arXiv subjects

Amin Doostmohammadi

Publications and source records attributed to Amin Doostmohammadi.

At least 37 records · Page 2Linked to original sources

arXiv2025

Multi-phase-field Models of Biological Tissues

The development of complex multicellular organisms from a single parent cell is a highly orchestrated process that cells conduct collectively without central guidance, creating intricate dynamic patterns essential for development and regeneration. Despite significant advances in imaging spatiotemporal dynamics of cell collectives and mechanical characterization techniques, the role of physical forces in biological functions remains poorly understood. Physics-based models are crucial in complementing experiments, providing high-resolution spatiotemporal fields in three dimensions. This review focuses on dense, soft multicellular systems, such as tissues, where mechanical deformation of one cell necessitates the re-organization of neighboring cells. The multi-phase-field model offers a rich physics-based framework to advance our understanding of biological systems and provides a robust playground for non-equilibrium physics of active matter. We discuss the foundational aspects of the multi-phase-field model and their applications in understanding physics of active matter. We also explore the integration of biological physics with experimental data, covering cell migration, heterogeneous cell populations, and confined systems. Finally, we highlight current trends, the importance of multi-phase-field models in biological and physics research, and future challenges.

cond-mat.soft↗

arXiv2025

Beyond Dipolar Activity: Quadrupolar Stress Drives Collapse of Nematic Order on Frictional Substrates

The field of active nematics has traditionally employed descriptions based on dipolar activity, with interactions that align along a single axis. However, it has been theoretically predicted that interactions with a substrate, prevalent in most biological systems, would require novel forms of activity, such as quadrupolar activity, that are governed by hydrodynamic screening. Here, by combining experiments and numerical simulations, we show that upon light-induced solidification of the underlying medium, microtubule-kinesin mixtures undergo a transformation that leads to a biphasic active suspension. Using an active lyotropic model, we prove that the transition is governed by screening effects that alter the dominant form of active stress. Specifically, the combined effect of friction and quadrupolar activity leads to a hierarchical folding that follows the intrinsic bend instability of the active nematic layer. Our results demonstrate the dynamics of the collapse of orientational order in active nematics and present a new route for controlling active matter by modifying the activity through changing the surrounding environment.

cond-mat.soft↗

arXiv2025

Phase diagram, confining strings, and a new universality class in nematopolar matter

We study a minimal model of a system with coexisting nematic and polar orientational orders, where one field tends to order and the other prefers isotropy. For strong coupling, the ordered field aligns the isotropic one, locking their orientations. The phase diagram reveals three distinct phases--nematopolar (aligned orders), nematic (independent orders), and isotropic (vanishing orders)--separated by continuous and discontinuous transitions, including a triple and a tricritical point. We find unique critical scaling for the nematopolar-nematic transition, distinct from standard nematic or polar universality classes. Additionally, in the locked nematopolar phase, we show nematic $+1/2$ topological defect pairs are connected and confined by strings with constant tension. These strings arise from frustration in locking the orientational orders and can be interpreted as elongated cores of $+1$ polar topological defects. When a sufficiently strong background field couples to the polar order, all topological defects are expelled from the region. Analytical predictions are quantitatively confirmed by numerical simulations.

cond-mat.soft↗

arXiv2024

Transport of topological defects in a biphasic mixture of active and passive nematic fluids

Collectively moving cellular systems often contain a proportion of dead cells or non-motile genotypes. When mixed, nematically aligning motile and non-motile agents are known to segregate spontaneously. However, the role that topological defects and active stresses play in shaping the distribution of the two phases remains unresolved. In this study, we investigate the behaviour of a two-dimensional binary mixture of active and passive nematic fluids to understand how topological defects are transported between the two phases and, ultimately, how this leads to the segregation of topological charges. When the activity of the motile phase is large, and the tension at the interface of motile and non-motile phases is weak, we find that the active phase tends to accumulate $+1/2$ defects and expel $-1/2$ defects so that the motile phase develops a net positive charge. Conversely, when the activity of the motile phase is comparatively small and interfacial tension is strong, the opposite occurs so that the active phase develops a net negative charge. We then use these simulations to develop a physical intuition of the underlying processes that drive the charge segregation. Lastly, we quantify the sensitivity of this process on the other model parameters, by exploring the effect that anchoring strength, orientational elasticity, friction, and volume fraction of the motile phase have on topological charge segregation. As $+1/2$ and $-1/2$ defects have very different effects on interface morphology and fluid transport, this study offers new insights into the spontaneous pattern formation that occurs when motile and non-motile cells interact.

