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Jean-Baptiste Fournier

Publications and source records attributed to Jean-Baptiste Fournier.

At least 37 records · Page 2Linked to original sources

Interplay of packing and flip-flop in local bilayer deformation. How phosphatidylglycerol could rescue mitochondrial function in a cardiolipin-deficient yeast mutant

In a previous work, we have shown that a spatially localized transmembrane pH gradient, produced by acid micro-injection near the external side of cardiolipin-containing giant unilamellar vesicles, leads to the formation of tubules that retract after the dissipation of this gradient. These tubules have morphologies similar to mitochondrial cristae. The tubulation effect is due to direct phospholipid packing modification in the outer leaflet that is promoted by protonation of cardiolipin headgroups. Here we compare the case of cardiolipin-containing giant unilamellar vesicles with that of phosphatidylglycerol-containing giant unilamellar vesicles. Local acidification also promotes formation of tubules in the latter. However, compared to cardiolipin-containing giant unilamellar vesicles the tubules are longer, exhibit a visible pearling and have a much longer lifetime after acid micro-injection is stopped. We attribute these differences to an additional mechanism that increases monolayer surface imbalance, namely inward PG flip-flop promoted by the local transmembrane pH-gradient. Simulations using a fully non-linear membrane model as well as geometrical calculations are in agreement with this hypothesis. Interestingly, among yeast mutants deficient in cardiolipin biosynthesis, only the crd1-null mutant, which accumulates phosphatidylglycerol, displays significant mitochondrial activity. Our work provides a possible explanation of such a property and further emphasizes the salient role of specific lipids in mitochondrial function.

cond-mat.soft↗

On the dynamics of the interaction between triggered active inclusions

In a one-dimensional elastic medium with finite correlation length and purely relaxational dynamics, we calculate the time dependence of the elastic force F(t) exchanged between two active inclusions that trigger an elastic deformation at t=0. We consider (i) linear inclusions coupling to the field with a finite force, and (ii) non-linear inclusions imposing a finite deformation. In the non-linear case, the force exhibits a transient maximum much larger than the equilibrium force, diverging as ~L^(-2) at separations L shorter than the field's correlation length. Both the mean-field and the Casimir component of the interaction are calculated. We also discuss the typical appearance time and equilibration time of the force, comparing the linear and the non-linear cases. The existence of a high transient force in the non-linear case should be a generic feature of elastically-mediated interactions.

cond-mat.stat-mech↗

Membrane properties revealed by spatiotemporal response to a local inhomogeneity

We study theoretically the spatiotemporal response of a lipid membrane submitted to a local chemical change of its environment, taking into account the time-dependent profile of the reagent concentration due to diffusion in the solution above the membrane. We show that the effect of the evolution of the reagent concentration profile becomes negligible after some time. It then becomes possible to extract interesting properties of the membrane response to the chemical modification. We find that a local density asymmetry between the two monolayers relaxes by spreading diffusively in the whole membrane. This behavior is driven by intermonolayer friction. Moreover, we show how the ratio of the spontaneous curvature change to the equilibrium density change induced by the chemical modification can be extracted from the dynamics of the local membrane deformation. Such information cannot be obtained by analyzing the equilibrium vesicle shapes that exist in different membrane environments in light of the area-difference elasticity model.

cond-mat.soft↗

Bilayer elasticity at the nanoscale: the need for new terms

Continuum elastic models that account for membrane thickness variations are especially useful in the description of nanoscale deformations due to the presence of membrane proteins with hydrophobic mismatch. We show that terms involving the gradient and the Laplacian of the area per lipid are significant and must be retained in the effective Hamiltonian of the membrane. We reanalyze recent numerical data, as well as experimental data on gramicidin channels, in light of our model. This analysis yields consistent results for the term stemming from the gradient of the area per molecule. The order of magnitude we find for the associated amplitude, namely 13-60 mN/m, is in good agreement with the 25 mN/m contribution of the interfacial tension between water and the hydrophobic part of the membrane. The presence of this term explains a systematic variation in previously published numerical data.

cond-mat.soft↗

Lipid membrane deformation in response to a local pH modification: theory and experiments

We study the deformation of a lipid membrane in response to a local pH modification. Experimentally, a basic solution is microinjected close to a giant unilamellar vesicle. A local deformation appears in the zone of the membrane that is closest to the micropipette, and relaxes when the injection is stopped. A theoretical description of this phenomenon is provided. It takes fully into account the spatiotemporal evolution of the concentration of hydroxide ions during and after the microinjection, as well as the linear dynamics of the membrane. This description applies to a local injection of any substance that reacts reversibly with the membrane lipids. We compare experimental data obtained in the domain of small deformations to the results of our linear description, and we obtain a good agreement between theory and experiments. In addition, we present direct experimental observations of the pH profile on the membrane during and after the microinjection, using pH-sensitive fluorescent lipids.

cond-mat.soft↗

Universal amplitudes of the Casimir-like interactions between four types of rods in fluid membranes

