arXiv subjects

Matthias F. Schneider

Publications and source records attributed to Matthias F. Schneider.

At least 19 recordsLinked to original sources

arXiv2026

Generation of two-dimensional pulses in lipid monolayers by rapid photoswitching

We study pressure pulse generation and propagation in lipid monolayers by an experimental approach employing rapid photoisomerization of photoswitchable lipids (azoPC). This allows us to generate longitudinal surface pressure pulses by optical flash excitation in both free and constrained layer geometries. We compare the observed pulse shapes with a theoretical approach based on a nonlinear fractional wave equation for a surface displacement field, where a fractional time derivative term captures the hydrodynamics of the monolayer subphase. We explore channel geometries of different lengths and widths and find quantitative agreement between theory and experiment regarding pulse speed and pulse shapes. For narrow channels, we employ a one-dimensional version of the fractional wave equation to study pulse propagation without any fit parameters by using the pressure signal at a close pressure sensor as boundary condition to predict the pressure signal at a second far sensor. A full two-dimensional description can capture all effects arising from the channel geometry for wider channels using one common set of fit parameters for the pulse excitation that can be applied to all geometries. The nonlinearity in the fractional wave equation plays no role in explaining the observed pulse shapes because pulse amplitudes generated by azoPC photoswitching remain very small.

cond-mat.soft↗

arXiv2022

Surface Deformation During an Action Potential in Pearled Cells

Electric pulses in biological cells (action potentials) have been reported to be accompanied by a propagating cell-surface deformation with a nano-scale amplitude. Typically, this cell surface is covered by external layers of polymer material (extracellular matrix, cell wall material etc.). It was recently demonstrated in excitable plant cells (Chara Braunii) that the rigid external layer (cell wall) hinders the underlying deformation. When the cell membrane was separated from the cell wall by osmosis, a mechanical deformation, in the micrometer range, was observed upon excitation of the cell. The underlying mechanism of this mechanical pulse has up to date remained elusive. Herein we report that Chara cells can undergo a pearling instability, and when the pearled fragments were excited even larger and more regular cell shape changes were observed (about 10 to 100 um in amplitude). These transient cellular deformations were captured by a curvature model that is based on three parameters: surface tension, bending rigidity and pressure difference across the surface. In this paper these parameters are extracted by curve-fitting to the experimental cellular shapes at rest and during excitation. This is a necessary step to identify the mechanical parameters that change during an action potential.

physics.bio-ph↗

arXiv2022

Unitary response of solvatochromic dye to pulse excitation in lipid and cell membranes

The existence of acoustic pulse propagation in lipid monolayers at the air-water interface is well known. These pulses are controlled by the thermodynamic state of the lipid membrane. Nevertheless, the role of acoustic pulses for intra- and intercellular communication are still a matter of debate. Herein, we used the dye di 4- -ANEPPDHQ, which is known to be sensitive to the physical state and transmembrane potential of membranes, in order to gain insight into compression waves in lipid-based membrane interfaces. The dye was incorporated into lipid monolayers made of phosphatidylserine or phosphatidylcholine at the air-water-interface. A significant blue shift of the emission spectrum was detected when the state of the monolayer was changed from the liquid expanded (LE) to the liquid condensed (LC) phase. This transition-sensitivity of di-4-ANEPPDHQ was generalized in experiments with the bulk solvent dimethyl sulfoxide (DMSO). Upon crystallization of solvent, the emission spectrum also underwent a blue shift. During compression pulses in lipid monolayers, a significant fluorescence response was only observed when in the main transition regime. The optical signature of these waves, in terms of sign and magnitude, was identical to the response of di-4-ANEPPDHQ during action potentials in neurons and excitable plant cells. These findings corroborated the suggestion that action potentials are nonlinear state changes that propagate in the cell membrane.