physics.bio-ph↗

arXiv2024

Active particles knead three-dimensional gels into open crumbs

Colloidal gels are prime examples of functional materials exhibiting disordered, amorphous, yet meta-stable forms. They maintain stability through short-range attractive forces and their material properties are tunable by external forces. Combining persistent homology analyses and simulations of three-dimensional colloidal gels doped with active particles, we reveal novel dynamically evolving structures of colloidal gels. Specifically, we show that the local injection of energy by active dopants can lead to highly porous, yet compact gel structures that can significantly affect the transport of active particles within the modified colloidal gel. We further show the substantially distinct structural behaviour between active doping of 2D and 3D systems by revealing how passive interfaces play a topologically different role in interacting with active particles in two and three dimensions. The results open the door to an unexplored prospect of forming a wide variety of compact but highly heterogeneous and percolated porous media through active doping of 3D passive matter, with diverse implications in designing new functional materials to active ground remediation.

cond-mat.soft↗

arXiv2024

Evidence of robust, universal conformal invariance in living biological matter

Collective cellular movement plays a crucial role in many processes fundamental to health, including development, reproduction, infection, wound healing, and cancer. The emergent dynamics that arise in these systems are typically thought to depend on how cells interact with one another and the mechanisms used to drive motility, both of which exhibit remarkable diversity across different biological systems. Here, we report experimental evidence of a universal feature in the patterns of flow that spontaneously emerges in groups of collectively moving cells. Specifically, we demonstrate that the flows generated by collectively moving dog kidney cells, human breast cancer cells, and by two different strains of pathogenic bacteria, all exhibit conformal invariance. Remarkably, not only do our results show that all of these very different systems display robust conformal invariance, but we also discovered that the precise form of the invariance in all four systems is described by the Schramm-Loewner Evolution (SLE), and belongs to the percolation universality class. A continuum model of active matter can recapitulate both the observed conformal invariance and SLE form found in experiments. The presence of universal conformal invariance reveals that the macroscopic features of living biological matter exhibit universal translational, rotational, and scale symmetries that are independent of the microscopic properties of its constituents. Our results show that the patterns of flows generated by diverse cellular systems are highly conserved and that biological systems can unexpectedly be used to experimentally test predictions from the theories for conformally invariant structures

cond-mat.soft↗

arXiv2023

Elasticity tunes mechanical stress localization around active topological defects

Mechanical stresses are increasingly found to be associated with various biological functionalities. At the same time, topological defects are being identified across a diverse range of biological systems and are points of localized mechanical stress. It is therefore important to ask how mechanical stress localization around topological defects is controlled. Here, we use continuum simulations of nonequilibrium, fluctuating and active nematics to explore the patterns of stress localization, as well as their extent and intensity around topological defects. We find that by increasing the orientational elasticity of the material, the isotropic stress pattern around topological defects is changed substantially from a stress dipole characterized by symmetric compression-tension regions around the core of the defect to a localized stress monopole at the defect position. Moreover, we show that elastic anisotropy alters the extent and intensity of the stresses, and can result in the dominance of tension or compression around defects. Finally, including both nonequilibrium fluctuations and active stress generation, we find that the elastic constant tunes the relative effect of each, leading to the flipping of tension and compression regions around topological defects. This flipping of the tension-compression regions only by changing the elastic constant presents an interesting, simple, way of switching the dynamic behavior in active matter by changing a passive material property. We expect these findings to motivate further exploration tuning stresses in active biological materials by varying material properties of the constituent units.

