Coupling of a slow and a fast oscillator can generate bursting

A general mechanism underlying bursting is proposed and described. It consists of two coupled nonlinear oscillators with different frequencies, where the slower oscillator alternatively switches the faster one on and off. This mechanism is shown to work in an extended Bonhoefer-van der Pol oscillator as well as in a modified version of the Hodgkin-Huxley equations.     

A Relaxation Oscillator Description of the Burst-Generating Mechanism in the Cardiac Ganglion of the Lobster,Homarus americanus

Properties of the neural mechanism responsible for generating the periodic burst of spike potentials in the nine ganglion neurons were investigated by applying brief, single shocks to the four small cells with extracellular electrodes placed near the trigger zones of the small cells. The shock elicited a burst if presented during the latter portion of the silent period, terminated a burst during the latter portion of the burst period, and was followed by a newly initiated burst during the early portion of the burst period. The resultant changes in burst and silent period durations were quantitatively described by a second-order non-linear differential equation similar to the van der Pol equation for a relaxation oscillator. The equation also qualitatively described changes in firing threshold of the small cells during the burst cycle. The first derivative of the solution to the equation is similar to slow transmembrane potentials in neurons that are involved in generation of burst activity in other crustacean cardiac ganglia.     

Mode analysis of a tubular structure of coupled non-linear oscillators for digestive-tract modelling

A commonly accepted mathematical model for the slow-wave electrical activity of the gastro-intestinal tract of humans and animals comprises a set of interconnected nonlinear oscillators. Using a van der Pol oscillator with third-power conductance characteristics as the unit oscillator a number of structures have been analysed using a matrix Krylov-Bogolioubov method linearisation. The mode analysis of one-dimensional chains and two-dimensional arrays has been reported. In this paper the method has been extended to consider a tubular structure which is relevant to modelling small-intestinal rhythms. It is shown that this structure is capable of producing stable single models, non-resonant double modes and degenerated modes. General expressions are obtained for anm×n structure and examples given of two special conditions of 3×4 (i.e. odd numbers of oscillators in a ring) and 4×3 cases. The analytical results obtained for these two cases have been vertified experimentally using an electronic implementation of coupled van der Pol oscillators. Results obtained using fifth-power non-linear oscillators are summarised.     

A simpler model of the human circadian pacemaker

Numerous studies have used the classic van der Pol oscillator, which contains a cubic nonlinearity, to model the effect of light on the human circadian pacemaker. Jewett and Kronauer demonstrated that Aschoff's rule could be incorporated into van der Pol type models and used a van der Pol type oscillator with higher order nonlinearities. Kronauer, Forger, and Jewett have proposed a model for light preprocessing, Process L, representing a biochemical process that converts a light signal into an effective drive on the circadian pacemaker. In the paper presented here, the authors use the classic van der Pol oscillator with Process L and Jewett and Kronauer's model of Aschoff's rule to model the human circadian pacemaker. This simpler cubic model predicts the results of a three-pulse human phase response curve experiment and a two-pulse amplitude reduction study with as much, or more, accuracy as the models of Jewett and Kronauer and Kronauer, Forger, and Jewett, which both employ a nonlinearity of degree 7. This suggests that this simpler cubic model should be considered as a potential alternative to other models of the human circadian system currently available.     

A statistical model of the human core-temperature circadian rhythm

We formulate a statistical model of the human core-temperature circadian rhythm in which the circadian signal is modeled as a van der Pol oscillator, the thermoregulatory response is represented as a first-order autoregressive process, and the evoked effect of activity is modeled with a function specific for each circadian protocol. The new model directly links differential equation-based simulation models and harmonic regression analysis methods and permits statistical analysis of both static and dynamical properties of the circadian pacemaker from experimental data. We estimate the model parameters by using numerically efficient maximum likelihood algorithms and analyze human core-temperature data from forced desynchrony, free-run, and constant-routine protocols. By representing explicitly the dynamical effects of ambient light input to the human circadian pacemaker, the new model can estimate with high precision the correct intrinsic period of this oscillator ( approximately 24 h) from both free-run and forced desynchrony studies. Although the van der Pol model approximates well the dynamical features of the circadian pacemaker, the optimal dynamical model of the human biological clock may have a harmonic structure different from that of the van der Pol oscillator.     

