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We formulate an adaptive version of Kelly's horse model in which the gambler learns from past race results using Bayesian inference. A known asymptotic scaling for the difference between the growth rate of the gambler and the optimal growth rate, known as the gambler'sregret, is recovered. We show how this adaptive strategy is related to the universal portfolio strategy, and we build improved adaptive strategies in which the gambler exploits information contained in the bookmaker odds distribution to reduce his/her initial loss of the capital during the learning phase., Comment: 13 pages, 8 figures

We study the stochastic thermodynamics of cell growth and division using a theoretical framework based on branching processes with resetting. Cell division may be split into two sub-processes: branching, by which a given cell gives birth to an identical copy of itself, and resetting, by which some properties of the daughter cells (such as their size or age) are reset to new values following division. We derive the first and second laws of stochastic thermodynamics for this process, and identify separate contributions due to branching and resetting. We apply our framework to well-known models of cell size control, such as the sizer, the timer, and the adder. We show that the entropy production of resetting is negative and that of branching is positive for these models in the regime of exponential growth of the colony. This property suggests an analogy between our model for cell growth and division and heat engines, and the introduction of a thermodynamic efficiency, which quantifies the conversion of one form of entropy production to another.

Homochirality, i.e. the dominance across all living matter of one enantiomer over the other among chiral molecules, is thought to be a key step in the emergence of life. Building on ideas put forward by Frank and many others, we proposed recently one such mechanism in Laurentet al.(Laurent, 2021Proc. Natl Acad. Sci. USA118, e2012741118. (doi:10.1073/pnas.2012741118)) based on the properties of large out of equilibrium chemical networks. We showed that in such networks, a phase transition towards a homochiral state is likely to occur as the number of chiral species in the system becomes large or as the amount of free energy injected into the system increases. This paper aims at clarifying some important points in that scenario, not covered by our previous work. We first analyse the various conventions used to measure chirality, introduce the notion of chiral symmetry of a network and study its implications regarding the relative chiral signs adopted by different groups of molecules. We then propose a generalization of Frank’s model for large chemical networks, which we characterize completely using methods of random matrices. This analysis is extended to sparse networks, which shows that the emergence of homochirality is a robust transition.

International audience; The selection of a single molecular handedness, or homochirality across all living matter, is a mystery in the origin of life. Frank's seminal model showed in the fifties how chiral symmetry breaking can occur in non-equilibrium chemical networks. However, an important shortcoming in this classic model is that it considers a small number of species, while there is no reason for the prebiotic system, in which homochirality first appeared, to have had such a simple composition. Furthermore, this model does not provide information on what could have been the size of the molecules involved in this homochiral prebiotic system. Here, we show that large molecular systems are likely to undergo a phase transition towards a homochiral state, as a consequence of the fact that they contain a large number of chiral species. Using chemoinformatics tools, we quantify how abundant are chiral species in the chemical universe of all possible molecules of a given length. Then, we propose that Frank's model should be extended to include a large number of species, in order to possess the transition towards homochirality as confirmed by numerical simulations. Finally, using random matrix theory, we prove that large non-equilibrium reaction networks possess a generic and robust phase transition towards a homochiral state.

International audience; We obtain general inequalities constraining the difference between the average of an arbitrary function ofa phenotypic trait, which includes the fitness landscape of the trait itself, in the presence or in the absence ofnatural selection. These inequalities imply bounds on the strength of selection, which can be measured fromthe statistics of trait values and divisions along lineages. The upper bound is related to recent generalizations oflinear response relations in stochastic thermodynamics, and shares common features with Fisher’s fundamentaltheorem of natural selection, and with its generalization by Price, although they define different measures ofselection. The lower bound follows from recent improvements on Jensen’s inequality, and both bounds dependon the variability of the fitness landscape. We illustrate our results using numerical simulations of growing cellcolonies and with experimental data of time-lapse microscopy experiments of bacteria cell colonies.

