Tonotopically arranged traveling waves in the miniature hearing organ of bushcrickets

Place based frequency discrimination (tonotopy) is a fundamental property of the coiled mammalian cochlea. Sound vibrations mechanically conducted to the hearing organ manifest themselves into slow moving waves that travel along the length of the organ, also referred to as traveling waves. These traveling waves form the basis of the tonotopic frequency representation in the inner ear of mammals. However, so far, due to the secure housing of the inner ear, these waves only could be measured partially over small accessible regions of the inner ear in a living animal. Here, we demonstrate the existence of tonotopically ordered traveling waves covering most of the length of a miniature hearing organ in the leg of bushcrickets in vivo using laser Doppler vibrometery. The organ is only 1 mm long and its geometry allowed us to investigate almost the entire length with a wide range of stimuli (6 to 60 kHz). The tonotopic location of the traveling wave peak was exponentially related to stimulus frequency. The traveling wave propagated along the hearing organ from the distal (high frequency) to the proximal (low frequency) part of the leg, which is opposite to the propagation direction of incoming sound waves. In addition, we observed a non-linear compression of the velocity response to varying sound pressure levels. The waves are based on the delicate micromechanics of cellular structures different to those of mammals. Hence place based frequency discrimination by traveling waves is a physical phenomenon that presumably evolved in mammals and bushcrickets independently.     

Multiple-spike waves in a one-dimensional integrate-and-fire neural network

This paper builds on the past study of single-spike waves in one-dimensional integrate-and-fire networks to provide a framework for the study of waves with arbitrary (finite or countably infinite) collections of spike times. Based on this framework, we prove an existence theorem for single-spike traveling waves, and we combine analysis and numerics to study two-spike traveling waves, periodic traveling waves, and general infinite spike trains. For a fixed wave speed, finite-spike waves, periodic waves, and other infinite-spike waves may all occur, and we discuss the relationships among them. We also relate the waves considered analytically to waves generated in numerical simulations by the transient application of localized excitation.Key words or phrases:Traveling waves, Integrate-and-fire network, Excitatory synaptic coupling     

Traveling electrical waves in cortex: insights from phase dynamics and speculation on a computational role

The theory of coupled phase oscillators provides a framework to understand the emergent properties of networks of neuronal oscillators. When the architecture of the network is dominated by short-range connections, the pattern of electrical output is predicted to correspond to traveling plane and rotating waves, in addition to synchronized output. We argue that this theory provides the foundation for understanding the traveling electrical waves that are observed across olfactory, visual, and visuomotor areas of cortex in a variety of species. The waves are typically present during periods outside of stimulation, while synchronous activity typically dominates in the presence of a strong stimulus. We suggest that the continuum of phase shifts during epochs with traveling waves provides a means to scan the incoming sensory stream for novel features. Experiments to test our theoretical approach are presented.     

Stimulus motion propels traveling waves in binocular rivalry

State transitions in the nervous system often take shape as traveling waves, whereby one neural state is replaced by another across space in a wave-like manner. In visual perception, transitions between the two mutually exclusive percepts that alternate when the two eyes view conflicting stimuli (binocular rivalry) may also take shape as traveling waves. The properties of these waves point to a neural substrate of binocular rivalry alternations that have the hallmark signs of lower cortical areas. In a series of experiments, we show a potent interaction between traveling waves in binocular rivalry and stimulus motion. The course of the traveling wave is biased in the motion direction of the suppressed stimulus that gains dominance by means of the wave-like transition. Thus, stimulus motion may propel the traveling wave across the stimulus to the extent that the stimulus motion dictates the traveling wave's direction completely. Using a computational model, we show that a speed-dependent asymmetry in lateral inhibitory connections between retinotopically organized and motion-sensitive neurons can explain our results. We argue that such a change in suppressive connections may play a vital role in the resolution of dynamic occlusion situations.     

A Discrete Velocity Kinetic Model with Food Metric: Chemotaxis Traveling Waves

We introduce a mesoscopic scale chemotaxis model for traveling wave phenomena which is induced by food metric. The organisms of this simplified kinetic model have two discrete velocity modes, \(\pm s\) and a constant tumbling rate. The main feature of the model is that the speed of organisms is constant \(s\,{>}\,0\) with respect to the food metric, not the Euclidean metric. The uniqueness and the existence of the traveling wave solution of the model are obtained. Unlike the classical logarithmic model case there exist traveling waves under super-linear consumption rates and infinite population pulse-type traveling waves are obtained. Numerical simulations are also provided.     