The spike generation zone of the ampullary electroreceptor

Sensory input to the central nervous system begins with a transduction step, specialized to the sensory modality involved, resulting in the production of postsynaptic electrical input to the outermost branches of a dendritic tree. Spatiotemporal summation of this slow input as it converges upon the axon then initiates the production of or modulates the rate of ongoing production of fast neural spikes destined for the central nervous system. We present a novel circuit design consisting of an operational amplifier, a tunnel diode and linear passive components, intended to model the spike generation zone at which the transformation of neural input from slow to fast format takes place. Our circuit is shown to be a relaxation oscillator of the van der Pol type. Simulated postsynaptic current modulates the frequency of spike production by the relaxation oscillator model, producing a stimulus-response characteristic which can be compared with those observed in vivo. Stimulus-response data for our model match data available in the literature for the ampullary electroreceptor of elasmobranch fish.     

A model for the speed of memory retrieval

The memory retrieval process of number problems with external noise is studied with the use of the Bonhoeffer–van der Pol oscillator model. Three cell assembly responses are simulated, coding one true number and two neighboring erroneous. The time of a correct response, T _c, was averaged over statistical assemblies of numerous trials. It is demonstrated that T _c takes a minimum value for a certain noise intensity. This result correlates well with experimental data by Usher and Feingold (2000). The location of the minimum as a function of the time delay between two consecutive simulation trials is investigated.     

The method of harmonic balance applied to coupled asymmetrical van der Pol oscillators for intestinal modelling

The spontaneous electrical rhythms recorded from the gastro-intestinal tract of humans and animals have been successfully modelled by an array of interconnected van der Pol oscillators. To account for asymmetry in the recorded waveforms (with particular reference to the human small intestine) an additional term in the van der Pol dynamics has been included. It is shown that the method of harmonic balance can be used to give analytical results for this asymmetrical condition. The non-linear algebraic equations are solved by hill-climbing to give values of d.c., fundamental and second harmonic amplitudes together with the entrained frequency. The results correlate well with actual measurements made on an analogue simulation by three different methods for waveshape factors of 0.1 and 1.0     

Unfolding an electronic integrate-and-fire circuit

Many physical and biological phenomena involve accumulation and discharge processes that can occur on significantly different time scales. Models of these processes have contributed to understand excitability self-sustained oscillations and synchronization in arrays of oscillators. Integrate-and-fire (I+F) models are popular minimal fill-and-flush mathematical models. They are used in neuroscience to study spiking and phase locking in single neuron membranes, large scale neural networks, and in a variety of applications in physics and electrical engineering. We show here how the classical first-order I+F model fits into the theory of nonlinear oscillators of van der Pol type by demonstrating that a particular second-order oscillator having small parameters converges in a singular perturbation limit to the I+F model. In this sense, our study provides a novel unfolding of such models and it identifies a constructible electronic circuit that is closely related to I+F.     

Analysis of limit-cycle conditions in intercoupled relay oscillators with reference to gastrointestinal modelling

This paper presents an exact analysis of a mutually coupled relay oscillator based on a method orignated by Tsypkin. Limit-cycle frequencies and phases can be determined exactly using this method, unlike other approximate methods based on describing functions and harmonic balance techniques. A new method of exact determination of limit-cycle stability is also shown to give excellent agreement with simulation studies. Different types of intercoupling are shown to give different stability conditions, and these are discussed in relation to gastrointestinal (GI) smooth muscle modelling. GI tract electrical activity has previously been modelled using bidirectionally coupled nonlinear oscillators, and the results of the present analysis of relay oscillators is compared with other studies using van der Pol dynamics.     

A novel approach to the van der pol oscillator: Natural frequency and entrainment

Abstract

The physics of the van der Pol oscillator as realized by the Meissner circuit is discussed by analogy to the beat phenomenon and by a consequent analysis of current balance. The current balance method leads to a new, very accurate equation for the dependence of the oscillator frequency on the feedback parameter. Several aspects of entrainment (existence, limited frequency range, dependence on parameters, phase shift) can be explained, too. Numerical results are presented which have been obtained by solving the homogeneous and inhomogeneous van der Pol equation with a Runge‐Kutta method.     