cond-mat.soft↗

arXiv2023

Periodic orbits, pair nucleation, and unbinding of active nematic defects on cones

Geometric confinement and topological constraints present promising means of controlling active materials. By combining analytical arguments derived from the Born-Oppenheimer approximation with numerical simulations, we investigate the simultaneous impact of confinement together with curvature singularity by characterizing the dynamics of an active nematic on a cone. Here, the Born-Oppenheimer approximation means that textures can follow defect positions rapidly on the time scales of interest. Upon imposing strong anchoring boundary conditions at the base of a cone, we find a a rich phase diagram of multi-defect dynamics including exotic periodic orbits of one or two $+1/2$ flank defects, depending on activity and non-quantized geometric charge at the cone apex. By characterizing the transitions between these ordered dynamical states, we can understand (i) defect unbinding, (ii) defect absorption and (iii) defect pair nucleation at the apex. Numerical simulations confirm theoretical predictions of not only the nature of the circular orbits but also defect unbinding from the apex.

cond-mat.soft↗

arXiv2023

Spontaneous Self-Constraint in Active Nematic Flows

Active processes drive and guide biological dynamics across scales -- from subcellular cytoskeletal remodelling, through tissue development in embryogenesis, to population-level bacterial colonies expansion. In each of these, biological functionality requires collective flows to occur while self-organized structures are protected; however, the mechanisms by which active flows can spontaneously constrain their dynamics to preserve structure have not previously been explained. By studying collective flows and defect dynamics in active nematic films, we demonstrate the existence of a self-constraint -- a two-way, spontaneously arising relationship between activity-driven isosurfaces of flow boundaries and mesoscale nematic structures. Our results show that self-motile defects are tightly constrained to viscometric surfaces -- contours along which vorticity and strain-rate balance. This in turn reveals that self-motile defects break mirror symmetry when they move along a single viscometric surface, in contrast with expectations. This is explained by an interdependence between viscometric surfaces and bend walls -- elongated narrow kinks in the orientation field. Although we focus on extensile nematic films, numerical results show the constraint holds whenever activity leads to motile half-charge defects. This mesoscale cross-field self-constraint offers a new framework for tackling complex 3D active turbulence, designing dynamic control into biomimetic materials, and understanding how biological systems can employ active stress for dynamic self-organization.

cond-mat.soft↗

arXiv2023

Symmetry-restoring crossover from defect-free to defect-laden turbulence in polar active matter

Coherent flows of self-propelled particles are characterized by vortices and jets that sustain chaotic flows, referred to as active turbulence. Here, we reveal a crossover between defect-free active turbulence and active turbulence laden with topological defects. Interestingly, we show that concurrent to the crossover from defect-free to defect-laden active turbulence is the restoration of the previously broken $\SO(2)$-symmetry signaled by the fast decay of the two-point correlations. By stability analyses of the topological charge density field, we provide theoretical insights on the criterion for the crossover to the defect-laden active turbulent state. Despite the distinct symmetry features between these two active turbulence regimes, the flow fluctuations exhibit universal statistical scaling behaviors at large scales, while the spectrum of polarity fluctuations decays exponentially at small length scales compared to the active energy injection length. These findings reveal a new dynamical crossover between distinct spatiotemporal organization patterns in polar active mater.

cond-mat.soft↗

arXiv2023

Mechanical basis and topological routes to cell elimination

Cell layers eliminate unwanted cells through the extrusion process, which underlines healthy versus flawed tissue behaviors. Although several biochemical pathways have been identified, the underlying mechanical basis including the forces involved in cellular extrusion remain largely unexplored. Utilizing a phase-field model of a three-dimensional cell layer, we study the interplay of cell extrusion with cell-cell and cell-substrate interactions, in a flat monolayer. Independent tuning of cell-cell versus cell-substrate adhesion forces reveals that extrusion events can be distinctly linked to defects in nematic and hexatic orders associated with cellular arrangements. Specifically, we show that by increasing relative cell-cell adhesion forces the cell monolayer can switch between the collective tendency towards five-fold, hexatic, disclinations relative to half-integer, nematic, defects for extruding a cell. We unify our findings by accessing three-dimensional mechanical stress fields to show that an extrusion event acts as a mechanism to relieve localized stress concentration.

