Effects of unstirred layers or transport number discontinuities on the transient and steady-state current-voltage relationships of membranes

The effects of current-induced electrolyte accumulation and depletion on the electrical properties of a two-layered membrane system have been examined. The membrane consisted of a charged, ion permselective layer and an uncharged, non-selective layer. The model was designed to reveal the properties of membranes possessing long pores with ionic charges at one end or of ion-selective membranes bounded by highly unstirred aqueous layers. Electrolyte concentration profiles in the inert layer and their time-dependent changes were obtained from solutions of the diffusion equation under the condition of constant current. The profiles were then used to calculate the voltage developed across the membrane at various times after the current is switched on. The theoretical results are presented in the form of $\text{i-V}$ curves with reduced coordinates that can be used to obtain time-current-voltage relationships for membranes of the type considered having any thickness of the non-selective layer and bathed in any concentration of any 1:1 electrolyte. Experimental results on a model composite membrane were in good agreement with calculations that assume that ion transport occurs only under the influence of electrical potential and concentration gradients, suggesting that in such systems, the combined effects of convection, osmosis, electro-osmosis, and concentration-dependence of diffusion coefficients, activity coefficients, and transference numbers are small. Voltage fluctuations in the form of periodic spikes were observed experimentally at the limiting current density (the current density at which the electrolyte concentration at one surface of the selective layer goes to 0). These phenomena were not seen when the current was in the direction leading to accumulation of electrolyte in the non-selective (unstirred) layer. Such composite membranes can exhibit S-shaped and N-shaped $\text{i-V}$ curves under ramp-voltage and ramp-current clamps, respectively.     

Description of turbulent convection in a plasma with the help of interacting Lorentz oscillators

A four-field model is proposed that describes turbulent plasma convection inside the separatrix during the L-H transition. It is shown that the Braginskii four-field hydrodynamic equations, which describe fluctuations of the electron and ion temperatures, plasma density, and electrostatic potential in tokamak edge plasmas, can be reduced to three Lorentz-like systems of equations coupled through the equation for the kinetic energy of the fluctuations, i.e., to a four-field edge turbulent layer model describing the nonlinear dynamics of convective cells in the presence of a sheared flow. For three coupled oscillators, the critical pressure gradient corresponding to transitions to both L-and H-modes is found to be much lower than that for an individual oscillator, which describes turbulent convection driven by fluctuations of one type. The edge turbulent layer model makes it possible to describe the formation of a transport barrier inside the separatrix during the L-H transition; calculate heat and particle fluxes via ion and electron channels; and, in combination with the transport code for a core plasma, compute the auxiliary heating power required for a transition to the H-mode.     

Unsteady Convection Flow and Heat Transfer over a Vertical Stretching Surface

This paper investigates the effect of thermal radiation on unsteady convection flow and heat transfer over a vertical permeable stretching surface in porous medium, where the effects of temperature dependent viscosity and thermal conductivity are also considered. By using a similarity transformation, the governing time-dependent boundary layer equations for momentum and thermal energy are first transformed into coupled, non-linear ordinary differential equations with variable coefficients. Numerical solutions to these equations subject to appropriate boundary conditions are obtained by the numerical shooting technique with fourth-fifth order Runge-Kutta scheme. Numerical results show that as viscosity variation parameter increases both the absolute value of the surface friction coefficient and the absolute value of the surface temperature gradient increase whereas the temperature decreases slightly. With the increase of viscosity variation parameter, the velocity decreases near the sheet surface but increases far away from the surface of the sheet in the boundary layer. The increase in permeability parameter leads to the decrease in both the temperature and the absolute value of the surface friction coefficient, and the increase in both the velocity and the absolute value of the surface temperature gradient.     

Kinetics of the fast electric signal from oriented purple membrane

The photoinduced electric response of oriented purple membranes associated with processes before the K-intermediate decay of bacteriorhodopsin was measured in the 180-300 K temperature range. These response signals consist of two kinetically distinct components (both temperature dependent). The experimental data show a correlation between the time constants of the rise of the signal and solution resistance. A model is proposed to assign these components to two diffusion-limited processes of charge displacement in the solution. The displacement is caused by the electric field of the photoinduced transient dipole which is formed in the primary act of the bacteriorhodopsin photocycle. The two processes are assigned as: (a) the conduction of electrical current through H-bonds (time resolved only in the temperature range 180-200 K) and (b) the diffusion of charges through the interfacial layer.     

Electromechanical stresses and the effect of pH on membrane structure.

