Collective movement of ions in the cytoplasm

A theoretical model of propagation in the cytoplasm of self-consistent electromagnetic waves of the millimeter-infrared range has been developed, cytoplasmic ions surrounded by water ??coats?? being the main carriers of these waves. It has been discovered that not only own long-wavelength transverse oscillations, but also longitudinal waves that cannot leave the cytoplasm can exist in tissues of living organisms. Frequencies and logarithmic decrements of such perturbations have been found, and it is shown that these frequencies are close to those of ion oscillations inside the ??coats.?? Passage of laser radiation in bioobjects at the indicated frequencies has been analyzed, revealing bands of body impenetrability for waves. A new mechanism of swinging of cytoplasm own oscillations is proposed, based on the existence of an extreme border of the ion movement area. It has been shown that with this mechanism, the electric field amplitude for the longitudinal waves is six to seven orders of magnitude greater than the Planck fluctuation level.     

Electrophysiological consequences of leukotrienes applied on isolated rat retina

Scotopic vision is the result of a cascade of light-dependent biochemical events in rod outer segments (ROS) involving mainly a cGMP-modulation of sodium current. This modification of ionic currents induces changes of membrane potential which generates electroretinographic (ERG) waves. As (i) ERG disturbances are commonly recorded in hypoxic and inflammatory retinal diseases (ii) leukotrienes (LTs), a very potent mediators of inflammation, disturb ionic exchanges in several artificial or natural membrane systems, we undertook the investigation of the effects of LTs on ERG record in mammalian isolated retina. LTB4, LTC4 and LTD4, all induced a dose-dependent marked reduction of the b wave amplitude of ERG. This effect is correlated with a significant decrease in the survival time of the retina. The analysis of the modification of ERG indicates that LTs exhibit a real toxic effect since b wave is mainly affected while P III wave is unchanged. Comparatively with other nervous cells, this phenomenon may be attributed to an increase in Na+ permeability of ROS. It is suggested that LTs may be involved in the development of inflammatory or ischemic retinal diseases.     

Computer simulation of bend propagation by axoplasmic microtubules

The generation of bending waves by microtubules in squid nerve axoplasm has been modelled using appropriately modified versions of computer programs developed previously for simulation of flagellar bending waves. The results confirm that a constant longitudinal force directed along the axis of the microtubule is sufficient to cause the generation of regular oscillations and propagated bending waves when the forward gliding movement of the microtubule is obstructed. No control mechanism is required to modulate the active force-generating system. In order to obtain bending waves similar to those observed experimentally, it was necessary to use a model for the force-generating system in which the active force decreases with increasing sliding velocity. If the elastic bending resistance of axoplasmic microtubules is similar to that of microtubules in sperm terminal filaments, the longitudinal force per unit length generated by the axoplasmic microtubules must be of the same order of magnitude as the force generated by dynein arms along the doublet microtubules of eukaryotic flagella.     

Spatiotemporal electrical and motility mapping of distension-induced propagating oscillations in the murine small intestine

Since the development of knockout animals, the mouse has become an important model to study gastrointestinal motility. However, little information is available on the electrical and contractile activities induced by distension in the murine small intestine. Spatiotemporal electrical mapping and mechanical recordings were made from isolated intestinal segments from different regions of the murine small intestine during distension. The electrical activity was recorded with 16 extracellular electrodes while motility was assessed simultaneously by tracking the border movements with a digital camera. Distension induced propagating oscillatory contractions in isolated intestinal segments. These propagating contractions were dictated by the underlying propagating slow wave with superimposed spikes. The frequencies, velocities, and direction of the propagating oscillations strongly correlated with the frequencies (r = 0.86), velocities (r = 0.84), and direction (r = 1) of the electrical slow waves. N(omega)-nitro-L-arginine methyl ester decreased the maximal diameter of the segment and reduced the peak contraction amplitude of the propagating oscillatory contractions, whereas atropine and verapamil blocked the propagating oscillations. Tetrodotoxin had little effect on the maximal diameter and peak contraction amplitude. In conclusion, distension in the murine small intestine does not initiate peristaltic reflexes but induces a propagating oscillatory motor pattern that is determined by propagating slow waves with superimposed spikes. These spikes are cholinergic and calcium dependent.     