cond-mat.soft↗

arXiv2023

Active topological defect absorption by a curvature singularity

Using the Born-Oppenheimer approximation, we present a general description of topological defects dynamics in $p$-atic materials on curved surfaces, and simplify it in the case of active nematics. We find that activity induces a geometric contribution to the motility of the $+1/2$ defect. Moreover, in the case of a cone, the simplest example of a geometry with curvature singularity, we find that the motility depends on the deficit angle of the cone and changes sign when the deficit angle is bigger than $π$, leading to the change in active behavior from contractile (extensile) to extensile (contractile) behavior. Using our analytical framework, we then identify for positively charged defects the basin of attraction to the cone apex and present closed-form predictions for defect trajectories near the apex. The analytical results are quantitatively corroborated against full numerical simulations. Provided the capture radius is small compared to the cone size, the agreement is excellent.

cond-mat.soft↗

arXiv2023

Spontaneous flows and dynamics of full-integer topological defects in polar active matter

Polar active matter of self-propelled particles sustain spontaneous flows through the full-integer topological defects. We study theoretically the effect of both polar and dipolar active forces on the flow profile around $\pm 1$ defects and their interaction in the presence of both viscosity and frictional dissipation. The vorticity induced by the active stress is non-zero at the $+1$ defect contributing to the active torque acting on the defect. A near-core flow reversal is predicted in absence of hydrodynamic screening (zero friction) as observed in numerical simulations. While $\pm 1$ defects are sources of spontaneous flows due to active stresses, they become sinks of flows induced by the polar active forces. We show analytically that the flow velocity induced by polar active forces increases away from a $\pm 1$ defect towards the uniform far-field, while its associated vorticity field decays as $1/r$ in the far-field. In the friction-dominated regime, we demonstrate that the flow induced by polar active forces enhances defect pair annihilation, and depends only on the orientation between a pair of oppositely charged defects relative to the orientation of the background polarization field. Interestingly, we find that this annihilation dynamics through mutual defect-defect interactions is distance independent, in contradiction with the effect of dipolar active forces which decay inversely proportional to the defect separation distance. As such, our analyses reveals a new, truly long-ranged mechanism for the pairwise interaction of oppositely-charged topological defects in polar active matter.

cond-mat.soft↗

arXiv2022

Distinct impacts of polar and nematic self-propulsion on active unjamming

Though jamming transitions are long studied in condensed matter physics and granular systems, much less is known about active jamming (or unjamming), which commonly takes place in living materials. In this paper, we explore, by molecular dynamic simulations, the jamming-unjamming transition in a dense system of active semi-flexible filaments. In particular we characterise the distinct impact of polar versus nematic driving for different filament rigidity and at varying density. Our results show that high densities of dynamic active filaments can be achieved by only changing the nature of the active force, nematic or polar. Interestingly, while polar driving is more effective at unjamming the system at high densities below confluency, we find that at even higher densities nematic driving enhances unjamming compared to its polar counterpart. The effect of varying the rigidity of filaments is also significantly different in the two cases: while for nematic driving lowering bending rigidity unjams the system, we find an intriguing re-entrant jamming-unjamming-jamming transition for polar driving as the filament rigidity is lowered. While the first transition (unjamming) is driven by softening due to reduced rigidity, the second transition (jamming) is a cooperative effect of ordering and coincides with the emergence of nematic order in the system. Together, through a generic model of self-propelled flexible filaments, our results demonstrate how tuning the nature of self-propulsion and flexibility can be employed by active materials to achieve high densities without getting jammed.

cond-mat.soft↗

arXiv2022

Stress percolation criticality of glass to fluid transition in active cell layers