Spatiotemporal analysis of simultaneous single-unit activity in the dragonfly

The present work describes a new technique for the identification of functional connectivity between neural firing patterns. The simultaneous singleunit recordings obtained from over 50 individual cells in the dragonfly mesothoracic ganglion during three consecutive behavioral states: pre-flight, flight and postflight were evaluated. Each individual spike train was converted into a synthesized analog gradient designed to capture crucial physiological characteristics of the cell from which the spike train emanated. Estimates of network functional connectivity were calculated using correlations between analog gradient spike trains for all possible cell pairings. Both functional excitation and inhibition could be detected in the correlations. The detection of functional connectivity was relatively independent of cell firing rate. More detailed analyses indicated the existence of cellular firing histories and connectivity patterns during flight that strongly resembled the characteristics of a bi-stable oscillator. Such an oscillator, hypothetically, could drive the elevator and depressor motor neuron firing paterns that support wing kinematics. There was no evidence for the functional existence of such an oscillator within either preor post-flight spike records. The detected spatiotemporal patterns of neural activity are hypothesized to be consistent with neural command sequences that the dragonfly might use to control flight. The demonstrated capability to define short-time scale functional relationships between spike trains obtained from dragonfly ganglia should have valuable applications to the comparative study of neural information processing strategies in a variety of other neural systems.     

Frequency and burst duration in oscillating neurons and two-cell networks

We study the relationship of injected current to oscillator period in single neurons and two-cell model networks formed by reciprocal inhibitory synapses. Using a Morris-Lecar-like model, we identify two qualitative types of oscillatory behavior for single model neurons. The classical oscillator behavior is defined as type A. Here the burst duration is relatively constant and the frequency increases with depolarization. For oscillator type B, the frequency first increases and then decreases when depolarized, due to the variable burst duration. Our simulations show that relatively modest changes in the maximal inward and outward conductances can move the oscillator from one type to another. Cultured stomatogastric ganglion neurons exhibit both A and B type behaviors and can switch between the two types with pharmacological manipulation. Our simulations indicate that the stability of a two-cell network with injected current can be extended with inhibitory coupling. In addition, two-cell networks formed from type A or type B oscillators behave differently from each other at lower synaptic strengths.     

Zum Mechanismus der biologischen 24-Stunden-Periodik

Summary The level (=arithmetic average of all instantaneous values)of a self-sustained oscillation in general influences all properties of the oscillation, including period, amplitude and shape of the oscillation, and the rate of exchange of energy between the oscillator and its environment. Only when the non-linear damping factor does not depend on the instantaneous value of the oscillating function, but only on the amplitude of the oscillation, are the other properties independent of the average level. The differential equations describing self-sustained oscillations cannot be solved exactly, but methods of approximation are applicable. Numerical solutions to several different forms of the equations will be discussed.In the simplest case (van der Pol equation) all properties of the self-sustained oscillation (e.g. period, amplitude) are extreme when the level is zero. The oscillation continues only within a given range of levels (oscillating range); outside this range, the oscillation damps out. In other modifications of the equation, the oscillating function cannot assume a zero value. In all cases, the extent to which the average level influences the different properties depends on the factor , which describes the position of the oscillation within the range between harmonic and relaxation types of oscillation.In the elementary van der Pol equation, the correlation between level and frequency changes sign within the oscillating range; that is, the circadian rule, demanding an always positive correlation between level and frequency, cannot be fulfilled. Only with an additional non-linearity in the energy of recoil does the correlation remain unchanged in sign throughout the oscillating range. A stability condition demands a positive sign for this non-linearity, and hence, for the correlation (fulfilling the circadian rule); if the sign is negative (violating the circadian rule), the oscillation becomes unstable. With an additional term of the third order, the oscillation acquires a two-peaked shape typical of many circadian oscillations.A simple differential equation describing all general properties of the circadian periodicity must fulfil these conditions: the oscillation must be self-sustained and limited to positive values; and the energy of recoil must be non-linear with a positive coefficient to obtain the appropriate correlation between level and frequency. In the equations here developed the environment directly influences only one parameter of the oscillation, i.e. the level. In addition to the circadian periodicity, the differential equations here examined describe the behavior of several other biological oscillations.

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Die benutzten mathematischen Begriffe folgen — soweit dort angeführt — den Benennungen des DIN-Blattes 1311; im Anhang I sind die wichtigsten Begriffe noch einmal zusammenfassend definiert.     