Intercellular water exchanges trigger soliton-like waves in multicellular systems

Oscillations and waves are ubiquitous in living cellular systems. Generations of these spatiotemporal patterns are generally attributed to some mechanochemical feedbacks. Here, we treat cells as open systems, i.e., water and ions can pass through the cell membrane passively or actively, and reveal a new origin of wave generation. We show that osmotic shocks above a shock threshold will trigger self-sustained cell oscillations and result in long-range waves propagating without decrement, a phenomenon that is analogous to the excitable medium. The traveling wave propagates along the intercellular osmotic pressure gradient, and its wave speed scales with the magnitude of intercellular water flows. Furthermore, we also find that the traveling wave exhibits several hallmarks of solitary waves. Together, our findings predict a new mechanism of wave generation in living multicellular systems. The ubiquity of intercellular water exchanges implies that this mechanism may be relevant to a broad class of systems.     

Resonance oscillations in the human arterial system]

The paper summarizes the results of the experiments aimed at obtaining sphygmograms of peripheral and carotid arteries with due regard to the values of longitudinal dimensions of body and extremities in healthy subjects. Mathematical equations expressing the fact that dicrotic waves recorded on sphygmograms are the reflections of blood eigentones coinciding with resonance oscillations have been derived. It is proved that at least two partial vibration systems oscillating with different own frequencies are present in human arteries. Conditions under which the resonance of constituent frequencies of pulsatile pressure waves and output waves in arteries occurs have been determined. From this point of view a new explanation for the well-known phenomenon of the pulsatile wave amplitude increase from the heart towards peripheric regions is proposed.     

Effect of a strong monochromatic electromagnetic wave with circular polarization on one-dimensional wake waves in plasma

A study is made of the excitation of wake waves by a one-dimensional bunch of charged particles in an electron plasma in the presence of an intense monochromatic pump wave with circular polarization. In the main state (in the absence of a bunch), the interaction between a pump wave and a plasma is described by the Maxwell equations and the nonlinear relativistic hydrodynamic equations for a cold plasma. The excitation of linear waves by a one-dimensional bunch is investigated against a cold plasma background. It is shown that, in a certain range of the parameter values of the bunch, pump wave, and plasma, the amplitude of the excited transverse waves grows as the energy of the bunch particles increases until the relativistic factor of the bunch reaches a certain threshold value above which the transverse wave amplitude becomes essentially independent of the bunch particle energy and grows as the intensity and frequency of the pump wave increase. The amplitude and wavelength of the longitudinal field, which is shown to depend weakly on the energy of the bunch particles, grows with increasing the pump wave intensity.     

Diagnostics of plasma oscillations by the field of surface waves guided by a dielectric waveguide

The potentialities of the diagnostic method for determining the plasma parameters by recording the surface waves guided by a dielectric waveguide and scattered by plasma oscillations are discussed. The use of surface (slowed) waves makes it possible to improve both the sensitivity and spatial resolution of measurements. The scattering is the most intense near the waveguide cutoff, at which the dependence of the wave propagation constant on the plasma density is the steepest. It is shown experimentally that the method proposed makes it possible to determine the discharge plasma density and electron energy and to estimate the amplitude of the RF field of the plasma waves forming the discharge and the amplitude of plasma density oscillations in these waves. The data obtained from the measurements of the amplitudes of both high-and low-frequency plasma density oscillations by the proposed method agree satisfactorily with theoretical predictions. The experimental data on the plasma density are confirmed by other diagnostic measurements. The ways of reducing measurement errors are proposed.     

Filling of partially collapsed compliant tubes

Filling of a thin-walled, highly compliant tube in a partially collapsed condition is studied. The theory, based on one-dimensional flow, takes account of friction, longitudinal tension, and the highly nonlinear pressure-area law for the tube. Various aspects of filling behavior are revealed by alternative calculations using: (i) the method of characteristics; (ii) numerical integration of the continuity, momentum, and tube-law equations; and (iii) a crude but simple lumped-element capacitance-inertance-resistance model. Varied phenomena appear. At high Reynolds number, these include dispersive wave trains associated with circumferential bending stiffness and longitudinal tension, nonlinear changes of wave form, development of highly asymmetrical wave reflections, and sloshing. At low Reynolds number, the area changes with time in a diffusivelike manner. The experiments exhibited the dispersive phenomena predicted by the theory.     