Kelly's criterion is a betting strategy that maximizes the long-term growth rate, but which is known to be risky. Here, we find optimal betting strategies that gives the highest capital growth rate while keeping a certain low value of risky fluctuations. We then analyze the trade-off between the average and the fluctuations of the growth rate, in models of horse races, first for two horses then for an arbitrary number of horses, and for uncorrelated or correlated races. We find an analog of a phase transition with a coexistence between two optimal strategies, where one has risk and the other one does not. The above trade-off is also embodied in a general bound on the average growth rate, similar to thermodynamic uncertainty relations. We also prove mathematically the absence of other phase transitions between Kelly's point and the risk-free strategy.

Autocatalysis is an essential property for theories of abiogenesis and chemical evolution. However, the different formalisms proposed so far seemingly address different forms of autocatalysis and it remains unclear whether all of them have been captured. Furthermore, the lack of unified framework thus far prevents a systematic study of autocatalysis. Here, we derive general stoichiometric conditions for catalysis and autocatalysis in chemical reaction networks from basic principles in chemistry. This allows for a classification of minimal autocatalytic motifs, which includes all known autocatalytic systems and motifs that had not been reported previously. We further examine conditions for kinetic viability of such networks, which depends on the autocatalytic motifs they contain. Finally, we show how this framework extends the range of conceivable autocatalytic systems, by applying our stoichiometric and kinetic analysis to autocatalysis emerging from coupled compartments. The unified approach to autocatalysis presented in this work lays a foundation towards the building of a systems-level theory of chemical evolution.

The problem of the emergence and survival of self-replicating molecules in origin-of-life scenarios is plagued by the error catastrophe, which is usually escaped by considering effects of compartmentalization, as in the stochastic corrector model. By addressing the problem in a simple system composed of a self-replicating molecule (a replicase) and a parasite molecule that needs the replicase for copying itself, we show that transient (rather than permanent) compartmentalization is sufficient to the task. We also exhibit a regime in which the concentrations of the two kinds of molecules undergo sustained oscillations. Our model should be relevant not only for origin-of-life scenarios but also for describing directed evolution experiments, which increasingly rely on transient compartmentalization with pooling and natural selection.

Using a population dynamics inspired by an ensemble of growing cells, a set of fluctuation theorems linking observables measured at the lineage and population levels are derived. One of these relations implies specific inequalities comparing the population doubling time with the mean generation time at the lineage or population levels. While these inequalities have been derived before for age controlled models with negligible mother-daughter correlations, we show that they also hold for a broad class of size-controlled models. We discuss the implications of this result for the interpretation of a recent experiment in which the growth of bacteria strains has been probed at the single cell level., 13 pages, 5 figures

In this paper, we derive a number of inequalities which express power-efficiency trade-offs that hold generally for thermodynamic machines operating in non-equilibrium stationary states. One of these inequalities concerns the output power, which is bounded by a quadratic function of the thermodynamic efficiency multiplied by a factor. Different factors can be obtained according to the level of knowledge one has about the underlying dynamics of the machine, they can depend for instance on the covariance of the input flux, the dynamical activity, or the non-equilibrium conductance., 18 pages, 3 figures

We extend a recently introduced framework for transient compartmentalization of replicators with selection dynamics, by including the effect of mutations and noise in such systems. In the presence of mutations, functional replicators (ribozymes) are turned into non-functional ones (parasites). We evaluate the phase diagram of a system undergoing transient compartmentalization with selection. The system can exhibit either coexistence of ribozymes and parasites, or a pure parasite phase. If the mutation rate exceeds a certain level called the error threshold, the only stable phase is the pure parasite one. Transient compartmentalization with selection can relax this error treshold with respect to a bulk quasispecies case, and even allow ribozymes to coexist with faster growing parasites.In order to analyze the role of noise, we also introduce a model for the replication of a template by an enzyme. This model admits two regimes: a diffusion limited regime which generates a high noise, and a replication limited regime, which generates a low noise at the population level. Based on this model, we find that, since the ribozyme dynamics belongs to the replication limited regime, the effects of noise on the phase diagram of the system are mostly negligible. Our results underlines the importance of transient compartmentalization for prebiotic scenarios, and may have implications for directed evolution experiments.

