannot cancel the Dirac residue. The apparent sign discrepancy between the shape equation used in the original papers and the modern convention is explained by the opposite definition of mean curvature. These results delimit precisely what can and cannot be inferred from modern existence and regularity theorems for Canham–Helfrich minimizers.
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Biological Physics · Cross-listed from cond-mat.soft

Position-dependent friction in protein folding from a GLE derived with a non-stationary localized projection distribution

The Generalized Langevin Equation (GLE) is an integro-differential equation of motion for a general observable of a many-body system and is rigorously derived using the projection operator formalism. In the standard derivation, a stationary canonical phase space distribution is used for the projection. Here, we derive a novel class of GLEs using a non-stationary projection distribution that constrains the initial ensemble to a hypersurface in phase space on which the observable has a fixed value $A_0$. As a result, all GLE parameters, and in particular the memory friction kernel, depend on $A_0$ and can be extracted from short simulations that do not sample the entire phase space. Applying this non-stationary GLE to protein folding trajectories, we find for villin and an $\alpha$-helical poly-alanine segment that the total friction is higher in the folded state. These results demonstrate that observable-dependent friction effects are non-negligible and can be accounted for using non-stationary GLEs derived by constrained projection schemes.
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All authors Salma Salem, Lucas Tepper, Benjamin J. A. Héry, Henrik Kiefer, David D. Girardier, Roland R. Netz

Biological Physics

Deep Generative Markov State Models with Experimental Restraints

A central challenge in molecular modeling is reconciling simulations with experimental observables, as force-field inaccuracies can distort equilibrium populations and long-timescale kinetics. While time-dependent restraints can improve simulated kinetics, time-dependent structural observables remain limited. Thus, most experimental data are time-averaged, informing equilibrium ensembles but not directly dynamics. Although ensemble refinement can improve thermodynamic accuracy, incorporating time-averaged observables into kinetic models remains difficult. Here, we introduce BICePs-reweighted Reversible DeepMSMs, combining Bayesian Inference of Conformational Populations (BICePs), variational learning of Markov processes, and maximum entropy (MaxEnt)/maximum caliber (MaxCal) principles to infer consistent thermodynamics and minimally perturbed kinetics. At its core is a reversible DeepMSM prior, obtained by applying an orthogonal transformation post-hoc to a pre-trained dynamics model (e.g., VAMPnet). This preserves the eigenspectrum exactly while yielding a valid transition matrix that is nonnegative, row-stochastic, and reversible. The reweighted model refines stationary populations, transition dynamics, and state-conditioned configurational landing densities. We validate the approach on a quadruple-well toy system and alanine dipeptide using backbone dihedral angles and J-coupling constants. In both systems, perturbing the observables induces predictable changes in the equilibrium ensemble and corresponding inferred kinetics. The resulting relaxation timescales and dynamical modes closely agree with analytical and MaxCal reference models. Furthermore, a diffusion-based generative model trained on MaxEnt landing densities produces physically realistic alanine dipeptide trajectories that reproduce thermodynamics and kinetics while satisfying the imposed experimental restraints.
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Biological Physics

Spatially Resolved Nucleated Polymerization: A Free-Boundary Model of Protein Aggregation in Concentrated Solutions

Kinetic models of protein aggregation describe populations by size, not by spatial organization or morphology. We extend Lumry-Eyring nucleated polymerization to a model in which the monomer is a density field and each aggregate is a region bounded by a level set. Growth is a flux condition on the available sites of a surface. Condensation is a reaction between the bonding sites of two surfaces in contact, at a rate set by the bond rate and the contact geometry. The availability of those sites is a field on the interface, and its equilibrium value follows from Wertheim's perturbation theory. The collision efficiency and the Fuchs stability ratio are therefore computed, not fitted. In a well-mixed limit the model's spatial averages satisfy the rate equations term by term; the monomer fraction agrees to eight parts in ten thousand, a difference that arises from equating aggregate size with volume. The condensation kernel's exponent is $0.5806\pm0.0013$ against the $0.600\pm0.010$ fitted to a monoclonal antibody. The computed stability ratio reproduces thirteen of fourteen published conditions at twelve $k_BT$, but only with the bond rate at the top of its range. In a many-body box, aggregates merge at $2.2$ to $3.8$ times the two-body rate.
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Tuesday 22 September