The presence of an electric field in a membrane creates stresses which lead to a mechanical compression of the membrane material. It is shown that considerations of this effect yield results consistent with the observed differential effect of pH on the plasma membrane substructure if the central electron-lucent layer in OsO4-fixed membranes is identified with the ionic depletion in the double-fixed charge model of cell membranes.     

Specific electric resistance of a narrow gap between two lipid bilayer membranes

The study deals with measurements of specific electric resistance of a narrow gap between two bilayer lipid membranes (BLM). The technique is based upon the influence of the outer electric field on the distance between two BLM which has been revealed earlier. The curve of specific resistance/solution ionic strength obtained suggests that surface conductivity is significant in similar systems. In gaps narrower than 40 nm possible increase of water viscosity must be also taken into consideration. A hypothetic mechanism of electric connection between cells is proposed.     

Electromechanical stresses and the effect of pH on membranes structure

The presence of an electric field in a membrane creates stresses which lead to a mechanical compression of the membrane material.It is shown that considerations of this effect yield results consistent with the observed differential effect of pH on the plasma membrane substructure if the central electron-lucent layer in OsO₄-fixed membranes is identified with the ionic depletion in the double-fixed charge model of cell membranes.     

Ion transport and the vibrating probe.

The theory of ion transport in the vicinity of a vibrating probe is developed. It is shown that the convection loops produced by the probe will not affect the electrical current density, assuming that the action of the probe does not affect the sources of the current in the biological system. However, the convection loops will significantly alter the ion concentration gradients in the unstirred layer near a tissue or cell surface. The concentration gradients within each convection loop will be reduced, while the concentration gradients between the loops and outside of the loops will be increased relative to the gradients existing without the probe. As a consequence, the electrical potential gradients can be changed relative to the potential gradients existing in the absence of the convection caused by the probe. If the mobility of the ion species carrying the electrical current is greater than the average ion mobility in the medium, then a decrease in ion concentration gradient will be accompanied by an increase in electrical potential gradient, while an increase in concentration gradient will be accompanied by a decrease or even a reversal of electrical potential gradient. Thus, the electrical potential gradient measured by the probe will depend on the concentration gradient in the vicinity of the probe, which will depend in turn on the spatial relation of the convection loops to the probe. An example of the effect of the convection loops on ion concentration and electrical potential is obtained from the theory via a numerical computer calculation. Experimental tests of this theory are discussed.     

Extracellular ion transport

It is a misconception to think that extracellular electric currents of biologic origin are carried by all of the ions in the extracellular medium. It is established that, for certain reasonable boundary conditions, only those ion species that are transported across the plasma membranes of the biological system, or that chemically derive from or contribute to these species, can contribute to the extracellular electric current at steady state. In the absence of convection, the extracellular current will be carried largely by diffusion of the transported ion species, at steady state. Extracellular electric potential gradients are shown to arise in a secondary manner, as a result of the electroneutrality condition. The effect of non-turbulent convection is included in the derivations and discussion.     

Traffic noise inhibits cognitive performance in a songbird

Noise pollution is commonly associated with human environments and mounting evidence indicates that noise has a variety of negative effects on wildlife. Noise has also been linked to cognitive impairment in humans and because many animals use cognitively intensive processes to overcome environmental challenges, noise pollution has the potential to interfere with cognitive function in animals living in urban areas or near roads. We experimentally examined how road traffic noise impacts avian cognitive performance by testing adult zebra finches (Taeniopygia guttata) on a battery of foraging tasks in the presence or absence of traffic noise playback. Here, we show that traffic noise reduces cognitive performance, including inhibitory control, motor learning, spatial memory and social learning, but not associative colour learning. This study demonstrates a novel mechanism through which anthropogenic noise can impact animals, namely through cognitive interference, and suggests that noise pollution may have previously unconsidered consequences for animals.     

Measurement of Extracellular Ion Fluxes Using the Ion-selective Self-referencing Microelectrode Technique