Can prelife proceed without cell division? A recently proposed mechanism suggests that transient compartmentalization could have preceded cell division in prebiotic scenarios. Here, we study transient compartmentalization dynamics in the presence of mutations and noise in replication, as both can be detrimental the survival of compartments. Our study comprises situations where compartments contain uncoupled autocatalytic reactions feeding on a common resource, and systems based on RNA molecules copied by replicases, following a recent experimental study. Using the theory of branching processes, we show analytically that two regimes are possible. In the diffusion-limited regime, replication is asynchronous which leads to a large variability in the composition of compartments. In contrast, in a replication-limited regime, the growth is synchronous and thus the compositional variability is low. Typically, simple autocatalysts are in the former regime, while polymeric replicators can access the latter. For deterministic growth dynamics, we introduce mutations that turn functional replicators into parasites. We derive the phase boundary separating coexistence or parasite dominance as a function of relative growth, inoculation size and mutation rate. We show that transient compartmentalization allows coexistence beyond the classical error threshold, above which the parasite dominates. Our findings invite to revisit major prebiotic transitions, notably the transitions towards cooperation, complex polymers and cell division., 37 pages, 13 figures

Kinetic theory and thermodynamics of reaction networks are extended to the out-of-equilibrium dynamics of continuous-flow stirred tank reactors (CSTR) and serial transfers. On the basis of their stoichiometry matrix, the conservation laws and the cycles of the network are determined for both dynamics. It is shown that the CSTR and serial transfer dynamics are equivalent in the limit where the time interval between the transfers tends to zero proportionally to the ratio of the fractions of fresh to transferred solutions. These results are illustrated with finite cross-catalytic reaction network and an infinite reaction network describing mass exchange between polymers. Serial transfer dynamics is typically used in molecular evolution experiments in the context of research on the origins of life. The present study is shedding a new light on the role played by serial transfer parameters in these experiments., 11 pages, 7 figures

Transient compartments have been recently shown to be able to maintain functional replicators in the context of prebiotic studies. Motivated by this experiment, we show that a broad class of selection dynamics is able to achieve this goal. We identify two key parameters, the relative amplification of non-active replicators (parasites) and the size of compartments. Since the basic ingredients of our model are the competition between a host and its parasite, and the diversity generated by small size compartments, our results are relevant to various phage-bacteria or virus-host ecology problems., 11 pages, 10 figures

The mobility of membrane proteins controls many biological functions. The application of the model of Saffman and Delbr\"uck to the diffusion of membrane proteins does not account for all the experimental measurements. These discrepancies have triggered a lot of studies on the role of the mechanical factors in the mobility. After a short review of the Saffman and Delbr\"uck model and of some key experiments, we explore the various ways to incorporate the effects of the different mechanical factors. Our approach focuses on the coupling of the protein to the membrane, which is the central element in the modelling. We present a general, polaron-like model, its recent application to the mobility of a curvature sensitive protein, and its various extensions to other couplings that may be relevant in future experiments., Comment: To appear as a Chapter in "Physics of Biological Membranes", edited by Dr. Patricia Bassereau & Dr. Pierre Sens, Springer Nature Switzerland

Inferring the directionality of interactions between cellular processes is a major challenge in systems biology. Time-lagged correlations allow to discriminate between alternative models, but they still rely on assumed underlying interactions. Here, we use the transfer entropy (TE), an information-theoretic quantity that quantifies the directional influence between fluctuating variables in a model-free way. We present a theoretical approach to compute the transfer entropy, even when the noise has an extrinsic component or in the presence of feedback. We re-analyze the experimental data from Kiviet et al. (2014) where fluctuations in gene expression of metabolic enzymes and growth rate have been measured in single cells of E. coli. We confirm the formerly detected modes between growth and gene expression, while prescribing more stringent conditions on the structure of noise sources. We furthermore point out practical requirements in terms of length of time series and sampling time which must be satisfied in order to infer optimally transfer entropy from times series of fluctuations., 24 pages, 7 figures