physics.bio-ph↗

arXiv2022

Sharp, localized phase transitions in single neuronal cells

The origin of nonlinear responses in cells has been suggested to be crucial for various cell functions including the propagation of the nervous impulse. In physics nonlinear behavior often originates from phase transitions. Evidence for such transitions on the single cell level, however, has so far not been provided leaving the field unattended by the biological community. Here we demonstrate that single cells of a human neuronal cell line, display all optical features of a sharp, highly nonlinear phase transition within their membrane. The transition is reversible and does not origin from protein denaturation. Triggered by temperature and modified by pH here, a thermodynamic approach, strongly suggests, that similar nonlinear state changes can be induced by other variables such as calcium or mechanical stress. At least in lipid membranes such state changes are accompanied by significant changes in permeability, enzyme activity, elastic and electrical properties.

physics.bio-ph↗

arXiv2020

The living state: how cellular excitability is controlled by the thermodynamic state of the membrane

The thermodynamic (TD) properties of biological membranes play a central role for living systems. It has been suggested, for instance, that nonlinear pulses such as action potentials (APs) can only exist if the membrane state is in vicinity of a TD transition. Herein, two membrane properties - excitability and AP velocity - are investigated for a broad spectrum of conditions in living systems (temperature (T), 3D-pressure (p) and pH dependence). Based on these data we predict parameter ranges in which a transition of the membrane is located (15-35°C below growth temperature; 1-3 pH units below pH 7; at ~800 atm) and propose the corresponding phase diagrams. The latter explain: (i) changes of AP velocity with T, p and pH. (ii) The existence and origin of two qualitatively different forms of loss of nonlinear excitability ("nerve blockage", anesthesia). (iii) The type and quantity of parameter changes that trigger APs. Finally, a quantitative comparison between the TD behaviour of 2D-lipid model membranes with living systems is attempted. The typical shifts in transition temperature with pH and p of model membranes agree closely with values obtained from cell physiological measurements (excitability and propagation velocity). Taken together, these results strongly suggest that it is not specific molecules that control the excitability of living systems but rather the TD properties of the quasi 2D-membrane interface. The approach as proposed herein can be extended to other quantities (surface potential, calcium concentration, etc.) and makes clearly falsifiable predictions, for example, that a transition exists within the specified parameter ranges in excitable cells.

physics.bio-ph↗

arXiv2018

On measuring the acoustic state changes in lipid membranes using fluorescent probes

Ultrasound is increasingly being used to modulate the properties of biological membranes for applications in drug delivery and neuromodulation. While various studies have investigated the mechanical aspect of the interaction such as acoustic absorption and membrane deformation, it is not clear how these effects transduce into biological functions, for example, changes in the permeability or the enzymatic activity of the membrane. A critical aspect of the activity of an enzyme is the thermal fluctuations of its solvation or hydration shell. Thermal fluctuations are also known to be directly related to membrane permeability. Here solvation shell changes of lipid membranes subject to an acoustic impulse were investigated using a fluorescence probe, Laurdan. Laurdan was embedded in multi-lamellar lipid vesicles in water, which were exposed to broadband pressure impulses of the order of 1MPa peak amplitude and 10μs pulse duration. An instrument was developed to monitor changes in the emission spectrum of the dye at two wavelengths with sub-microsecond temporal resolution. The experiments show that changes in the emission spectrum, and hence the fluctuations of the solvation shell, are related to the changes in the thermodynamic state of the membrane and correlated with the compression and rarefaction of the incident sound wave. The results suggest that acoustic fields affect the state of a lipid membrane and therefore can potentially modulate the kinetics of channels and proteins embedded in the membrane.

cond-mat.soft↗

arXiv2018

It sounds like an action potential: unification of electrical, chemical and mechanical aspects of acoustic pulses in lipids

