arXiv subjects

Ramin Golestanian

Publications and source records attributed to Ramin Golestanian.

At least 91 records · Page 5Linked to original sources

arXiv2021

Stochastic dynamics of chemotactic colonies with logistic growth

The interplay between cellular growth and cell-cell signaling is essential for the aggregation and proliferation of bacterial colonies, as well as for the self-organization of cell tissues. To investigate this interplay, we focus here on the collective properties of dividing chemotactic cell colonies by studying their long-time and large-scale dynamics through a renormalization group (RG) approach. The RG analysis reveals that a relevant but unconventional chemotactic interaction -- corresponding to a polarity-induced mechanism -- is generated by fluctuations at macroscopic scales, even when an underlying mechanism is absent at the microscopic level. This emerges from the interplay of the well-known Keller--Segel (KS) chemotactic nonlinearity and cell birth and death processes. At one-loop order, we find no stable fixed point of the RG flow equations. We discuss a connection between the dynamics investigated here and the celebrated Kardar--Parisi--Zhang (KPZ) equation with long-range correlated noise, which points at the existence of a strong-coupling, nonperturbative fixed point.

cond-mat.stat-mech↗

arXiv2021

Synchronization and enhanced catalysis of mechanically coupled enzymes

We examine the stochastic dynamics of two enzymes that are mechanically coupled to each other, e.g., through an elastic substrate or a fluid medium. The enzymes undergo conformational changes during their catalytic cycle, which itself is driven by stochastic steps along a biased chemical free energy landscape. We find conditions under which the enzymes can synchronize their catalytic steps, and discover that the coupling can lead to a significant enhancement in their overall catalytic rate. Both effects can be understood as arising from a global bifurcation in the underlying dynamical system at sufficiently strong coupling. Our findings suggest that, despite their molecular scale, enzymes can be cooperative and improve their performance in metabolic clusters.

cond-mat.stat-mech↗

arXiv2021

Comment on "Relative Diffusivities of Bound and Unbound Protein Can Control Chemotactic Directionality''

This is a comment on a recently published paper in Langmuir: Mandal, N. S.; Sen, A. Relative Diffusivities of Bound and Unbound Protein Can Control Chemotactic Directionality. Langmuir 2021, PMID: 34647749 [arXiv:2103.13469]. In this study, Mandal and Sen claim to propose a new kinetic model to analyze the directional movement of enzyme molecules in response to a gradient of their substrate, with the supposedly new prediction that net movement occurs up the substrate gradient when the diffusivity of the substrate-bound enzyme is lower than that of the unbound enzyme, and movement down the substrate gradient when the diffusivity of the substrate-bound enzyme is higher than that of the unbound enzyme. They develop this theoretical scheme to present an alternative to our previously published theoretical framework [Agudo-Canalejo, J.; Illien, P.; Golestanian, R. Phoresis and enhanced diffusion compete in enzyme chemotaxis. Nano Lett. 2018, 18, 2711-2717; arXiv:2104.02394]. We demonstrate that despite their claim, what they present is exactly the same theory as our work, and therefore, we conclude that their claim of novelty is unsubstantiated.

physics.bio-ph↗

arXiv2021

Molecular Diffusivity of Click Reaction Components: The Diffusion Enhancement Question

Micrometer-sized objects are widely known to exhibit chemically-driven motility in systems away from equilibrium. Experimental observation of reaction-induced motility or enhancement in diffusivity at the much shorter length scale of small molecules is however still a matter of debate. Here, we investigate the molecular diffusivity of reactants, catalyst and product of a model reaction, the copper-catalyzed azide-alkyne cycloaddition click reaction, and develop new NMR diffusion approaches that allow the probing of reaction-induced diffusion enhancement in nano-sized molecular systems with higher precision than the state of the art. Following two different approaches that enable the accounting of time-dependent concentration changes during NMR experiments, we closely monitored the diffusion coefficient of reaction components during the reaction. The reaction components showed distinct changes in the diffusivity: while the two reactants underwent a time-dependent decrease in their diffusivity, the diffusion coefficient of the product gradually increased and the catalyst showed only slight diffusion enhancement within the range expected for reaction-induced sample heating. The decrease in diffusion coefficient of the alkyne, one of the two reactants of click reaction, was not reproduced during its copper coordination when the second reactant, azide, was absent. Our results do not support the catalysis-induced diffusion enhancement of the components of click reaction and, instead, point to the role of a relatively large intermediate species within the reaction cycle with diffusivity lower than both the reactants and product molecule.

physics.chem-ph↗

arXiv2021

Non-equilibrium phase separation in mixtures of catalytically active particles: size dispersity and screening effects