The effects of the size and shape of landscape features on the formation of traveling waves in cyclic populations

Recent field data indicate that in a number of cyclic populations, the cycles are organized spatially with the form of a periodic traveling wave. One way in which this type of wave is generated is when dispersing individuals encounter landscape features that impede movement in certain directions. In this article, we investigate the dependence of such periodic waves on ecological parameters and on the form of the landscape feature. Using a standard predator-prey model as a prototype for a cyclic population, we calculate the speed and amplitude of waves generated by a large landscape feature. This enables us to determine parameters for which the waves are stable; in other cases, they evolve into irregular oscillations. We then undertake for the first time a detailed study of the effects of the size and shape of a landscape feature on the waves that it generates. We show that size rather than shape is the key wave-forming property, with smaller obstacles generating waves with longer wavelength and waves from larger landscape features dominating those from smaller ones. Our results suggest that periodic traveling waves may be much more common than has previously been assumed in real ecological systems, and they enable quantitative predictions on the properties of these waves for particular cases.     

Learning poly-synaptic paths with traveling waves

Traveling waves are commonly observed across the brain. While previous studies have suggested the role of traveling waves in learning, the mechanism remains unclear. We adopted a computational approach to investigate the effect of traveling waves on synaptic plasticity. Our results indicate that traveling waves facilitate the learning of poly-synaptic network paths when combined with a reward-dependent local synaptic plasticity rule. We also demonstrate that traveling waves expedite finding the shortest paths and learning nonlinear input/output mapping, such as exclusive or (XOR) function.     

Are buffers boring? Uniqueness and asymptotical stability of traveling wave fronts in the buffered bistable system

Traveling waves of calcium are widely observed under the condition that the free cytosolic calcium is buffered. Thus it is of physiological interest to determine how buffers affect the properties of calcium waves. Here we summarise and extend previous results on the existence, uniqueness and stability of traveling wave solutions of the buffered bistable equation, which is the simplest possible model of the upstroke of a calcium wave. Taken together, the results show that immobile buffers do not change the existence, uniqueness or stability of the traveling wave, while mobile buffers can eliminate a traveling wave. However, if a wave exists in the latter case, it remains unique and stable.      

Calcium-Actin Waves and Oscillations of Cellular Membranes

We propose a mechanism for the formation of membrane oscillations and traveling waves, which arise due to the coupling between the actin cytoskeleton and the calcium flux through the membrane. In our model, the fluid cell membrane has a mobile but constant population of proteins with a convex spontaneous curvature, which act as nucleators of actin polymerization and adhesion. Such a continuum model couples the forces of cell-substrate adhesion, actin polymerization, membrane curvature, and the flux of calcium through the membrane. Linear stability analysis shows that sufficiently strong coupling among the calcium, membrane, and protein dynamics may induce robust traveling waves on the membrane. This result was checked for a reduced feedback scheme and is compared to the results without the effects of calcium, where permanent phase separation without waves or oscillations is obtained. The model results are compared to the published observations of calcium waves in cell membranes, and a number of testable predictions are proposed.     

Spreading disease: integro-differential equations old and new

We investigate an integro-differential equation for a disease spread by the dispersal of infectious individuals and compare this to Mollison's [Adv. Appl. Probab. 4 (1972) 233; D. Mollison, The rate of spatial propagation of simple epidemics, in: Proc. 6th Berkeley Symp. on Math. Statist. and Prob., vol. 3, University of California Press, Berkeley, 1972, p. 579; J. R. Statist. Soc. B 39 (3) (1977) 283] model of a disease spread by non-local contacts. For symmetric kernels with moment generating functions, spreading infectives leads to faster traveling waves for low rates of transmission, but to slower traveling waves for high rates of transmission. We approximate the shape of the traveling waves for the two models using both piecewise linearization and a regular-perturbation scheme.     

Hypothesis on acoustic receptive fields

A hypothesis of acoustic receptive fields is studied, which is based on the fact that the cochlea of the internal ear is a wave guide with traveling waves and the resonance in the critical layer. When a harmonic sound influences the ear, the traveling wave reaches the critical layer for the corresponding frequency and generates there a train of decaying waves about 25 periods in duration, which form a steep slope of the envelope. The funnel-shaped convergence of all neurones innervating the acoustic receptors of the Corti organ along the slope of the envelope gives rise to acoustic receptive fields. The hypothesis is consistent with some other experimental data. Such an acoustic receptive field makes it possible to use the whole train of waves in the critical layer to measure the frequency of the acting sinusoidal sound with the greatest possible accuracy. Similarly, a high accuracy of recognition of short-time sound pulses is provided, which could not be explained earlier.     