Periodic and non-periodic responses of a periodically forced Hodgkin-Huxley oscillator

Membrane potential responses of a Hodgkin-Huxley oscillator to an externally-applied sinusoidal current were numerically calculated with relation to bifurcation parameters of the amplitude and the frequency of the stimulating current. The Hodgkin-Huxley oscillator, or the Hodgkin-Huxley axon in the state of self-sustained oscillation of action potentials, was realized by immersing the axon in calcium-deficient sea water. The forced oscillations were analysed by the stroboscopic plots and/or the Lorenz plots. The results show that the periodically forced Hodgkin-Huxley oscillator exhibits not only periodic motions (harmonic or sub-harmonic synchronization) but also non-periodic motions (quasi-periodic or chaotic oscillation), that the motions were determined by the amplitude and the frequency of the stimulating current, and that the characteristic motions obtained in the present study were in reasonable agreement with those of our previous results, found experimentally in squid giant axons. Also, two kinds of routes to the chaotic oscillations were found; successive period-doubling bifurcations and formation of the intermittently chaotic oscillation from sub-harmonic synchronization.     

Coupled Van der Pol oscillators — A model of excitatory and inhibitory neural interactions

A system of mutually coupled Van der Pol equations is derived from an extended version of the Wilson and Cowan model for the dynamics of a number of excitatory and inhibitory neural subsets. In the lowest order of approximation, interactions between excitatory and inhibitory subsets appear as linear elastic coupling, while those within and between excitatory and excitatory subsets appear as nonlinear frictional coupling. The case of two coupled oscillators is investigated by the method of averaging and the stability conditions for two mode oscillations are obtained. Internal resonance is also discussed briefly in the case of identical oscillators.     

A van der Pol coupled-oscillator model as a basis for developing a system for suppressing MHD instabilities in a tokamak

The main parameters of tokamak discharges are known to be limited by large-scale MHD instabilities. Sometimes, the instabilities lead to a rapid (on time scales of tens of microseconds) disruption of the discharge current and to the release of all the energy stored in the plasma column at the discharge chamber wall. This process, which is called the disruptive instability, may have irreversible catastrophic consequences for the operation of a fusion reactor. In the present paper, a study is made of the dynamics of self-oscillations in systems of two and six van der Pol coupled oscillators. A van der Pol coupled-oscillator model is used to develop a multivariable feedback controller based on the combined principle of compensating for internal cross feedbacks within the object and introducing damping feedbacks in each control channel. By using mathematical simulation methods, it is shown that the controller designed guarantees the suppression of self-oscillations in a system of van der Pol oscillators over a fairly broad range of parameters of the object under control (and thereby provides the structural stability of the object). The nonlinear control system model makes it possible to suppress coupled MHD perturbations developing in a tokamak plasma.     

Neural rate equations for bursting dynamics derived from conductance-based equations

A method of obtaining rate equations from conductance-based equations is developed and applied to fast-spiking and bursting neocortical neurons. It involves splitting systems of conductance-based equations into fast and slow subsystems, and averaging the effects of fast terms that drive the slowly varying quantities by showing that their average is closely proportional to the firing rate. The dependence of the firing rate on the injected current is then approximated in the analysis. The resulting behavior of the slow variables is then substituted back into the fast equations, with the further approximation of replacing the fast voltages in these terms by effective values. For bursting neurons the method yields two coupled limit-cycle oscillators: a self-exciting oscillator for the slow variables that commences limit-cycle oscillations at a critical current and modulates a fast spike-generating oscillator, thereby leading to slowly modulated bursts with a group of spikes in each burst. The dynamics of these coupled oscillators are then verified against those of the conductance-based equations. Finally, it is shown how to place the results in a form suitable for use in mean-field equations for neural population dynamics.     

Coupled oscillators utilised as gait rhythm generators of a two-legged walking machine

The gait of current two-legged walking machines differs from that of humans, although the kinematic structures of these machines' legs frequently imitate human limbs. This paper presents a method of generating the trajectories of hip and knee joint angles resulting in a gait pattern similar to that of a human. For this purpose the solutions of coupled van der Pol oscillator equations are utilised. There is much evidence that these equations can be treated as a good model of the central pattern generator generating functional (also locomotional) rhythms in living creatures. The oscillator equations are solved by numerical integration. The method of changing the type of gait by changing appropriate parameter values in the oscillator equations is presented (change of velocity and trajectory of leg-ends). The results obtained enable enhanced control of twolegged walking systems by including gait pattern generators which will assume a similar role to that of biological generators.