Biological Physics · Cross-listed from cs.RO

Water Surface Swimming in a Centipede and its Robophysical Model

Elongate multi-legged robots use coordinated body waves and distributed legs to move through cluttered terrestrial environments. However, as housing actuators for independent leg control can require bulky body segments, their non-streamlined body and limb structure makes it difficult to achieve swimming capability comparable to their terrestrial locomotor performance. At the water surface, we found that the multi-legged robots we tested unexpectedly moved backward: their body waves traveled in the same direction as their displacement, i.e., swimming with a direct wave. We found similar behavior in the centipede Lithobius forficatus, which swims with a direct body wave and periodic leg movement. To study how distributed legs contribute to direct-wave swimming, we analyze animal kinematics and develop a multi-legged robophysical model that allows independent variation of leg morphology and stiffness, body-wave direction, and leg coordination. Robophysical experiments show that direct body waves produce consistent forward motion under the tested conditions and that swimming performance depends on body–leg coordination. Additionally, directionally compliant legs increase displacement from approximately 0.08 to 0.21 body lengths per cycle relative to rigid legs under matched anti-phase actuation. These findings clarify how distributed appendages contribute to surface swimming and establish gait and morphology design principles for extending multi-legged field robots from terrestrial locomotion into aquatic environments.
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All authors Zhaochen J. Xu, Delfin Aydin, Abdullah Mustafa, Margarita B. Levin, Jianfeng Lin, Tianyu Wang, Daniel I. Goldman

Biological Physics · Cross-listed from cond-mat.soft

An interdimensional funnel enhances topologically protected oscillations in high-dimensional stochastic systems

Biochemical systems such as protein complexes often occupy configuration spaces whose dimensionality grows with the number of molecular components. Although usually viewed as an obstacle to coherent dynamics, here we show that high dimensionality can enhance topologically protected oscillations. In the topological regime, a stochastic system with a $D$-dimensional configuration space develops a steady-state current confined to a one-dimensional edge cycle. The intervening boundary faces between the $D$-dimensional bulk and the one-dimensional cycle form an interdimensional funnel, sequentially pushing the system towards lower dimensional faces, down to the one-dimensional cycle. As a consequence, increasing $D$ enhances the funnel, leading to improved one-dimensional localization and oscillatory coherence. A quantized biorthogonal Zak phase identifies the transition to the topological regime, which additionally exhibits a highly structured non-Hermitian skin effect. In contrast with previous realizations of topological states in stochastic systems, the phenomena we uncover have no counterpart in quantum condensed matter or active matter systems.
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Biological Physics · Cross-listed from cond-mat.soft

Experimental Characterization of the Passive Mechanical Behavior of Rat Pelvic Floor Muscles: Effects of Test Conditions and Pregnancy

Purpose: Pelvic floor muscles undergo large deformations during vaginal delivery, yet the passive tensile behavior of pregnant and non-pregnant pelvic floor muscle and the influence of testing protocols remain incompletely characterized. This study evaluated how pregnancy, loading rate, preconditioning, and storage affect passive tensile mechanics of rat pelvic floor muscle. Methods: Levator ani samples from pregnant and non-pregnant rats were tested in uniaxial tension to failure across multiple preconditioning amplitudes, two loading rates, and fresh or frozen storage conditions. Peak and transition metrics, strain energy density, toe-region stiffness, and high-strain stiffness were extracted using automated region-detection methods. Rat properties were also compared with digitized human cadaver levator ani data. Results: Pregnant rat samples reached greater peak strains than controls, although the difference was not significant. Preconditioning affected peak strain in a loading-rate-dependent manner: responses tapered near 2% preconditioning strain at 0.5% strain/s, whereas no comparable tapering occurred through 15% at 1% strain/s. Compared with human cadaver data, rat samples reached substantially larger peak strains and lower high-strain stiffness. Conclusion: This study provides one of the first direct experimental comparisons of passive pelvic floor muscle mechanics between pregnant and non-pregnant tissue, while systematically evaluating how multiple testing-protocol variables influence the measured response. Distinguishing protocol-dependent variability from biological differences provides a more reproducible basis for cross-study comparison and constitutive model calibration. Rat pelvic floor muscle is a useful model for pregnancy-associated adaptation, but absolute mechanical values should be translated to human tissue cautiously.
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All authors Jose L. Monclova, Matthew Shankel, Ava M. English, Mona Eskandari, Heidi P. Feigenbaum

Biological Physics · Cross-listed from cond-mat.soft

Entropy-Driven Initiation and Cytoskeletal Viscoelasticity in Endocytosis: An Onsager Variational Framework