Traveling waves and pulses in a one-dimensional network of excitable integrate-and-fire neurons

 We study the existence and stability of traveling waves and pulses in a one-dimensional network of integrate-and-fire neurons with synaptic coupling. This provides a simple model of excitable neural tissue. We first derive a self-consistency condition for the existence of traveling waves, which generates a dispersion relation between velocity and wavelength. We use this to investigate how wave-propagation depends on various parameters that characterize neuronal interactions such as synaptic and axonal delays, and the passive membrane properties of dendritic cables. We also establish that excitable networks support the propagation of solitary pulses in the long-wavelength limit. We then derive a general condition for the (local) asymptotic stability of traveling waves in terms of the characteristic equation of the linearized firing time map, which takes the form of an integro-difference equation of infinite order. We use this to analyze the stability of solitary pulses in the long-wavelength limit. Solitary wave solutions are shown to come in pairs with the faster (slower) solution stable (unstable) in the case of zero axonal delays; for non-zero delays and fast synapses the stable wave can itself destabilize via a Hopf bifurcation. Received: 27 October 1998     

Traveling waves in the prefrontal cortex during working memory

Neural oscillations are evident across cortex but their spatial structure is not well- explored. Are oscillations stationary or do they form “traveling waves”, i.e., spatially organized patterns whose peaks and troughs move sequentially across cortex? Here, we show that oscillations in the prefrontal cortex (PFC) organized as traveling waves in the theta (4-8Hz), alpha (8-12Hz) and beta (12-30Hz) bands. Some traveling waves were planar but most rotated. The waves were modulated during performance of a working memory task. During baseline conditions, waves flowed bidirectionally along a specific axis of orientation. Waves in different frequency bands could travel in different directions. During task performance, there was an increase in waves in one direction over the other, especially in the beta band.     

Mathematical modeling of Erk activity waves in regenerating zebrafish scales

Erk signaling regulates cellular decisions in many biological contexts. Recently, we have reported a series of Erk activity traveling waves that coordinate regeneration of osteoblast tissue in zebrafish scales. These waves originate from a central source region, propagate as expanding rings, and impart cell growth, thus controlling tissue morphogenesis. Here, we present a minimal reaction-diffusion model for Erk activity waves. The model considers three components: Erk, a diffusible Erk activator, and an Erk inhibitor. Erk stimulates both its activator and inhibitor, forming a positive and negative feedback loop, respectively. Our model shows that this system can be excitable and propagate Erk activity waves. Waves originate from a pulsatile source that is modeled by adding a localized basal production of the activator, which turns the source region from an excitable to an oscillatory state. As Erk activity periodically rises in the source, it can trigger an excitable wave that travels across the entire tissue. Analysis of the model finds that positive feedback controls the properties of the traveling wavefront and that negative feedback controls the duration of Erk activity peak and the period of Erk activity waves. The geometrical properties of the waves facilitate constraints on the effective diffusivity of the activator, indicating that waves are an efficient mechanism to transfer growth factor signaling rapidly across a large tissue.     

Formation and Dynamics of Waves in a Cortical Model of Cholinergic Modulation

Acetylcholine (ACh) is a regulator of neural excitability and one of the neurochemical substrates of sleep. Amongst the cellular effects induced by cholinergic modulation are a reduction in spike-frequency adaptation (SFA) and a shift in the phase response curve (PRC). We demonstrate in a biophysical model how changes in neural excitability and network structure interact to create three distinct functional regimes: localized asynchronous, traveling asynchronous, and traveling synchronous. Our results qualitatively match those observed experimentally. Cortical activity during slow wave sleep (SWS) differs from that during REM sleep or waking states. During SWS there are traveling patterns of activity in the cortex; in other states stationary patterns occur. Our model is a network composed of Hodgkin-Huxley type neurons with a M-current regulated by ACh. Regulation of ACh level can account for dynamical changes between functional regimes. Reduction of the magnitude of this current recreates the reduction in SFA the shift from a type 2 to a type 1 PRC observed in the presence of ACh. When SFA is minimal (in waking or REM sleep state, high ACh) patterns of activity are localized and easily pinned by network inhomogeneities. When SFA is present (decreasing ACh), traveling waves of activity naturally arise. A further decrease in ACh leads to a high degree of synchrony within traveling waves. We also show that the level of ACh determines how sensitive network activity is to synaptic heterogeneity. These regimes may have a profound functional significance as stationary patterns may play a role in the proper encoding of external input as memory and traveling waves could lead to synaptic regularization, giving unique insights into the role and significance of ACh in determining patterns of cortical activity and functional differences arising from the patterns.     