In this paper, we introduce a real symmetric and positive semi-definite matrix, which we call the non-equilibrium conductance matrix, and which generalizes the Onsager response matrix for a system in a non-equilibrium stationary state. We then express the thermodynamic efficiency in terms of the coefficients of this matrix using a parametrization similar to the one used near equilibrium. This framework, then valid arbitrarily far from equilibrium allows to set bounds on the thermodynamic efficiency by a universal function depending only on the degree of coupling between input and output currents. It also leads to new general power-efficiency trade-offs valid for macroscopic machines that are compared to trade-offs previously obtained from uncertainty relations. We illustrate our results on an unicycle heat to heat converter and on a discrete model of molecular motor., Comment: 24 pages, 5 figures

We describe the kinetics and thermodynamics of copolymers undergoing recombination reactions, which are important for prebiotic chemistry. We use two approaches: the first one, based on chemical rate equations and the mass-action law describes the infinite size limit, while the second one, based on the chemical master equation, describes systems of finite size. We compare the predictions of both approaches for the relaxation of thermodynamic quantities towards equilibrium. We find that for some choice of initial conditions, the entropy of the sequence distribution can be lowered at the expense of increasing the entropy of the length distribution. We consider mainly energetically neutral reactions, except for one simple case of non-neutral reactions., Comment: 15 pages, 6 figures and 1 table

Depolymerizing cytoskeletal filaments are involved in cell division, cell motility, and other cellular functions. Understanding the dynamics of depolymerization is as important as understanding the dynamics of polymerization. We study nonequilibrium depolymerization of actin filaments using a simple two-state model. We show that the polymerization history influences the dynamics of depolymerization as well as the length fluctuations during depolymerization. We also simulate depolymerization under different experimentally feasible conditions. Under conditions of constant concentration, we show that the depolymerization happens in two regimes. Under the conditions of mass conservation, the depolymerization can have three regimes.

Uncovering mechanisms that control the dynamics of microtubules is fundamental for our understanding of multiple cellular processes such as chromosome separation and cell motility. Building on previous theoretical work on the dynamic instability of microtubules, we propose here a stochastic model that includes all relevant biochemical processes that affect the dynamics of microtubule plus-end, namely, the binding of GTP-bound monomers, unbinding of GTP- and GDP-bound monomers, and hydrolysis of GTP monomers. The inclusion of dissociation processes, present in our approach but absent from many previous studies, is essential to guarantee the thermodynamic consistency of the model. Our theoretical method allows us to compute all dynamic properties of microtubules explicitly. Using experimentally determined rates, it is found that the cap size is similar to 3.6 layers, an estimate that is compatible with several experimental observations. In the end, our model provides a comprehensive description of the dynamic instability of microtubules that includes not only the statistics of catastrophes but also the statistics of rescues.

Fluctuation theorems have become an important tool in single molecule biophysics to measure free energy differences from non-equilibrium experiments. When significant coarse-graining or noise affect the measurements, the determination of the free energies becomes challenging. In order to address this thermodynamic inference problem, we propose improved estimators of free energy differences based on fluctuation theorems, which we test on a number of examples. The effect of the noise can be described by an effective temperature, which only depends on the signal to noise ratio, when the work is Gaussian distributed and uncorrelated with the error made on the work. The notion of effective temperature appears less useful for non-Gaussian work distributions or when the error is correlated with the work, but nevertheless, as we show, improved estimators can still be constructed for such cases. As an example of non-trivial correlations between the error and the work, we also consider measurements with delay, as described by linear Langevin equations., 19 pages, 8 figures. Submitted to Phys. Rev. E

We describe the oligosaccharides-exchange dynamics performed by so-called D-enzymes on polysaccharides. To mimic physiological conditions, we treat this process as an open chemical network by assuming some of the polymer concentrations fixed (chemostatting). We show that three different long-time behaviors may ensue: equilibrium states, nonequilibrium steady states, and continuous growth states. We dynamically and thermodynamically characterize these states and emphasize the crucial role of conservation laws in identifying the chemostatting conditions inducing them., Comment: 12 pages, 9 figures and 1 table