Receptor-mediated endocytosis requires a particle to approach the cell membrane to within a few nanometers before ligand–receptor binding can occur. Existing continuum models often start from an already established contact and do not explicitly describe how crowding particles on the extracellular side influence the distribution of the particle near the membrane. We examine entropic depletion forces as one possible nonspecific contribution to this initial approach. For ideal depletants, the Asakura–Oosawa excluded-volume construction gives an exact depletion potential for the planar geometry before contact. The potential and force vanish continuously at the onset of excluded-volume overlap. This interaction provides a possible contribution to membrane proximity before specific binding, while its extension to curved wrapping geometries requires additional approximation. Within a reduced continuum model, we combine depletion attraction, ligand–receptor binding, membrane deformation, and cytoskeletal viscoelastic dissipation. The viscoelastic contact is formulated through a hereditary integral and a standard linear solid. The kinetic model gives a conditional minimum ligand density for complete engulfment, a finite particle-size window, and a stiffness-dependent upper limit. When the stationary radius lies inside the domain of finite positive wrapping times, the estimated wrapping time has a minimum at a radius that decreases with increasing binding energy density. At fixed viscosity and other independent parameters, the same time approximation predicts slower wrapping as cell stiffness increases. The two positive roots defining the size window merge at a limiting parameter value, which characterizes closure of the admissible size interval. Depletion attraction is interpreted as one possible contribution to particle-membrane association
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Biological Physics

Traveling Waves Enhance Transport during Cytoplasmic Streaming in Drosophila Oogenesis

Cytoplasmic streaming is believed crucial to large developing cells, such as oocytes, where flow provides a much faster route to transport and mixing of cellular components than molecular diffusion. Molecular motors carrying cargoes on cytoskeletal elements are known to drive streaming in a variety of biological systems and, in Drosophila oocytes, cell-spanning vortical flows are tied to kinesin motors moving on dense beds of microtubules. Recent theories and simulations suggest such streaming self-organizes through interacting microtubules, motors, and flow. Whether these theoretically derived flows are adequate to the tasks of transport and mixing has been unclear. Here we report on new observations of coherent waves traveling persistently through microtubule beds during streaming, and investigate their impact on fluid transport and mixing. Leveraging recent advances in simulating the complex fluid-structure problems of cellular environments, we probe a biophysical model set in an oocytal geometry. Our simulations identify previously unknown waving states that show concordance with our experimental observation and the capacity for functional transport and mixing.
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All authors Olenka Jain, David B. Stein, Reza Farhadifar, Elizabeth R. Gavis, Stanislav Y. Shvartsman, Michael J. Shelley

Biological Physics

The nanoscopic submyelin space mediates efficient K$^+$ uptake and shapes nodal action potentials

Myelin membranes are tightly connected to the axon, forming an electrical route for rapid saltatory conduction of action potentials from node to node. Beneath the myelin, adjacent to the node, lie voltage-gated potassium (K$^+$) channels, whose fundamental role in conduction remains poorly understood. Constrained by biological data, we developed a finite element model of a node of Ranvier encompassing an adaxonal membrane separating oligodendrocyte cytoplasm from the periaxonal space, and solved the full coupled set of physical equations governing electrodiffusion and voltage profiles at the nanoscale. Remarkably, we find that nodal action potentials invert at the inner adaxonal myelin membrane. The hyperpolarization-driven opening of myelinic Kir4.1 channels produces an instantaneous, powerful K$^+$ prebuffering, clamping the periaxonal K$^+$ concentration and enabling high-frequency action potential generation. Together, these findings indicate that the periaxon maintains an electrical feedback loop with the node, and suggest a key evolutionary advantage of juxtaparanodal K$^+$ channel positioning.
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All authors T. M. Kamsma, R. van Roij, C. Spitoni, M. H. P. Kole

Biological Physics

Coherence in Biological Systems

When does a collection of autonomous cells become a multicellular individual? We propose that coherence provides a physical description of this transition. Coherence is treated as a global property arising when distinguishable constituents admit a physically meaningful collective state-space description. Using the center of mass and an interaction-based construction, we show that such collective states can be defined for classical bodies before dynamics is introduced, with normal modes emerging as a particular dynamical realization. We apply this framework to multicellular organization, where cells retain their identities while their independent individuality is replaced by participation in the organized whole. In Dictyostelium discoideum, cAMP-mediated coupling produces population-level collective modes, while starvation provides an experimentally controlled energetic constraint on the transition to multicellularity. The framework yields direct tests through interaction-derived collective eigenstates and the energetic cost of maintaining autonomous versus collective organization. Coherence may thus provide a general physical description of multicellular individuality without requiring microscopic quantum coherence or intrinsic wave character.
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Biological Physics · Cross-listed from physics.optics

Optical-Memory Transport Imaging: Extension to Stochastic Diffusion

Finite-memory optical tracers can encode transport properties through their previously experienced excitation. Using such memory features, harboured in the population of excitable long-lived electronic states within the emitters, instantaneous localization assessments are not needed to determine their transport properties. Here we formulate this principle for monitoring of stochastic transport extracted from transport-history integral over conditional transport histories. Under structured illumination, this history yields a measurable transfer-function response from which scalar and tensor diffusion are reconstructed without time-resolved acquisition. Fisher-information analysis identifies the optimal operating regime and yields first-principles, parameter-free predictions of scalar and tensor reconstruction precision, in quantitative agreement with independent Monte Carlo simulations. These results establish finite optical memory as a general principle for stochastic transport imaging.
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Biological Physics

How do incorrect ligands help detect a correct ligand?