Biomolecular condensates in cells are often rich in catalytically-active enzymes. This is particularly true in the case of the large enzymatic complexes known as metabolons, which contain different enzymes that participate in the same catalytic pathway. One possible explanation for this self-organization is the combination of the catalytic activity of the enzymes and a chemotactic response to gradients of their substrate, which leads to a substrate-mediated effective interaction between enzymes. These interactions constitute a purely non-equilibrium effect and show exotic features such as non-reciprocity. Here, we analytically study a model describing the phase separation of a mixture of such catalytically-active particles. We show that a Michaelis-Menten-like dependence of the particles' activities manifests itself as a screening of the interactions, and that a mixture of two differently-sized active species can exhibit phase separation with transient oscillations. We also derive a rich stability phase diagram for a mixture of two species with both concentration-dependent activity and size dispersity. This work highlights the variety of possible phase separation behaviours in mixtures of chemically-active particles, which provides an alternative pathway to the passive interactions more commonly associated with phase separation in cells. Our results highlight non-equilibrium organizing principles that can be important for biologically relevant liquid-liquid phase separation.

q-bio.SC↗

arXiv2021

Diffusiophoretic propulsion of an isotropic active colloidal particle near a finite-sized disk embedded in a planar fluid-fluid interface

Breaking spatial symmetry is an essential requirement for phoretic active particles to swim at low Reynolds number. This fundamental prerequisite for swimming at the micro-scale is fulfilled either by chemical patterning of the surface of active particles or alternatively by exploiting geometrical asymmetries to induce chemical gradients and achieve self-propulsion. In the present manuscript, a far-field analytical model is employed to quantify the leading-order contribution to the induced phoretic velocity of a chemically homogeneous isotropic active colloid near a finite-sized disk of circular shape resting on an interface separating two immiscible viscous incompressible Newtonian fluids. To this aim, the solution of the phoretic problem is formulated as a mixed-boundary-value problem which is subsequently transformed into a system of dual integral equations on the inner and outer domains. Depending on the ratio of different involved viscosities and solute solubilities, the sign of phoretic mobility and chemical activity, as well as the ratio of particle-interface distance to the radius of the disk, the isotropic active particle is found to be either repelled from the interface, attracted to it, or reach a stable hovering state and remains immobile near the interface. Our results may prove useful in controlling and guiding the motion of self-propelled phoretic active particles near aqueous interfaces.

cond-mat.soft↗

arXiv2021

A topological fluctuation theorem

Fluctuation theorems specify the non-zero probability to observe negative entropy production, contrary to a naive expectation from the second law of thermodynamics. For closed particle trajectories in a fluid, Stokes theorem can be used to give a geometric characterization of the entropy production. Building on this picture, we formulate a topological fluctuation theorem that depends only by the winding number around each vortex core and is insensitive to other aspects of the force. The probability is robust to local deformations of the particle trajectory, reminiscent of topologically protected modes in various classical and quantum systems. We demonstrate that entropy production is quantized in these strongly fluctuating systems, and it is controlled by a topological invariant. We demonstrate that the theorem holds even when the probability distributions are non-Gaussian functions of the generated heat.

cond-mat.stat-mech↗

arXiv2021

Topology protects chiral edge currents in stochastic systems

Constructing systems that exhibit time-scales much longer than those of the underlying components, as well as emergent dynamical and collective behavior, is a key goal in fields such as synthetic biology and materials self-assembly. Inspiration often comes from living systems, in which robust global behavior prevails despite the stochasticity of the underlying processes. Here, we present two-dimensional stochastic networks that consist of minimal motifs representing out-of-equilibrium cycles at the molecular scale and support chiral edge currents in configuration space. These currents arise in the topological phase due to the bulk-boundary correspondence and dominate the system dynamics in the steady-state, further proving robust to defects or blockages. We demonstrate the topological properties of these networks and their uniquely non-Hermitian features such as exceptional points and vorticity, while characterizing the edge state localization. As these emergent edge currents are associated to macroscopic timescales and length scales, simply tuning a small number of parameters enables varied dynamical phenomena including a global clock, dynamical growth and shrinkage, and synchronization. Our construction provides a novel topological formalism for stochastic systems and fresh insights into non-Hermitian physics, paving the way for the prediction of robust dynamical states in new classical and quantum platforms.

cond-mat.stat-mech↗

arXiv2021

Optimal swimmer can be puller, pusher, or neutral depending on the shape

The ability of microswimmers to deploy optimal propulsion strategies is of paramount importance for their locomotory performance and survival at low Reynolds numbers. Although for perfectly spherical swimmers minimum dissipation requires a neutral type swimming, any departure from the spherical shape may lead the swimmer to adopt a new propulsion strategy, namely those of puller- or pusher-type swimming. In this study, by using the minimum dissipation theorem for microswimmers, we determine the flow field of an optimal nearly spherical swimmer, and show that indeed depending on the shape profile, the optimal swimmer can be a puller, pusher, or neutral. Using an asymptotic approach, we find that amongst all the modes of the shape function, only the third mode determines to leading order the swimming type of the optimal swimmer.