In an ongoing debate on the physical nature of the action potential, one group adheres to the electrical model of Hodgkin and Huxley, while the other describes the action potential as a non-linear acoustic pulse propagating within an interface near a transition. However, despite remarkable similarities, acoustics remains a non-intuitive mechanism for action potentials for the following reason. While acoustic pulses are typically associated with the propagation of density, pressure and temperature variation, action potentials are most commonly measured electrically. Here, we show that this discrepancy is lifted upon considering the electrical and chemical aspects of the interface, in addition to its mechanical properties. Specifically, we demonstrate how electrical and pH aspects of acoustic pulses emerge from an idealized description of the lipid interface, which is based on classical physical principles and contains no fit parameters. The pulses that emerge from the model show striking similarities to action potentials not only in qualitative shape and scales (time, velocity and voltage), but also demonstrate saturation of amplitude and annihilation upon collision.

physics.bio-ph↗

arXiv2018

Similarities between action potentials and acoustic pulses in a van der Waals fluid

An action potential is typically described as a purely electrical change that propagates along the membrane of excitable cells. However, recent experiments have demonstrated that non-linear acoustic pulses that propagate along lipid interfaces and traverse the melting transition, share many similar properties with action potentials. Despite the striking experimental similarities, a comprehensive theoretical study of acoustic pulses in lipid systems is still lacking. Here we demonstrate that an idealized description of an interface near phase transition captures many properties of acoustic pulses in lipid monolayers, as well as action potentials in living cells. The possibility that action potentials may better be described as acoustic pulses in soft interfaces near phase transition is illustrated by the following similar properties: correspondence of time and velocity scales, qualitative pulse shape, sigmoidal response to stimulation amplitude (an `all-or-none' behavior), appearance in multiple observables (particularly, an adiabatic change of temperature), excitation by many types of stimulations, as well as annihilation upon collision. An implication of this work is that crucial functional information of the cell may be overlooked by focusing only on electrical measurements.

physics.bio-ph↗

arXiv2017

Collision of two action potentials in a single excitable cell

It is a common incident in nature, that two waves or pulses run into each other head-on. The outcome of such an event is of special interest, because it allows conclusions about the underlying physical nature of the pulses. The present experimental study dealt with the head-on meeting of two action potentials (AP) in a single excitable plant cell (Chara braunii internode). The membrane potential was monitored at the two extremal regions of an excitable cell. In control experiments, an AP was excited electrically at either end of the cell cylinder. Subsequently, stimuli were applied simultaneously at both ends of the cell in order to generate two APs that met each other head-on. When two action potentials propagated into each other, the pulses did not penetrate but annihilated (N=26 experiments in n=10 cells). APs in excitable plant cells did not penetrate upon meeting head-on. In the classical electrical model, this behavior is specifically attributed to relaxation of ion channel proteins. From an acoustic point of view, annihilation can be viewed as a result of nonlinear material properties the entire system. The present results suggest that APs in excitable animal and plant cells belong to a similar class of nonlinear phenomena. Intriguingly, other excitation waves in biology (intracellular waves, cortical spreading depression, etc.) also annihilate upon collision and are thus expected to follow the same underlying principles as the observed action potentials.

physics.bio-ph↗

arXiv2017

On cell surface deformation during an action potential

The excitation of many cells and tissues is associated with cell mechanical changes. The evidence presented herein corroborates that single cells deform during an action potential (AP). It is demonstrated that excitation of plant cells (Chara braunii internodes) is accompanied by out-of-plane displacements of the cell surface in the micrometer range (1-10 micron). The onset of cellular deformation coincides with the depolarization phase of the AP. The mechanical pulse (i) propagates with the same velocity as the electrical pulse (within experimental accuracy; 10 mm/s), (ii) is reversible, (iii) in most cases of biphasic nature (109 out of 152 experiments) and (iv) presumably independent of actin-myosin-motility. The existence of transient mechanical changes in the cell cortex is confirmed by micropipette aspiration experiments. A theoretical analysis demonstrates that this observation can be explained by a reversible change in the mechanical properties of the cell surface (transmembrane pressure, surface tension and bending rigidity). Taken together, these findings contribute to the ongoing debate about the physical nature of cellular excitability.