Using three-dimensional representation of confluent cell layers, we map the amorphous solid to fluid phase transition in active cell layers onto the two-dimensional (2D) site percolation universality class. Importantly, we unify two distinct, predominant, pathways associated with this transition; namely (i) cell-cell adhesion and (ii) active traction forces. For each pathway, we independently vary the corresponding control parameter and focus on the emergent mechanical stress patterns as the monolayer transitions from a glassy- to a fluid-like state. Through finite-size scaling analyses, our results lead us to establish the glassy- to fluid-like transition as a critical phenomena in terms of stress development in the cell layer and show that the associated criticality belongs to the 2D site percolation universality class. Our findings offer a fresh perspective on solid (glass-like) to fluid phase transition in active cell layers and can bridge our understanding of glassy behaviors in active matter with potential implications in biological processes such as wound healing, development, and cancer progression.

cond-mat.soft↗

arXiv2022

Fluctuation-induced dynamics of nematic topological defects

Topological defects are increasingly being identified in various biological systems, where their characteristic flow fields and stress patterns are associated with continuous active stress generation by biological entities. Here, using numerical simulations of continuum fluctuating nematohydrodynamics we show that even in the absence of any specific form of active stresses associated with self-propulsion, mesoscopic fluctuations in either orientational alignment or hydrodynamics can independently result in flow patterns around topological defects that resemble the ones observed in active systems. Our simulations further show the possibility of extensile- and contractile-like motion of fluctuation-induced positive half-integer topological defects. Remarkably, isotropic stress fields also reproduce the experimentally measured stress patterns around topological defects in epithelia. Our findings further reveal that extensile- or contractile-like flow and stress patterns around fluctuation-induced defects are governed by passive elastic stresses and flow-aligning behavior of the nematics.

cond-mat.soft↗

arXiv2022

Collective rotational motion of freely-expanding T84 epithelial cell colonies

Coordinated rotational motion is an intriguing, yet still elusive mode of collective cell migration, which is relevant in pathological and morphogenetic processes. Most of the studies on this topic have been carried out on confined epithelial cells. The driver of collective rotation in such conditions has not been clearly elucidated, although it has been speculated that spatial confinement can play an essential role in triggering cell rotation. Here, we study the growth of epithelial cell colonies freely expanding (i.e., with no physical constraints) on the surface of cell culture plates, a case which has received scarce attention in the literature. We find that coordinated cell rotation spontaneously occurs in cell clusters in the free growth regime, thus implying that cell confinement is not necessary to elicit collective rotation as previously suggested. The collective rotation was size and shape dependent: a highly coordinated disk-like rotation was found in small cell clusters with a round shape, while collective rotation was suppressed in large irregular cell clusters generated by merging of different clusters in the course of their growth. The angular motion was persistent in the same direction, although clockwise and anticlockwise rotations were equally likely to occur among different cell clusters. Radial cell velocity was low as compared to the angular velocity. A clear difference in morphology was observed between cells at the periphery and the ones in the core of the clusters, the former being more elongated and spread out as compared to the latter. Overall, our results provide the first quantitative and systematic evidence that coordinated cell rotation does not require a spatial confinement and occurs spontaneously in freely expanding epithelial cell colonies.

cond-mat.soft↗

arXiv2022

Differential elasticity in lineage segregation of embryonic stem cells

The question of what guides lineage segregation is central to development, where cellular differentiation leads to segregated cell populations destined for specialized functions. Here, using optical tweezers measurements of mouse embryonic stem cells (mESCs), we reveal a mechanical mechanism based on differential elasticity in the second lineage segregation of the embryonic inner cell mass into epiblast (EPI) cells - that will develop into the fetus - and primitive endoderm (PrE) - which will form extraembryonic structures such as the yolk sac. Remarkably, we find that these mechanical differences already occur during priming and not just after a cell has committed to differentiation. Specifically, we show that the mESCs are highly elastic compared to any other reported cell type and that the PrE cells are significantly more elastic than EPI-primed cells. Using a model of two cell types differing only in elasticity we show that differential elasticity alone can lead to segregation between cell types, suggesting that the mechanical attributes of the cells contribute to the segregation process. Our findings present differential elasticity as a previously unknown mechanical contributor to the lineage segregation during the embryo morphogenesis.

physics.bio-ph↗