We derive a set of isometric fluctuation relations, which constrain the order parameter fluctuations in finite-size systems at equilibrium and in the presence of a broken symmetry. These relations are exact and should apply generally to many condensed-matter physics systems. Here, we establish these relations for magnetic systems and nematic liquid crystals in a symmetry-breaking external field, and we illustrate them on the Curie-Weiss and the XY models. Our relations also have implications for spontaneous symmetry breaking, which are discussed., SCOPUS: ar.j, info:eu-repo/semantics/published

We study the time-dependent height fluctuations of an active membrane containing energy-dissipating pumps that drive the membrane out of equilibrium. Unlike previous investigations based on models that neglect either curvature couplings or random fluctuations in pump activities, our formulation explores two new models that take both of these effects into account. In the first model, the magnitude of the nonequilibrium forces generated by the pumps is allowed to fluctuate temporally. In the second model, the pumps are allowed to switch between "on" and "off" states. We compute the mean squared displacement of a membrane point for both models, and show that they exhibit distinct dynamical behaviors from previous models, and in particular, a superdiffusive regime specifically arising from the shot noise., 7 pages, 4 figures

We study the stochastic thermodynamics of an overdamped harmonic chain, which can be viewed equivalently as a 1D Rouse chain or as an approximate model of single file diffusion. We discuss mainly two levels of description of this system: the Markovian level for which the trajectories of all the particles of the chain are known and the non-Markovian level in which only the motion of a tagged particle is available. For each case, we analyze the energy dissipation and its dependence on initial conditions. Surprisingly, we find that the average coarse-grained entropy production rate can become transiently negative when an oscillating force is applied to the tagged particle. This occurs due to memory effects as shown in a framework based on path integrals or on a generalized Langevin equation., 17 pages, 7 figures, REVTeX, minor revisions

Using the framework of stochastic thermodynamics, we present an experimental study of a doublet of magnetic colloidal particles which is manipulated by a time-dependent magnetic field. Due to hydrodynamic interactions, each bead experiences a state-dependent friction, which we characterize using a hydrodynamic model. In this work, we compare two estimates of the dissipation in this system: the first one is energy based since it relies on the measured interaction potential, while the second one is information based since it uses only the information content of the trajectories. While the latter only offers a lower bound of the former, we find it to be simple to implement and of general applicability to more complex systems., Main text: 5 pages, 4 figures. Supplementary material: 5 pages, 5 figures

We describe an experimental and theoretical study of the effect of optical absorption on the photonic Hall effect in a passive matrix containing magnetoactive scatterers. We find that for the case of absorbing scatterers, the magnetotransverse light current changes sign and increases with increasing absorption. Good agreement is obtained with numerical calculations. For the case of an absorbing matrix, no effect was observed.

We show how to extract from a sufficiently long time series of stationary fluctuations of chemical reactions an estimate of the entropy production. This method, which is based on recent work on fluctuation theorems, is direct, non-invasive, does not require any knowledge about the underlying dynamics, and is applicable even when only partial information is available. We apply it to simple stochastic models of chemical reactions involving a finite number of states, and for this case, we study how the estimate of dissipation is affected by the degree of coarse-graining present in the input data., Accepted in Journal of Chemical Physics

We analyze thermodynamic bounds on equilibrium fluctuations of an order parameter, which are analogous to relations, which have been derived recently in the context of non-equilibrium fluctuations of currents. We discuss the case of {\it global} fluctuations when the order parameter is measured in the full system of interest, and {\it local} fluctuations, when the order parameter is evaluated only in a sub-part of the system. Using isometric fluctuation theorems, we derive thermodynamic bounds on the fluctuations of the order parameter in both cases. These bounds could be used to infer the value of symmetry breaking field or the relative size of the observed sub-system to the full system from {\it local} fluctuations., Comment: 8 pages, 6 figures, in press for Europhys. Lett

We develop a model to describe the force generated by the polymerization of an array of parallel biofilaments. The filaments are assumed to be coupled only through mechanical contact with a movable barrier. We calculate the filament density distribution and the force-velocity relation with a mean-field approach combined with simulations. We identify two regimes: a non-condensed regime at low force in which filaments are spread out spatially, and a condensed regime at high force in which filaments accumulate near the barrier. We confirm a result previously known from other related studies, namely that the stall force is equal to N times the stall force of a single filament. In the model studied here, the approach to stalling is very slow, and the velocity is practically zero at forces significantly lower than the stall force., Comment: 21 pages, 6 figures: Combined figures, fixed typos, added extra material, altered symbolism to avoid confusion. Accepted by New Journal of Physics