physics.flu-dyn↗

arXiv2021

Transient fluctuation-induced forces in driven electrolytes after an electric field quench

Understanding how electrolyte solutions behave out of thermal equilibrium is a long-standing endeavor in many areas of chemistry and biology. Although mean-field theories are widely used to model the dynamics of electrolytes, it is also important to characterize the effects of fluctuations in these systems. In a previous work, we showed that the dynamics of the ions in a strong electrolyte that is driven by an external electric field can generate long-ranged correlations manifestly different from the equilibrium screened correlations; in the nonequilibrium steady state, these correlations give rise to a novel long-range fluctuation-induced force (FIF). Here, we extend these results by considering the dynamics of the strong electrolyte after it is quenched from thermal equilibrium upon the application of a constant electric field. We show that the asymptotic long-distance limit of both charge and density correlations is generally diffusive in time. These correlations give rise to long-ranged FIFs acting on the neutral confining plates with long-time regimes that are governed by power-law temporal decays toward the steady-state value of the force amplitude. These findings show that nonequilibrium fluctuations have nontrivial implications on the dynamics of objects immersed in a driven electrolyte, and they could be useful for exploring new ways of controlling long-distance forces in charged solutions.

cond-mat.soft↗

arXiv2021

'Fuelled' motion: phoretic motility and collective behaviour of active colloids

Designing microscopic and nanoscopic self-propelled particles and characterising their motion has become a major scientific challenge over the past decades. To this purpose, phoretic effects, namely propulsion mechanisms relying on local field gradients, have been the focus of many theoretical and experimental studies. In this review, we adopt a tutorial approach to present the basic physical mechanisms at stake in phoretic motion, and describe the different experimental works that lead to the fabrication of active particles based on this principle. We also present the collective effects observed in assemblies of interacting active colloids, and the theoretical tools that have been used to describe phoretic and hydrodynamic interactions.

cond-mat.soft↗

arXiv2021

Enhanced diffusion and chemotaxis at the nanoscale

Enzymes have been recently proposed to have mechanical activity associated with their chemical activity. In a number of recent studies, it has been reported that enzymes undergo enhanced diffusion in the presence of their corresponding substrate, when this substrate is uniformly distributed in solution. Moreover, if the concentration of the substrate is non-uniform, enzymes and other small molecules have been reported to show chemotaxis -- biased stochastic movement in the direction of the substrate gradient -- typically towards higher concentrations of this substrate, with a few exceptions. The underlying physical mechanisms responsible for enhanced diffusion and chemotaxis at the nanoscale, however, are still not well understood. Understanding these processes is important both for fundamental biological research, e.g. in the context of spatial organization of enzymes in metabolic pathways (metabolon formation), as well as for engineering applications, such as in the design of new vehicles for targeted drug delivery. In this Account, we will review the available experimental observations of both enhanced diffusion and chemotaxis, and we will discuss critically the different theories that have been proposed to explain the two. We first focus on enhanced diffusion, beginning with an overview of the experimental results. We then discuss the two main types of mechanisms that have been proposed, namely active mechanisms relying on the catalytic step of the enzymatic reaction, and equilibrium mechanisms which consider the reversible binding and unbinding of the substrate to the enzyme. We put particular emphasis on an equilibrium model recently introduced by us, which describes how the diffusion of dumbbell-like modular enzymes can be enhanced in the presence of substrate, thanks to a binding-induced reduction of the internal fluctuations of the enzyme. We then turn to chemotaxis, [...]

physics.bio-ph↗

arXiv2021

Phoresis and Enhanced Diffusion Compete in Enzyme Chemotaxis

Chemotaxis of enzymes in response to gradients in the concentration of their substrate has been widely reported in recent experiments, but a basic understanding of the process is still lacking. Here, we develop a microscopic theory for chemotaxis, valid for enzymes and other small molecules. Our theory includes both non-specific interactions between enzyme and substrate, as well as complex formation through specific binding between the enzyme and the substrate. We find that two distinct mechanisms contribute to enzyme chemotaxis: a diffusiophoretic mechanism due to the non-specific interactions, and a new type of mechanism due to binding-induced changes in the diffusion coefficient of the enzyme. The latter chemotactic mechanism points towards lower substrate concentration if the substrate enhances enzyme diffusion, and towards higher substrate concentration if the substrate inhibits enzyme diffusion. For a typical enzyme, attractive phoresis and binding-induced enhanced diffusion will compete against each other. We find that phoresis dominates above a critical substrate concentration, whereas binding-induced enhanced diffusion dominates for low substrate concentration. Our results resolve an apparent contradiction regarding the direction of urease chemotaxis observed in experiments, and in general clarify the relation between enhanced diffusion and chemotaxis of enzymes. Finally, we show that the competition between the two distinct chemotactic mechanisms may be used to engineer nanomachines that move towards or away from regions with a specific substrate concentration.