Cells from animals, plants and single cells are enclosed by a barrier called the cell membrane that separates the cytoplasm from the outside. Cell layers such as epithelia also form a barrier that separates the inside from the outside or different compartments of multicellular organisms. A key feature of these barriers is the differential distribution of ions across cell membranes or cell layers. Two properties allow this distribution: 1) membranes and epithelia display selective permeability to specific ions; 2) ions are transported through pumps across cell membranes and cell layers. These properties play crucial roles in maintaining tissue physiology and act as signaling cues after damage, during repair, or under pathological condition. The ion-selective self-referencing microelectrode allows measurements of specific fluxes of ions such as calcium, potassium or sodium at single cell and tissue levels. The microelectrode contains an ionophore cocktail which is selectively permeable to a specific ion. The internal filling solution contains a set concentration of the ion of interest. The electric potential of the microelectrode is determined by the outside concentration of the ion. As the ion concentration varies, the potential of the microelectrode changes as a function of the log of the ion activity. When moved back and forth near a source or sink of the ion (i.e. in a concentration gradient due to ion flux) the microelectrode potential fluctuates at an amplitude proportional to the ion flux/gradient. The amplifier amplifies the microelectrode signal and the output is recorded on computer. The ion flux can then be calculated by Fick’s law of diffusion using the electrode potential fluctuation, the excursion of microelectrode, and other parameters such as the specific ion mobility. In this paper, we describe in detail the methodology to measure extracellular ion fluxes using the ion-selective self-referencing microelectrode and present some representative results.     

Generation of large-scale structures by strong drift turbulence

The previously existing quasilinear theory of the generation of a large-scale radial electric field by small-scale drift turbulence in a plasma is generalized for the case of strong turbulence which is usually observed in experiments. The geostrophic equation (i.e., the reduced Charney-Hasegawa-Mima equation) is used to construct a systematic theory in the two-scale direct interaction approximation. It is shown that, as in the quasilinear case, drift turbulence results in a turbulent viscosity effect and leads to the renormalization of the Poisson bracket in the Charney-Hasegawa-Mima equation. It is found that, for strong drift turbulence, the viscosity coefficient is represented as a sum of two parts, which are comparable in magnitude. As in quasilinear theory, the first part is determined by the second-order correlation functions of the turbulent field and is always negative. The second part is proportional to the third-order correlation functions, and the sign of its contribution to the turbulent viscosity coefficient depends strongly on the turbulence spectrum. The turbulent viscosity coefficient is calculated numerically for the Kolmogorov spectra, which characterize the inertial interval of the drift turbulence.     

Nonequilibrium voltage fluctuations in biological membranes. II. Voltage and current noise generated by ion carriers, channels and electrogenic pumps

As applications of the general theoretical framework of charge transport in biological membranes and related voltage and current noise, a number of model calculations are presented for ion carriers, rigid channels, channels with conformational substates and electrogenic pumps. The results are discussed with special reference to the problem of threshold values for sensory transduction processes and their limitations by voltage fluctuations. Furthermore, starting from the special results of model calculations, an attempt is made to determine more general aspects of electric fluctuations generated by charge-transport processes in biological membranes: different frequency dependences of voltage and current noise, and dependence of noise intensities with increasing distance from the equilibrium state.     

Conductance versus current noise in a neuronal model for noisy subthreshold oscillations and related spike generation

Biological systems are notoriously noisy. Noise, therefore, also plays an important role in many models of neural impulse generation. Noise is not only introduced for more realistic simulations but also to account for cooperative effects between noisy and nonlinear dynamics. Often, this is achieved by a simple noise term in the membrane equation (current noise). However, there are ongoing discussions whether such current noise is justified or whether rather conductance noise should be introduced because it is closer to the natural origin of noise. Therefore, we have compared the effects of current and conductance noise in a neuronal model for subthreshold oscillations and action potential generation. We did not see any significant differences in the model behavior with respect to voltage traces, tuning curves of interspike intervals, interval distributions or frequency responses when the noise strength is adjusted. These findings indicate that simple current noise can give reasonable results in neuronal simulations with regard to physiological relevant noise effects.     

Multispectral imaging of ion transport in neutral carrier-based cation-selective membranes.

BACKGROUND: High-resolution spectroscopic imaging of the cross section of ion-selective membranes during real-time electrochemical measurements is termed spectroelectrochemical microscopy (SpECM). SpECM is aimed for optimizing the experimental conditions in mass transport controlled ion-selective electrode (ISE) membranes for improved detection limit. METHODS: The SpECM measurements are performed in a thin layer electrochemical cell. The key element of the cell is a membrane strip spacer ring assembly which forms a two compartment electrochemical cell. The cell is placed onto the stage of a microscope and the membrane strip is positioned in the center of the field of view. A slice of the image is focused onto the entrance slit of the imaging spectrometer. RESULTS: SpECM has been used for the determination of the diffusion coefficients of different membrane ingredients and for the quantitative assessment of the charged site concentrations in ISE membranes and membrane plasticizers. In addition, changes in the concentration profiles of the ionophore (free and complexed) and charged mobile sites inside the ISE membranes are documented upon the application of large external voltages. CONCLUSIONS: This account demonstrates the power and advantages of SpECM, a multispectral imaging method for investigations of mass transport processes in ISE membranes during electrochemical measurements.     