The fluctuation-induced, Casimir-like interaction between two parallel rods of length L adsorbed on a fluid membrane is calculated analytically at short separations d<<L. The rods are modeled as constraints imposed on the membrane curvature along a straight line. This allows to define four types of rods, according to whether the membrane can twist along the rod and/or curve across it. For stiff constraints, all the interaction potentials between the different types of rods are attractive and proportional to L/d. Two of the four types of rods are then equivalent, which yields six universal Casimir amplitudes. Repulsion can occur between different rods for soft constraints. Numerical results obtained for all ranges of d/L show that the attraction potential reaches kT for d/L\simeq0.2. At separations smaller than d_c \approx L(L/l_p)^(1/3), where l_p is the rod persistence length, two rods with fixed ends will bend toward each other and finally come into contact because of the Casimir interaction.

cond-mat.soft↗

Dynamical membrane curvature instability controlled by intermonolayer friction

We study a dynamical curvature instability caused by a local chemical modification of a phospholipid membrane. In our experiments, a basic solution is microinjected close to a giant unilamellar vesicle, which induces a local chemical modification of some lipids in the external monolayer of the membrane. This modification causes a local deformation of the vesicle, which then relaxes. We present a theoretical description of this instability, taking into account both the change of the equilibrium lipid density and the change of the spontaneous membrane curvature induced by the chemical modification. We show that these two types of changes of the membrane properties yield different dynamics. In contrast, it is impossible to distinguish them when studying the equilibrium shape of a vesicle subjected to a global modification. In our model, the longest relaxation timescale is related to the intermonolayer friction, which plays an important part when there is a change in the equilibrium density in one monolayer. We compare our experimental results to the predictions of our model by fitting the measured time evolution of the deformation height to the solution of our dynamical equations. We obtain good agreement between theory and experiments. Our fits enable us to estimate the intermonolayer friction coefficient, yielding values that are consistent with previous measurements.

cond-mat.soft↗

Forces exerted by a correlated fluid on embedded inclusions

We investigate the forces exerted on embedded inclusions by a fluid medium with long-range correlations, described by an effective scalar field theory. Such forces are the basis for the medium-mediated Casimir-like force. To study these forces beyond thermal average, it is necessary to define them in each microstate of the medium. Two different definitions of these forces are currently used in the literature. We study the assumptions underlying them. We show that only the definition that uses the stress tensor of the medium gives the sought-after force exerted by the medium on an embedded inclusion. If a second inclusion is embedded in the medium, the thermal average of this force gives the usual Casimir-like force between the two inclusions. The other definition can be used in the different physical case of an object that interacts with the medium without being embedded in it. We show in a simple example that the two definitions yield different results for the variance of the Casimir-like force.

cond-mat.soft↗

Membrane stress tensor in the presence of lipid density and composition inhomogeneities

We derive the expression of the stress tensor for one and two-component lipid membranes with density and composition inhomogeneities. We first express the membrane stress tensor as a function of the free-energy density by means of the principle of virtual work. We then apply this general result to a monolayer model which is shown to be a local version of the area-difference elasticity (ADE) model. The resulting stress tensor expression generalizes the one associated with the Helfrich model, and can be specialized to obtain the one associated with the ADE model. Our stress tensor directly gives the force exchanged through a boundary in a monolayer with density and composition inhomogeneities. Besides, it yields the force density, which is also directly obtained in covariant formalism. We apply our results to study the forces induced in a membrane by a local perturbation.

cond-mat.soft↗

Fluctuations of the Casimir-like force between two membrane inclusions

Although Casimir forces are inseparable from their fluctuations, little is known about these fluctuations in soft matter systems. We use the membrane stress tensor to study the fluctuations of the membrane-mediated Casimir-like force. This method enables us to recover the Casimir force between two inclusions and to calculate its variance. We show that the Casimir force is dominated by its fluctuations. Furthermore, when the distance d between the inclusions is decreased from infinity, the variance of the Casimir force decreases as -1/d^2. This distance dependence shares a common physical origin with the Casimir force itself.

cond-mat.soft↗

On the stress and torque tensors in fluid membranes

We derive the membrane elastic stress and torque tensors using the standard Helfrich model and a direct variational method in which the edges of a membrane are infinitesimally translated and rotated. We give simple expressions of the stress and torque tensors both in the local tangent frame and in projection onto a fixed frame. We recover and extend the results of Capovilla and Guven [J. Phys. A, 2002, \textbf{35}, 6233], which were obtained using covariant geometry and Noether's theorem: we show that the Gaussian rigidity contributes to the torque tensor and we include the effect of a surface potential in the stress tensor. Many interesting situations may be investigated directly using force and torque balances instead of full energy minimization. As examples, we consider the force exerted at the end of a membrane tubule, membrane adhesion and domain contact conditions.

cond-mat.soft↗

Effective creases and contact angles between membrane domains with high spontaneous curvature