Intrigued by the response of T cell receptors to the presence of a few agonist ligands, we propose a minimal model that can achieve similar performance. The model consists of a small cluster of immobile receptors that bind reversibly to two types (correct/incorrect) of ligands in the environment, with slightly weaker binding strength for the incorrect one. It features binding-state coupling between nearest-neighbor receptors, and receptors in the bound/free states are activated/deactivated by specific enzymes, with rates that allow kinetic proofreading. It is found that, for a range of binding-state coupling strength, incorrect ligands alone cannot activate the receptors, but the binding of merely one correct ligand to a receptor is sufficient to promote the activation of other receptors via induced binding to incorrect ligands. Both response time and signal amplification increase as the receptor binding-state coupling strength increases until it reaches an optimal range to achieve the most rapid and sensitive response. These results suggest a possible mechanism for a speedy and specific response of receptors to very few correct ligands in biological and artificial systems at the subcellular scale.
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Monday 21 September

Biological Physics · Cross-listed from q-bio.PE

Stochastic Field Theory of HIV Latency: Instanton Dynamics and the Path to Viral Rebound

The transition from clinical latency to active HIV infection is a stochastic process. Traditional deterministic models based on ordinary differential equations (ODEs) fail to capture the extinction boundary and the subsequent rebound of the virus because they neglect demographic noise in the small-population limit. We map the resulting Master Equation onto a coherent-state path integral using the Doi-Peliti second-quantization formalism and verify that the stochastic Hamiltonian reproduces the correct mean-field ODEs on the classical manifold $\bar{\phi}_i = 1$. In the semiclassical (large system-size) limit we derive eight coupled Hamilton-Jacobi equations, identify the virus-free and endemic fixed points, and obtain the basic reproduction number $R_0$ for the four-species network. For the analytically tractable case of single-virion bursting ($n=1$) we show that the stochastic Hamiltonian factorizes into two bilinear terms, exposing a non-trivial zero-energy surface that constitutes the instanton trajectory. A quasi-steady-state reduction then yields a first-order linear ODE whose closed-form solution gives the instanton action $S_{\rm inst}$ in terms of biological parameters. Calibrating to clinical data—infected-cell half-life $\delta_I^{-1}\approx 2\,\rm d$, viral burst size $N\approx 10^3$, clearance rate $\delta_V\approx 23\,\rm d^{-1}$, reactivation rate $\eta\approx 10^{-3}\,\rm d^{-1}$, and latent reservoir size $L_0\approx 10^6$ cells—we obtain a mean first-passage time (MFPT) to viral rebound of $\approx 6$ months for a typical patient, with a range of days to years depending on reservoir size. These predictions are qualitatively consistent with observed post-interruption rebound timescales.
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Biological Physics

Towards Accurate Prediction of Mutation-Induced Changes in Protein Structure

Proteins can possess numerous mutations relative to their wild-type amino acid sequences with minimal impact to their structure and function. However, in other cases, even a single amino acid mutation relative to the wild-type sequence can lead to a large change in structure or even a disease phenotype. While the accuracy of wild-type protein structure prediction has improved significantly in recent years, it remains difficult to accurately predict the structure of mutant proteins. Here, we characterize the local mutation-induced structural changes in proteins for a dataset of wildtype and the corresponding single-amino acid mutant x-ray crystal structures from the Protein Data Bank (PDB). We find that mutation-induced structural changes in these proteins are localized at the site of the mutation, decaying rapidly with increasing spatial distance from the mutation site. In addition, we evaluate how well AlphaFold3 can recapitulate the observed mutation-induced structural deformations in the x-ray crystal structures. We find that the accuracy of the AlphaFold3 predictions decreases strongly with increasing mutation-induced deformation. In contrast to the results for AlphaFold3, the Pearson correlation between a single physical feature, i.e. the change in solvent accessibility, and the mutation-induced deformation does not depend on the magnitude of the deformation. Our results and analyses provide a framework for further studies aimed at predicting the structural changes in proteins caused by single amino acid mutations.
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