Generation of auroral kilometric radiation by a finite-size source in a dipole magnetic field

Generation, amplification, and propagation of auroral kilometric radiation in a narrow three-dimensional plasma cavity in which a weakly relativistic electron beam propagates is studied in the geometrical optics approximation. It is shown that the waves that start with a group velocity directed earthward and have optimal relation between the wave vector components determining the linear growth rate and the wave residence time inside the amplification region undergo the largest amplification. Taking into account the longitudinal velocity of fast electrons results in the shift of the instability domain toward wave vectors directed to the Earth and leads to a change in the dispersion relation, due to which favorable conditions are created for the generation of waves with frequencies above the cutoff frequency for the cold background plasma at the wave generation altitude. The amplification factor for these waves is lower than for waves that have the same wave vectors but are excited by the electron beams with lower velocities along the magnetic field. For waves excited at frequencies below the cutoff frequency of the background plasma at the generation altitude, the amplification factor increases with increasing longitudinal electron velocity, because these waves reside for a longer time in the amplification region.     

Functional state of the axonemal dyneins during flagellar bend propagation

When mouse spermatozoa swim in media of high viscosity, additional waves of bending are superimposed on the primary traveling wave. The additional (secondary) waves are relatively small in scale and high in frequency. They originate in the proximal part of the interbend regions. The initiation of secondary bending happens only in distal parts of the flagellum. The secondary waves propagate along the interbends and then tend to die out as they encounter the next-most-distal bend of the primary wave, if that bend exceeds a certain angle. The principal bends of the primary wave, being of greater angle than the reverse bends, strongly resist invasion by the secondary waves; when a principal bend of the primary wave propagates off the flagellar tip, the secondary wave behind it suddenly increases in amplitude. We claim that the functional state of the dynein motors in relation to the primary wave can be deduced from their availability for recruitment into secondary wave activity. Therefore, only the dyneins in bends are committed functionally to the maintenance and propagation of the flagellar wave; dyneins in interbend regions are not functionally committed in this way. We equate functional commitment with tension-generating activity, although we argue that the regions of dynein thus engaged nevertheless permit sliding displacements between the doublets.     

Electrophysiological consequences of leukotrienes applied on isolated rat retina

Scotopic vision is the result of a cascade of light-dependent biochemical events in rod outer segments (ROS) involving mainly a cGMP-modulation of sodium current. This modification of ionic currents induces changes of membrane potential which generates electroretinographic (ERG) waves. As (i) ERG disturbances are commonly recorded in hypoxic and inflammatory retinal diseases (ii) leukotrienes (LTs), a very potent mediators of inflammation, disturb ionic exchanges several artificial or natural membrane systems, we undertook the investigation of the effects of LTs on ERG record in mammalian isolated retina. LTB₄, LTC₄ and LTD₄, all induced a dose-dependent marked reduction of the b wave amplitude of ERG. This effect is correlated with a significant decrease in the survival time of the retina. The analysis of the modification of ERG indicates that LTs exhibit a real toxic effect since b wave is mainly affected while P III wave is unchanged. Comparatively with other nervous cells, this phenomenon may be attributed to an increase in Na⁺ permeability of ROS. It is suggested that LTs may be involved in the development of inflammatory or ischemic retinal diseases.     

Wave Propagation through a Viscous Incompressible Fluid Contained in an Initially Stressed Elastic Tube

To have a better understanding of the flow of blood in arteries a theoretical analysis of the pressure wave propagation through a viscous incompressible fluid contained in an initially stressed tube is considered. The fluid is assumed to be Newtonian. The tube is taken to be elastic and isotropic. The analysis is restricted to tubes with thin walls and to waves whose wavelengths are very large compared with the radius of the tube. It is further assumed that the amplitude of the pressure disturbance is sufficiently small so that nonlinear terms of the inertia of the fluid are negligible compared with linear ones. Both circumferential and longitudinal initial stresses are considered; however, their origins are not specified. Initial stresses enter equations as independent parameters. A frequency equation, which is quadratic in the square of the propagation velocity is obtained. Two out of four roots of this equation give the velocity of propagation of two distinct outgoing waves. The remaining two roots represent incoming waves corresponding to the first two waves. One of the waves propagates more slowly than the other. As the circumferential and/or longitudinal stress of the wall increases, the velocity of propagation and transmission per wavelength of the slower wave decreases. The response of the fast wave to a change in the initial stress is on the opposite direction.     