We present a general framework for systems which are prepared in a non-stationary non-equilibrium state in the absence of any perturbation, and which are then further driven through the application of a time-dependent perturbation. We distinguish two different situations depending on the way the non-equilibrium state is prepared, either it is created by some driving; or it results from a relaxation following some initial non-stationary conditions. Our approach is based on a recent generalization of the Hatano-Sasa relation for non-stationary probability distributions. We also investigate whether a form of second law holds for separate parts of the entropy production, in a way similar to the work of M. Esposito et al., Phys. Rev. Lett., 104:090601 (2010), but for a non-stationary reference process instead of a stationary one. We find that although the special structure of the theorems derived in this reference is not recovered in the general case, detailed fluctuation theorems still hold separately for parts of the entropy production in this case. These detailed fluctuation theorems lead to interesting generalizations of the second-law of thermodynamics off equilibrium., Comment: 36 pages, 4 figures

We discuss the consequences of a variant of the Hatano-Sasa relation in which a non-stationary distribution is used in place of the usual stationary one. We first show that this non-stationary distribution is related to a difference of traffic between the direct and dual dynamics. With this formalism, we extend the definition of the adiabatic and non-adiabatic entropies introduced by M. Esposito and C. Van den Broeck in Phys. Rev. Lett. 104, 090601 (2010) for the stationary case. We also obtain interesting second-law like inequalities for transitions between non-stationary states., Comment: 4 pages, 2 figures

Fluctuation relations are a set of remarkable relations obeyed by a large class of systems and arbitrarily far from equilibrium. It is interesting to discuss the implications of these relations for molecular motors, which are chemically driven enzymes. These enzymes operate stochastically at the molecular level and for these reasons undergo large thermal fluctuations. Using simple ratchet models of molecular motors, the various forms of fluctuation relations can be illustrated in a simple way. In the linear regime, finite time fluctuation relations imply specific modified fluctuation‐dissipation relations.

This review is focused on the application of specific fluctuation relations, such as the Gallavotti-Cohen relation, to ratchet models of a molecular motor. A special emphasis is placed on two-states models such as the flashing ratchet model. We derive the Gallavotti-Cohen fluctuation relation for these models and we discuss some of its implications., 27 pages, 11 figures

We present a theoretical framework to understand a modified fluctuation-dissipation theorem valid for systems close to non-equilibrium steady-states and obeying markovian dynamics. We discuss the interpretation of this result in terms of trajectory entropy excess. The framework is illustrated on a simple pedagogical example of a molecular motor. We also derive in this context generalized Green-Kubo relations similar to the ones derived recently by Seifert., Phys. Rev. Lett., 104, 138101 (2010) for more general networks of biomolecular states., Comment: 6 pages, 2 figures, submitted in EPL

The behavior of a conductive membrane in a static (DC) electric field is investigated theoretically. An effective zero-thickness model is constructed based on a Robin-type boundary condition for the electric potential at the membrane, originally developed for electrochemical systems. Within such a framework, corrections to the elastic moduli of the membrane are obtained, which arise from charge accumulation in the Debye layers due to capacitive effects and electric currents through the membrane and can lead to an undulation instability of the membrane. The fluid flow surrounding the membrane is also calculated, which clarifies issues regarding these flows sharing many similarities with flows produced by induced charge electro-osmosis (ICEO). Non-equilibrium steady states of the membrane and of the fluid can be effectively described by this method. It is both simpler, due to the zero thickness approximation which is widely used in the literature on fluid membranes, and more general than previous approaches. The predictions of this model are compared to recent experiments on supported membranes in an electric field., 14 pages, 5 figures