We show that the short-scale elastic distortions that are excited in the vicinity of the joint between different lipidic membrane domains (at a scale of \~10 nm) may produce a "crease" from the point of view of the standard elastic description of membranes, i.e., an effective discontinuity in the membrane slope at the level of Helfrich's theory. This "discontinuity" may be accounted for by introducing a line tension with an effective angular dependence. We show that domains bearing strong spontaneous curvatures, such as biological rafts, should exhibit creases with a finite contact-angle, almost prescribed, corresponding to a steep extremum of the line energy. Finite contact-angles might also occur in symmetric membranes from the recruitment of impurities at the boundary.

cond-mat.soft↗

A Gaussian model for the membrane of red blood cells with cytoskeletal defects

We study a Gaussian model of the membrane of red blood cells: a "phantom" triangular network of springs attached at its vertices to a fluid bilayer with curvature elasticity and tension. We calculate its fluctuation spectrum and we discuss the different regimes and non-monotonic features, including the precise crossover at the mesh size between the already known limits with two different tensions and the renormalisation of the bending rigidity at low wavevectors. We also show that the non-diagonal correlations reveal, in "dark field", the cytoskeletal defects. As a first step toward a non-invasive defect spectroscopy, the specific case of lacking bonds is studied numerically and analytically.

cond-mat.soft↗

Modeling planar degenerate wetting and anchoring in nematic liquid crystals

We propose a simple surface potential favoring the planar degenerate anchoring of nematic liquid crystals, i.e., the tendency of the molecules to align parallel to one another along any direction parallel to the surface. We show that, at lowest order in the tensorial Landau-de Gennes order-parameter, fourth-order terms must be included. We analyze the anchoring and wetting properties of this surface potential. In the nematic phase, we find the desired degenerate planar anchoring, with positive scalar order-parameter and some surface biaxiality. In the isotropic phase, we find, in agreement with experiments, that the wetting layer may exhibit a uniaxial ordering with negative scalar order-parameter. For large enough anchoring strength, this negative ordering transits towards the planar degenerate state.

cond-mat.soft↗

General up to next-nearest neighbour elasticity of triangular lattices in three dimensions

We establish the most general form of the discrete elasticity of a 2D triangular lattice embedded in three dimensions, taking into account up to next-nearest neighbour interactions. Besides crystalline system, this is relevant to biological physics (e.g., red blood cell cytoskeleton) and soft matter (e.g., percolating gels, etc.). In order to correctly impose the rotational invariance of the bulk terms, it turns out to be necessary to take into account explicitly the elasticity associated with the vertices located at the edges of the lattice. We find that some terms that were suspected in the litterature to violate rotational symmetry are in fact admissible

cond-mat.soft↗

Fluctuation spectrum of fluid membranes coupled to an elastic meshwork: jump of the effective surface tension at the mesh size

We identify a class of composite membranes: fluid bilayers coupled to an elastic meshwork, that are such that the meshwork's energy is a function $F_\mathrm{el}[A_ξ]$ \textit{not} of the real microscopic membrane area $A$, but of a \textit{smoothed} membrane's area $A_ξ$, which corresponds to the area of the membrane coarse-grained at the mesh size $ξ$. We show that the meshwork modifies the membrane tension $σ$ both below and above the scale $ξ$, inducing a tension-jump $Δσ=dF_\mathrm{el}/dA_ξ$. The predictions of our model account for the fluctuation spectrum of red blood cells membranes coupled to their cytoskeleton. Our results indicate that the cytoskeleton might be under extensional stress, which would provide a means to regulate available membrane area. We also predict an observable tension jump for membranes decorated with polymer "brushes".

cond-mat.soft↗

Effective interactions between inclusions in complex fluids driven out of equilibrium

The concept of fluctuation-induced effective interactions is extended to systems driven out of equilibrium. We compute the forces experienced by macroscopic objects immersed in a soft material driven by external shaking sources. We show that, in contrast with equilibrium Casimir forces induced by thermal fluctuations, their sign, range and amplitude depends on specifics of the shaking and can thus be tuned. We also comment upon the dispersion of these shaking-induced forces, and discuss their potential application to phase ordering in soft-materials.

cond-mat.soft↗

Elastic interaction between "hard'' or "soft" pointwise inclusions on biological membranes

We calculate the induced elastic-interaction between pointwise membrane inclusions that locally interact up to quadratic order with the membrane curvature tensor. For isotropic inclusions, we recover the usual interaction proportional to the inverse fourth power of the separation, however with a prefactor showing a non-trivial dependence on the rigidity $Γ$ of the quadratic potential. In the large $Γ$ limit, corresponding to ``hard'' inclusions, we recover the standard prefactor first obtained by Goulian et al. [Europhys. Lett. \textbf{22}, 145 (1993)]. In the small $Γ$ limit, corresponding to "soft" inclusions, we recover the recent result of Marchenko and Misbah [Eur. Phys. J. E \textbf{8}, 477 (2002)]. This shows that the latter result bears no fundamental discrepancy with previous works, but simply corresponds to the limit of soft inclusions. We discuss how the same inclusion can be depicted as hard or soft according to the degree of coarse-graining of the pointwise description. Finally, we argue that conical transmembrane proteins should be fundamentally considered as hard inclusions.

cond-mat.soft↗