The equation of motion for sperm flagella.

The equation of motion for sperm flagella, in which the elastic bending moment and the active contractile moment are balanced by the moment from the viscous resistance of the surrounding fluid, is solved for a wave solution that superimposes partial solutions. Substitution of the expression for the wave solution into the equation leads to an expression for the active contractile moment. This active moment can be decomposed into two parts. The first part describes an active moment that travels over the flagellum with the mechanical flagellar wave, the second part represents a moment in phase over the entire length of the flagellum, which decreases linearly towards the distal tip. The linear synchronous moment, to which an amount of traveling moment has been added as a perturbation, leads to wave solutions that closely resemble flagellar waves. Properties such as wavelength and wave amplitudes and also the shape of the waves in sea urchin sperm flagella at different frequencies are accurately described by the theory. The change in wave shape in sea urchin sperm flagella at raised viscosity is predicted well by the theory. The different wave properties caused in bull sperm flagella by different boundary conditions at the proximal junction are explained. When only a traveling active moment is present in a flagellum, the wave solutions describe waves of a small wave length in a long flagellum. Some properties of the wave motion of sperm flagella are derived from the theory and verified experimentally.     

Modeling of glycolytic wave propagation in an open spatial reactor with inhomogeneous substrate influx

The spatio-temporal dynamics of traveling waves in glycolysis as it occurs in yeast extract have been studied, both theoretically and experimentally. We describe this phenomenon with the distributed Selkov model that accounts for the reactions of phosphofructokinase, which is a key enzyme of the glycolytic reaction cascade. To describe the experimentally observed phase waves in an open spatial reactor we introduce a non-homogeneous flux of substrate in the model. The experimental observation that waves can change their direction of propagation during the experiment is considered in the model. The mechanism for such a change in wave direction is discussed.     

Traveling wave solutions of a reaction diffusion model for competing pioneer and climax species

Presented is a reaction-diffusion model for the interaction of pioneer and climax species. For certain parameters the system exhibits bistability and traveling wave solutions. Specifically, we show that when the climax species diffuses at a slow rate there are traveling wave solutions which correspond to extinction waves of either the pioneer or climax species. A leading order analysis is used in the one-dimensional spatial case to estimate the wave speed sign that determines which species becomes extinct. Results of these analyses are then compared to numerical simulations of wave front propagation for the model on one and two-dimensional spatial domains. A simple mechanism for harvesting is also introduced.     

Existence of traveling waves for integral recursions with nonmonotone growth functions

A class of integral recursion models for the growth and spread of a synchronized single-species population is studied. It is well known that if there is no overcompensation in the fecundity function, the recursion has an asymptotic spreading speed c*, and that this speed can be characterized as the speed of the slowest non-constant traveling wave solution. A class of integral recursions with overcompensation which still have asymptotic spreading speeds can be found by using the ideas introduced by Thieme (J Reine Angew Math 306:94–121, 1979) for the study of space-time integral equation models for epidemics. The present work gives a large subclass of these models with overcompensation for which the spreading speed can still be characterized as the slowest speed of a non-constant traveling wave. To illustrate our results, we numerically simulate a series of traveling waves. The simulations indicate that, depending on the properties of the fecundity function, the tails of the waves may approach the carrying capacity monotonically, may approach the carrying capacity in an oscillatory manner, or may oscillate continually about the carrying capacity, with its values bounded above and below by computable positive numbers. B. Li’s research was partially supported by the National Science Foundation under Grant DMS-616445. M. A. Lewis research was supported by “The Canada Research Chairs program,” and a grant from the Natural Sciences and Engineering Research Council of Canada.     

Plant Dynamics,Birth-Jump Processes,and Sharp Traveling Waves

Motivated by the importance of understanding the dynamics of the growth and dispersal of plants in various environments, we introduce and analyze a discrete agent-based model based on a birth-jump process, which exhibit wave-like solutions. To rigorously analyze these traveling wave phenomena, we derive the diffusion limit of the discrete model and prove the existence of traveling wave solutions (sharp and continuously differentiable) assuming a logarithmic-type growth. Furthermore, we provide a variational speed for the minimum speed of the waves and perform numerical experiments that confirm our results.