We study the stochastic dynamics of growth and shrinkage of single actin filaments taking into account insertion, removal, and ATP hydrolysis of subunits either according to the vectorial mechanism or to the random mechanism. In a previous work, we developed a model for a single actin or microtubule filament where hydrolysis occurred according to the vectorial mechanism: the filament could grow only from one end, and was in contact with a reservoir of monomers. Here we extend this approach in several ways, by including the dynamics of both ends and by comparing two possible mechanisms of ATP hydrolysis. Our emphasis is mainly on two possible limiting models for the mechanism of hydrolysis within a single filament, namely the vectorial or the random model. We propose a set of experiments to test the nature of the precise mechanism of hydrolysis within actin filaments., To appear in Biophysical Journal (2010)

We study the thermal expansion of chains formed by self-assembly of magnetic colloidal particles in a magnetic field. Using video-microscopy, complete positional data of all the particles of the chains is obtained. By changing the ionic strength of the solution and the applied magnetic field, the interaction potential can be tuned. We analyze the thermal expansion of the chain using a simple model of a one dimensional anharmonic crystal of finite size., 5 pages and 3 figures

International audience; In 1994 Leal Calderon et al. (Phys. Rev. Lett. 72, 2959 (1994)) introduced the magnetic chaining technique to directly probe the force-distance profile between colloidal particles. In this paper, we revisit this approach in two ways. First, we describe a new experimental design which allows us to utilize sample volumes as low as a few microliters, involving femtomoles of surface active macromolecules. Secondly, we extensively describe the characterization and preparation of the magnetic colloids, and we give a quantitative evaluation of performance and resolution of the technique in terms of force and interparticle separation.

Molecular motors convert chemical energy derived from the hydrolysis of ATP into mechanical energy. A well-studied model of a molecular motor is the flashing ratchet model. We show that this model exhibits a fluctuation relation known as the Gallavotti-Cohen symmetry. Our study highlights the fact that the symmetry is present only if the chemical and mechanical degrees of freedom are both included in the description., Comment: 5 pages, 4 figures

We study the stochastic dynamics of growth and shrinkage of single actin filaments or microtubules taking into account insertion, removal, and ATP/GTP hydrolysis of subunits. The resulting phase diagram contains three different phases: a rapidly growing phase, an intermediate phase and a bound phase. We analyze all these phases, with an emphasis on the bound phase. We also discuss how hydrolysis affects force-velocity curves. The bound phase shows features of dynamic instability, which we characterize in terms of the time needed for the ATP/GTP cap to disappear as well as the time needed for the filament to reach a length of zero, i.e., (to collapse) for the first time. We obtain exact expressions for all these quantities, which we test using Monte Carlo simulations., Comment: 34 pages

We present measurements of the fluctuation spectrum of giant vesicles containing bacteriorhodopsin (BR) pumps using video-microscopy. When the pumps are activated, we observe a significant increase of the fluctuations in the low wavevector region, which we interpret as due to a lowering of the effective tension of the membrane., 4 pages, 2 figures

We discuss the electrostatic contribution to the elastic moduli of a cell or artificial membrane placed in an electrolyte and driven by a DC electric field. The field drives ion currents across the membrane, through specific channels, pumps or natural pores. In steady state, charges accumulate in the Debye layers close to the membrane, modifying the membrane elastic moduli. We first study a model of a membrane of zero thickness, later generalizing this treatment to allow for a finite thickness and finite dielectric constant. Our results clarify and extend the results presented in [D. Lacoste, M. Cosentino Lagomarsino, and J. F. Joanny, Europhys. Lett., {\bf 77}, 18006 (2007)], by providing a physical explanation for a destabilizing term proportional to $\kps^3$ in the fluctuation spectrum, which we relate to a nonlinear ($E^2$) electro-kinetic effect called induced-charge electro-osmosis (ICEO). Recent studies of ICEO have focused on electrodes and polarizable particles, where an applied bulk field is perturbed by capacitive charging of the double layer and drives flow along the field axis toward surface protrusions; in contrast, we predict "reverse" ICEO flows around driven membranes, due to curvature-induced tangential fields within a non-equilibrium double layer, which hydrodynamically enhance protrusions. We also consider the effect of incorporating the dynamics of a spatially dependent concentration field for the ion channels., Comment: 22 pages, 10 figures. Under review for EPJE