physics.bio-ph↗

arXiv2021

Designing metachronal waves of cilia

On surfaces with many motile cilia, beats of the individual cilia coordinate to form metachronal waves. We present a theoretical framework that connects the dynamics of an individual cilium to the collective dynamics of a ciliary carpet via systematic coarse-graining. We uncover the criteria that control the selection of frequency and wavevector of stable metchacronal waves of the cilia and examine how they depend on the geometric and dynamical characteristics of single cilia, as well as the geometric properties of the array. We perform agent-based numerical simulations of arrays of cilia with hydrodynamic interactions and find quantitative agreement with the predictions of the analytical framework. Our work sheds light on the question of how the collective properties of beating cilia can be determined using information about the individual units, and as such exemplifies a bottom-up study of a rich active matter system.

cond-mat.soft↗

arXiv2021

Long-range fluctuation-induced forces in driven electrolytes

We study the stochastic dynamics of an electrolyte driven by a uniform external electric field and show that it exhibits generic scale invariance despite the presence of Debye screening. The resulting long-range correlations give rise to a Casimir-like fluctuation-induced force between neutral boundaries that confine the ions; this force is controlled by the external electric field, and it can be both attractive and repulsive with similar boundary conditions, unlike other long-range fluctuation-induced forces. This work highlights the importance of nonequilibrium correlations in electrolytes and shows how they can be used to tune interactions between uncharged biological or synthetic structures at large separations.

cond-mat.soft↗

arXiv2021

Active beating of a reconstituted synthetic minimal axoneme

Propelling microorganisms through fluids and moving fluids along cellular surfaces are essential biological functions accomplished by long, thin structures called motile cilia and flagella, whose regular, oscillatory beating breaks the time-reversal symmetry required for transport. Although top-down experimental approaches and theoretical models have allowed us to broadly characterize such organelles and propose mechanisms underlying their complex dynamics, constructing minimal systems capable of mimicking ciliary beating and identifying the role of each component remains a challenge. Here we report the bottom-up assembly of a minimal synthetic axoneme, which we call a synthoneme, using biological building blocks from natural organisms, namely pairs of microtubules and cooperatively associated axonemal dynein motors. We show that upon provision of energy by ATP, microtubules undergo rhythmic bending by cyclic association-dissociation of dyneins. Our simple and unique beating minimal synthoneme represents a self-organized nanoscale biomolecular machine that can also help understand the mechanisms underlying ciliary beating.

cond-mat.soft↗

arXiv2021

Scaling regimes of active turbulence with external dissipation

Active fluids exhibit complex turbulent-like flows at low Reynolds number. Recent work predicted that 2d active nematic turbulence follows universal scaling laws. However, experimentally testing these predictions is conditioned by the coupling to the 3d environment. Here, we measure the spectrum of the kinetic energy, $E(q)$, in an active nematic film in contact with a passive oil layer. At small and intermediate scales, we find the scaling regimes $E(q)\sim q^{-4}$ and $E(q)\sim q^{-1}$, respectively, in agreement with the theoretical prediction for 2d active nematics. At large scales, however, we find a new scaling $E(q)\sim q$, which emerges when the dissipation is dominated by the 3d oil layer. In addition, we derive an explicit expression for the spectrum that spans all length scales, thus explaining and connecting the different scaling regimes. This allows us to fit the data and extract the length scale that controls the crossover to the new large-scale regime, which we tune by varying the oil viscosity. Overall, our work experimentally demonstrates the emergence of universal scaling laws in active turbulence, and it establishes how the spectrum is affected by external dissipation.

cond-mat.soft↗

arXiv2021

Minimum Dissipation Theorem for Microswimmers

We derive a theorem for the lower bound on the energy dissipation rate by a rigid surface-driven active microswimmer of arbitrary shape in a fluid at low Reynolds number. We show that, for any swimmer, the minimum dissipation at a given velocity can be expressed in terms of the resistance tensors of two passive bodies of the same shape with a no-slip and perfect-slip boundary. To achieve the absolute minimum dissipation, the optimal swimmer needs a surface velocity profile that corresponds to the flow around the perfect-slip body, and a propulsive force density that corresponds to the no-slip body. Using this theorem, we propose an alternative definition of the energetic efficiency of microswimmers that, unlike the commonly-used Lighthill efficiency, can never exceed unity. We validate the theory by calculating the efficiency limits of spheroidal swimmers.

physics.flu-dyn↗