On the transmission of external pressure fluctuations to the cerebrospinal fluid

A theory has been formulated to explain the manner in which external pressure fluctuations are transmitted to the cerebrospinal fluid (CSF). The theory is based upon a three-compartment model which consists of the cerebral ventricles, the basal cisterns and spinal subarachnoid space, and the cortical subarachnoid space. The external pressure disturbance is represented by a Fourier series summed over the frequency ω. The mathematical analysis leads to a time constant τ which depends upon the compliances of the spinal region and sources of external pressure fluctuations, the rate of CSF absorption and the rate of fluid transfer between compartments. For arterial pulsations where ωτ ? 1, the theory is in accord with the experimental observations that (i) the arterial and CSF pulse waves are nearly identical in shape, and (ii) the amplitude of the CSF pulse wave increases with intracranial pressure. Moreover, it predicts that the amplitude of the wave will be larger in the spinal region than in the ventricles. The theory also accounts for the observation of one per minute pulse waves observed in hydrocephalic patients with decreased absorption rates.     

The physical basis of alpha waves in the electroencephalogram and the origin of the “Berger effect”

Synchronised activity, differing in phase in different populations of neurons, plays an important role in existing theories on the function of brain oscillations (e.g., temporal correlation hypothesis). A prerequisite for this synchronisation is that stimuli are capable of affecting (resetting) the phase of brain oscillations. Such a change in the phase of brain waves is also assumed to underlie the Berger effect: when observers open their eyes, the amplitude of EEG oscillations in the alpha band (8–13 Hz) decreases significantly. This finding is usually thought to involve a desynchronisation of activity in different neurons. For functional interpretations of brain oscillations in the visual system, it therefore seems to be crucial to find out whether or not the phase of brain oscillations can be affected by visual stimuli. To answer this question, we investigated whether alpha waves are generated by a linear or a nonlinear mechanism. If the mechanism is linear – in contrast to nonlinear ones – phases cannot be reset by a stimulus. It is shown that alpha-wave activity in the EEG comprises both linear and nonlinear components. The generation of alpha waves basically is a linear process and flash-evoked potentials are superimposed on ongoing alpha waves without resetting their phase. One nonlinear component is due to light adaptation, which contributes to the Berger effect. The results call into question theories about brain-wave function based on temporal correlation or event-related desynchronisation.Electronic Supplementary Material: Supplementary material is available for this article at     

Role of membrane potential in vasomotion of isolated pressurized rat arteries

Vasomotion, the phenomenon of vessel diameter oscillation, regulates blood flow and resistance. The main parameters implicated in vasomotion are particularly the membrane potential and the cytosolic free calcium in smooth muscle cells. In this study, these parameters were measured in rat perfused-pressurized mesenteric artery segments. The application of norepinephrine (NE) caused rhythmic diameter contractions and membrane potential oscillations (amplitude; 5.3 +/- 0.3 mV, frequency; 0.09 +/- 0.01 Hz). Verapamil (1 microM) abolished this vasomotion. During vasomotion, 10(-5) M ouabain (Na(+)-K(+) ATPase inhibitor) decreased the amplitude of the electrical oscillations but not their frequency (amplitude; 3.7 +/- 0.3 mV, frequency; 0.08 +/- 0.002 Hz). Although a high concentration of ouabain (10(-3) M) (which exhibits non-specific effects) abolished both electrical membrane potential oscillations and vasomotion, we conclude that the Na+-K+ ATPase could not be implicated in the generation of the membrane potential oscillations. We conclude that in rat perfused-pressurized mesenteric artery, the slow wave membrane type of potential oscillation by rhythmically gating voltage-dependent calcium channels, is responsible for the oscillation of intracellular calcium and thus vasomotion.     

On the roles of Ca2+ diffusion, Ca2+ buffers, and the endoplasmic reticulum in IP3-induced Ca2+ waves.