Skin-friction drag analysis from the forced convection modeling in simplified underwater swimming

This study deals with skin-friction drag analysis in underwater swimming. Although lower than profile drag, skin-friction drag remains significant and is the second and only other contribution to total drag in the case of underwater swimming. The question arises whether varying the thermal gradient between the underwater swimmer and the pool water may modify the surface shear stress distribution and the resulting skin-friction drag acting on a swimmer's body. As far as the authors are aware, such a question has not previously been addressed. Therefore, the purpose of this study was to quantify the effect of this thermal gradient by using the integral formalism applied to the forced convection theory. From a simplified model in a range of pool temperatures (20-30 degrees C) it was demonstrated that, whatever the swimming speeds, a 5.3% reduction in the skin-friction drag would occur with increasing average boundary-layer temperature provided that the flow remained laminar. However, as the majority of the flow is actually turbulent, a turbulent flow analysis leads to the major conclusion that friction drag is a function of underwater speed, leading to a possible 1.5% reduction for fast swimming speeds above 1m/s. Furthermore, simple correlations between the surface shear stress and resulting skin-friction drag are derived in terms of the boundary-layer temperature, which may be readily used in underwater swimming situations.     

Correlation analysis of electrical noise in lipid bilayer membranes: kinetics of gramicidin A channels.

If a membrane contains ion-conducting channels which form and disappear in a random fashion, an electric current which is passed through the membrane under constant voltage shows statistical fluctuations. Information on the kinetics of channel formation and on the conductance of the single channel may be obtained by analyzing the electrical noise generated in a membrane containing a great number of channels. For this purpose the autocorrelation function of the current noise is measured at different concentrations of the channel-forming substance. As a test system for the application of this technique we have used lipid bilayer membranes doped with gramicidin A. From the correlation time of the current noise generated by the membrane, the rate constants of formation (k-R) and dissociation (k-D) of the channels could be determined. In addition, the mean square of the current fluctuations yielded the single-channel conductance lambda. The values of k-R, k-D, and lambda obtained from the noise analysis agreed closely with the values determined by relaxation measurments and single-channel experiments.     

Protein blotting in uniform or gradient electric fields

Experimental data and computer mapping were used to analyze electric fields generated by a variety of electrode arrays in protein blotting apparatus. Asymmetric electrode arrays were found to generate nonuniform fields that effected uneven transfers of 125I-labeled albumin from gels to nitrocellulose membranes. Symmetric arrays with multiple (four), independent wire electrodes, supplied individually with electric current, generated the most uniform fields and effected the most even transfers of the test protein. With multiple independent electrodes, gradient electric fields can be generated in which differences in electrophoretic elution between large and small proteins can be eliminated. Transfer apparatus with either uniform or gradient electric fields are expected to improve qualitative results and make possible quantitation of protein blotting.     

5-Methylphenazinium methylsulfate mediates cyclic electron flow and proton gradient dissipation in chromaffin-vesicle membranes

When 5-methylphenazinium methylsulfate and a reductant (ascorbate or NADH) are added together to a suspension of resealed chromaffin-vesicle membranes, the pH gradient (inside acidic) and the membrane potential (inside positive) established by the H(+)-translocating adenosine triphosphatase (ATPase) are rapidly dissipated. Dissipation of the pH gradient may be observed using either the optical probe acridine orange or the weak base methylamine. Dissipation of the membrane potential may be observed using the potential-dependent dye oxonol VI. A reductant and 5-methylphenazinium methylsulfate added in combination will also abolish a K+ diffusion potential across chromaffin-vesicle membranes but not across liposome membranes. 5-Methylphenazinium methylsulfate oxidizes cytochrome b561 in chromaffin-vesicle ghosts. Ascorbate readily reduces cytochrome b561, but reduction of cytochrome b561 by NADH is greatly enhanced in the presence of 5-methylphenazinium methylsulfate. These results are consistent with a mechanism in which proton gradient dissipation (a net efflux of H+) is caused by an influx of electrons through the membrane-protein cytochrome b561 coupled with an efflux of H carried by the reduced species 5-methyl-10-hydrophenazine. Although 5-methylphenazinium has been thought to accumulate within acidic vesicles as a weak base, this accounts for neither proton gradient dissipation nor for intravesicular accumulation of the compound.