physics.bio-ph↗

arXiv2017

Nonlinear fractional waves at elastic interfaces

We derive the nonlinear fractional surface wave equation that governs compression waves at an interface that is coupled to a viscous bulk medium. The fractional character of the differential equation comes from the fact that the effective thickness of the bulk layer that is coupled to the interface is frequency dependent. The nonlinearity arises from the nonlinear dependence of the interface compressibility on the local compression, which is obtained from experimental measurements and reflects a phase transition at the interface. Numerical solutions of our nonlinear fractional theory reproduce several experimental key features of surface waves in phospholipid monolayers at the air-water interface without freely adjustable fitting parameters. In particular, the propagation length of the surface wave abruptly increases at a threshold excitation amplitude. The wave velocity is found to be of the order of 40 cm/s both in experiments and theory and slightly increases as a function of the excitation amplitude. Nonlinear acoustic switching effects in membranes are thus shown to arise purely based on intrinsic membrane properties, namely the presence of compressibility nonlinearities that accompany phase transitions at the interface.

physics.flu-dyn↗

arXiv2016

Biological signaling by interfacial sound pulses. A physics approach

Biological signaling is imagined as a combination of activation and transport. The former is triggered by local molecular interactions and the latter is the result of molecular diffusion. However, other fundamental physical principles of communication have yet to be addressed. We have recently shown, that lipid interfaces allow for the excitation and propagation of sound pulses. Here we demonstrate, that these reversible perturbations can control the activity of membrane embedded enzymes without the necessity of molecular transport. They therefore allow for the rapid communication between distant biological entities (e.g. receptor and enzyme) at the speed of sound, which is here in the order of 1 m/s within the membrane. The mechanism reported provides a new physical framework for biological signaling.

physics.bio-ph↗

arXiv2015

Protons at the speed of sound: Specific biological signaling from physics

Local changes in pH are known to significantly alter the state and activity of proteins and in particular enzymes. pH variations induced by pulses propagating along soft interfaces (e.g. the lipid bilayer) would therefore constitute an important pillar towards a new physical mechanism of biochemical regulation and biological signaling. Here we investigate the pH-induced physical perturbation of a lipid interface and the physiochemical nature of the subsequent acoustic propagation. Pulses are stimulated by local acidification of a lipid monolayer and propagate, in analogy to sound, at velocities controlled by the two-dimensional compressibility of the interface. With transient local pH changes of 0.6 units directly observed at the interface and velocities up to 1.4 m/s this represents hitherto the fastest protonic communication observed. Furthermore simultaneously propagating mechanical and electrical changes in the lipid interface up to 8 mN/m and 100 mV are detected, exposing the thermodynamic nature of these pulses. Finally, these pulses are excitable only beyond a threshold for protonation, determined by the pKa of the lipid head groups. When combined with an enzymatic pH-optimum, the proposed communication can be very specific, thus providing a new physical basis for intra- and intercellular signaling via two-dimensional sound waves at interfaces.

physics.bio-ph↗

arXiv2015

Solitary Shock Waves and Adiabatic Phase Transition in Lipid Interfaces and Nerves

This study shows that the stability of solitary waves excited in a lipid monolayer near a phase boundary requires positive curvature of the adiabats, a known necessary condition in shock compression science. It is further shown that the condition results in a threshold for excitation, saturation of the wave amplitude and the splitting of the wave at the phase boundaries. Splitting in particular confirms that a hydrated lipid interface can undergo condensation on adiabatic heating thus showing retrograde behavior. Finally, using the new theoretical insights and state dependence of conduction velocity in nerves, the curvature of the adiabatic state diagram is shown to be closely tied to the thermodynamic blockage of nerve pulse propagation.

physics.bio-ph↗

arXiv2014

On the excitation of action potentials by protons and its potential implications for cholinergic transmission