We have investigated the effects of Ca2+ diffusion, mobile and stationary Ca2+ buffers in the cytosol, and Ca2+ handling by the endoplasmic reticulum on inositol 1,4,5-trisphosphate-induced Ca2+ wave propagation. Rapid equilibration of free and bound Ca2+ is used to describe Ca2+ sequestration by buffers in both the cytosol and endoplasmic reticulum (ER) lumen. Cytosolic Ca2+ regulation is based on a kinetic model of the inositol 1,4,5-trisphosphate (IP3) receptor of De Young and Keizer that includes activation and inhibition of the IP3 receptor Ca2+ channel in the ER membrane and SERCA Ca2+ pumps in the ER. Diffusion of Ca2+ in the cytosol and the ER and the breakdown and diffusion of IP3 are also included in our calculations. Although Ca2+ diffusion is severely limited because of buffering, when conditions are chosen just below the threshold for Ca2+ oscillations, a pulse of IP3 or Ca2+ results in a solitary trigger wave that requires diffusion of Ca2+ for its propagation. In the oscillatory regime repetitive wave trains are observed, but for this type of wave neither the wave shape nor the speed is strongly dependent on the diffusion of Ca2+. Local phase differences lead to waves that are predominately kinematic in nature, so that the wave speed (c) is related to the wavelength (lambda) and the period of the oscillations (tau) approximately by the formula c = lambda/tau. The period is determined by features that control the oscillations, including [IP3] and pump activity, which are related to recent experiments. Both solitary waves and wave trains are accompanied by a Ca2+ depletion wave in the ER lumen, similar to that observed in cortical preparations from sea urchin eggs. We explore the effect of endogenous and exogenous Ca2+ buffers on wave speed and wave shape, which can be explained in terms of three distinct effects of buffering, and show that exogenous buffers or Ca2+ dyes can have considerable influence on the amplitude and width of the waves.     

Kinematics of human spermatozoa

A microcinematographic (50 f/s) study was performed on motile human spermatozoa. Eighty percent were found to have a linear trajectory and a pseudo-sinusoidal head displacement pattern. Throughout their progression, the spermatozoa periodically rotated on their longitudinal axis at a frequency equal to that of flagellar wave formation. These waves were found always to begin on the same side of the cell and to propagate in the flattened plane of the head until the moment of rotation. At this time the wave had reached a point near the middle of the flagellum. Beyond this point, the flagellum moves out of the plane of the head. Different variables used to characterize the movement of spermatozoa included the velocity of progression, amplitude and velocity of head displacement, frequency of rotation, wave amplitude, and propagation velocity of the flagellar wave. Among these variables, it was the propagation velocity of the wave that was found to be best correlated with the velocity of spermatozoan progression. This flagellar wave exhibited two stages, one of initiation and one of propagation.     

Molecular dynamics simulation of structural changes of lipid bilayers induced by shock waves: Effects of incident angles

Unsteady and nonequilibrium molecular dynamics simulations of the response of dipalmitoylphosphatidylcholine (DPPC) bilayers to the shock waves of various incident angles are presented. The action of an incident shock wave is modeled by adding a momentum in an oblique direction to water molecules adjacent to a bilayer. We thereby elucidate the effects of incident shock angles on (i) collapse and rebound of the bilayer, (ii) lateral displacement of headgroups, (iii) tilts of lipid molecules, (iv) water penetration into the hydrophobic region of the bilayer, and (v) momentum transfer across the bilayer. The number of water molecules delivered into the hydrophobic region is found to be insensitive to incident shock angles. The most important structural changes are the lateral displacement of headgroups and tilts of lipid molecules, which are observed only in the half of the bilayer directly exposed to a shock wave for all incident shock angles studied here. As a result, only the normal component of the added oblique momentum is substantially transferred across the bilayer. This also suggests that the irradiation by shock waves may induce a jet-like streaming of the cytoplasm toward the nucleus.     

Molecular dynamics simulation of structural changes of lipid bilayers induced by shock waves: Effects of incident angles

Unsteady and nonequilibrium molecular dynamics simulations of the response of dipalmitoylphosphatidylcholine (DPPC) bilayers to the shock waves of various incident angles are presented. The action of an incident shock wave is modeled by adding a momentum in an oblique direction to water molecules adjacent to a bilayer. We thereby elucidate the effects of incident shock angles on (i) collapse and rebound of the bilayer, (ii) lateral displacement of headgroups, (iii) tilts of lipid molecules, (iv) water penetration into the hydrophobic region of the bilayer, and (v) momentum transfer across the bilayer. The number of water molecules delivered into the hydrophobic region is found to be insensitive to incident shock angles. The most important structural changes are the lateral displacement of headgroups and tilts of lipid molecules, which are observed only in the half of the bilayer directly exposed to a shock wave for all incident shock angles studied here. As a result, only the normal component of the added oblique momentum is substantially transferred across the bilayer. This also suggests that the irradiation by shock waves may induce a jet-like streaming of the cytoplasm toward the nucleus.