One of the most conserved mechanisms for transmission of a nerve pulse across a synapse relies on acetylcholine. Ever since the Nobel-prize winning works of Dale and Loewi, it has been assumed that acetylcholine - subsequent to its action on a postsynaptic cell - is split into inactive by-products by acetylcholinesterase. Herein, this widespread assumption is falsified. Excitable cells (Chara australis internodes), which had previously been unresponsive to acetylcholine, became acetylcholine-sensitive in presence of acetylcholinesterase. The latter was evidenced by a striking difference in cell membrane depolarisation upon exposure to 10 mM intact acetylcholine (deltaV=-2plus/minus5 mV) and its hydrolysate respectively (deltaV=81plus/minus19 mV) for 60 sec. This pronounced depolarization, which also triggered action potentials, was clearly attributed to one of the hydrolysis products: acetic acid (deltaV=87plus/minus9 mV at pH 4.0; choline ineffective in range 1-10 mM). In agreement with our findings, numerous studies in the literature have reported that acids excite gels, lipid membranes, plant cells, erythrocytes as well as neurons. Whether excitation of the postsynaptic cell in a cholinergic synapse is due to protons or due to intact acetylcholine is a most fundamental question that has not been addressed so far.

physics.bio-ph↗

arXiv2014

Evidence for 2D Solitary Sound Waves in a Lipid Controlled Interface and its Biological Implications

Biological membranes by virtue of their elastic properties should be capable of propagating localized perturbations analogous to sound waves. However, the existence and the possible role of such waves in communication in biology remains unexplored. Here we report the first observations of 2D solitary elastic pulses in lipid interfaces, excited mechanically and detected by FRET. We demonstrate that the nonlinearity near a maximum in the susceptibility of the lipid monolayer results in solitary pulses that also have a threshold for excitation. These experiments clearly demonstrate that the state of the interface regulates the propagation of pulses both qualitatively and quantitatively. We elaborate on the striking similarity of the observed phenomenon to nerve pulse propagation and a thermodynamic basis of cell signaling in general.

physics.bio-ph↗

arXiv2013

Phase Transition and dissipation driven budding in lipid vesicles

Membrane budding has been extensively studied as an equilibrium process attributed to the formation of coexisting domains or changes in the vesicle area to volume ratio (reduced volume). In contrast, non-equilibrium budding remains experimentally widely unexplored especially when time scales fall well below the characteristic diffusion time of lipidsτ . We show that localized mechanical perturbations, initiated by driving giant unilamellar vesicles (GUVs) through their lipid phase transition, leads to the immediate formation of rapidly growing, multiply localized, non-equilibrium buds, when the transition takes place at short timescales (<τ). We show that these buds arise from small fluid-like perturbations and grow as spherical caps in the third dimension, since in plane spreading is obstructed by the continuous rigid gel-like matrix. Accounting for both three and two dimensional viscosity, we demonstrate that dissipation decreases the size scale of the system and therefore favours the formation of multiple buds as long as the perturbation takes place above a certain critical rate. This rate depends on membrane and media viscosity and is qualitatively and quantitatively correctly predicted by our theoretical description.

physics.bio-ph↗

arXiv2010

Inertia effects and stress accumulation in a constricted duct: A combined experimental and lattice Boltzmann study

We experimentally and numerically investigate the flow of a Newtonian fluid through a constricted geometry for Reynolds numbers in the range $0.1 - 100$. The major aim is to study non-linear inertia effects at larger Reynolds numbers (>10) on the shear stress evolution in the fluid. This is of particular importance for blood flow as some biophysical processes in blood are sensitive to shear stresses, e.g., the initialization of blood clotting. We employ the lattice Boltzmann method for the simulations. The conclusion of the predictions is that the peak value of shear stress in the constriction grows disproportionally fast with the Reynolds number which leads to a non-linear shear stress accumulation. As a consequence, the combination of constricted blood vessel geometries and large Reynolds numbers may increase the risk of undesired blood clotting.

cond-mat.soft↗