On oscillatory responses of the Limulus retina

Published observations of the dynamic properties of lateral and self-inhibition in the Limulus retina lead to a non-linear integral equation for the response of ommatidia located near the center of a uniformly illuminated region. Coleman and Renninger (1976, 1978) showed that when the excitation is constant in time and the sum of the inhibitory coefficients for the illuminated region exceeds a critical value, the integral equation has a stable periodic solution describing a sustained, spatially synchronized, oscillatory response in which bursts of activity alternate with silent periods. Such spatially synchronized bursting has been observed in the Limulus retina in situ by Barlow and Fraioli (1978), using the preparation of Barlow and Kaplan (1971). Employing experimental data on the temporal dependence of lateral and self-inhibition, which were then available only for the excised eye, Coleman and Renninger calculated a value of 0.34 s for the period p of the bursting response, which is significantly above the range, 0.11–0.20 s, of values of p observed for the Limulus eye in situ. Brodie et al. (1978) have recently published measurements of the temporal dependence of lateral and self-inhibition for the in situ preparation. Here we show that when the kernel functions in Coleman and Renninger's integral equation are chosen in accord with these new data, the periodic solutions of the equation have a period of approximately 0.13s, which is in the range (0.11–0.20 s) required for agreement with experiment. Other properties of the periodic solutions, i.e., their general form and the threshold levels of inhibition required for their existence, are also in accord with published observations of the behavior of the retina in situ.     

Approximate expressions of the bifurcating periodic solutions in a neuron model with delay-dependent parameters by perturbation approach

This paper is interested in gaining insights of approximate expressions of the bifurcating periodic solutions in a neuron model. This model shares the property of involving delay-dependent parameters. The presence of such dependence requires the use of suitable criteria which usually makes the analytical work so harder. Most existing methods for studying the nonlinear dynamics fail when applied to such a class of delay models. Although Xu et al. (Phys Lett A 354:126–136, 2006) studied stability switches, Hopf bifurcation and chaos of the neuron model with delay-dependent parameters, the dynamics of this model are still largely undetermined. In this paper, a detailed analysis on approximation to the bifurcating periodic solutions is given by means of the perturbation approach. Moreover, some examples are provided for comparing approximations with numerical solutions of the bifurcating periodic solutions. It shows that the dynamics of the neuron model with delay-dependent parameters is quite different from that of systems with delay-independent parameters only.     

Periodic solutions of population models in a periodically fluctuating environment.

A periodically fluctuating environment is assumed in a population-modeling process that generates nonautonomous difference equations. The existence and uniqueness of periodic solutions are studied. A sufficient condition for existence and a necessary condition for uniqueness are obtained. Stability of the periodic solutions is investigated. Several numerical examples are given to illustrate the basic results, and a brief discussion is presented.     

Degenerate Hopf bifurcation and isolated periodic solutions of the Hodgkin-Huxley model with varying sodium ion concentration

Points of degenerate Hopf bifurcation in the Hodgkin-Huxley model are found as parameters temperature T and voltage level of sodium VNa are varied. Local techniques of degenerate Hopf bifurcation analysis are used to show the existence of families of periodic solutions of the model: isolated branches of periodic solutions (i.e. branches not connected to the stationary branch) are found in addition to Hopf branches. Purely numerical techniques are used to show that the isolas persist for VNa up to a value slightly greater than 114 mV. Under some conditions there are multiple stable periodic solutions, so "jumping" between action potentials of different amplitudes might be observed.     

Generation of the nervous impulse and periodic oscillations

Usually the models for the excitation and propagation of the nervous impulse are studied either in the space-clamp situation or on a model axon extended on both sides to infinity. Following Fitzhugh in the present paper the release of an impulse train at the axon hillock is studied within the scope of Fitzhugh's BVP model. The existence and stability of periodic oscillations are studied by direct methods, also the relation to Liénard's equation. The exact correspondence between the BVP model and the socalled Nagumo-equation is established. For typical examples the solutions are computed by numerical methods.     

Synaptic regulation of the state of excitable membranes

Hodgkin-Huxly equations accounting for (stationary) control signals (synaptic conductivity and current) have stationary and periodic solutions. The domain of unstability of stationary solutions determined on the control signals plane (excitation and inhibition). On one part of the domain boundary a decrease of excitation and increase of inhibition suppress oscillations. On another part of the boundary they induced oscillations, which were not explained previously in accordance with the experimental data.     

Bifurcation of periodic solutions of Hodgkin-Huxley model for the squid giant axon

The Hodgkin-Huxley model of the space-clamped squid giant axon is shown to admit unstable periodic solutions for current stimuli less than the stimulus at which the rest state becomes linearly unstable. The periodic solutions are demonstrated both by bifurcation theory and by numerical integration. The presence of subcritical unstable oscillations explains the discontinuous behaviour of the amplitude of the repetitive response as a function of current stimulus     

Cyclic AMP oscillations in suspensions of Dictyostelium discoideum

A model developed previously for signal relay and adaptation in the cellular slime mould Dictyostelium discoideum is shown to account for the observed oscillations of calcium and cyclic AMP in cellular suspensions. A qualitative argument is given which explains how the oscillations arise, and numerical computations show how characteristics such as the period and amplitude of the periodic solutions depend on parameters in the model. Several extensions of the basic model are investigated, including the effect of cell aggregation and the effect of time delays in the activation and adaptation processes. The dynamics of mixed cell populations in which only a small fraction of the cells are capable of autonomous oscillation are also studied.     

具有三时滞Lotka-Volterra互惠系统的Hopf分支

建立了具有三个时滞的Lotka-Volterra互惠系统;获得了正平衡点和Hopf分支存在的条件等;并对所获得的结果进行了数值模拟．     

Consequences of delayed lateral inhibition in the retina of Limulus. I. Elementary theory of spatially uniform fields.

Although the magnitude of lateral inhibition in the retina of the compound eye of Limulus polyphemus depends strongly on the distance between ommatidia, the delay time τ between the response of one ommatidium and the consequent inhibition of another is independent of the distance between them and is approximately 0·1 sec (cf., e.g., Ratliff, Knight, Dodge &; Hartline,1973). We have recently shown (Coleman &; Renninger, 1974) that under appropriate circumstances the response r of such a retina to a temporally constant and spatially uniform excitation of amount e should be a succession of “bursts” and “rest periods”, each of duration τ, so that r, although spatially uniform, is a periodic function of time with period 2τ and mean value $\text{1}\text{2}\text{ϵ}$. Further, the periodic function r can have a fine structure in which there is repeated information about the duration and sequence, but not the amplitude, of any short pulses which occurred immediately (i.e. within a time interval of length τ) before the onset of steady uniform excitation. We derive here implications of several plausible general hypotheses about the functional form of lateral inhibition. We show that, under those of the considered hypotheses which are expected to hold for Limulus in strong light, periodic bursting behavior for r is stable and is rapidly approached if the excitation is held steady and uniform.     

Two species competition in a periodic environment

The classical Lotka-Volterra equations for two competing species have constant coefficients. In this paper these equations are studied under the assumption that the coefficients are periodic functions of a common period. As a generalization of the existence theory for equilibria in the constant coefficient case, it is shown that there exists a branch of positive periodic solutions which connects (i.e. bifurcates from) the two nontrivial periodic solutions lying on the coordinate axes. This branch exists for a finite interval or spectrum of bifurcation parameter values (the bifurcation parameter being the average of the net inherent growth rate of one species). The stability of these periodic solutions is studied and is related to the theory of competitive exclusion. A specific example of independent ecological interest is examined by means of which it is shown under what circumstances two species, which could not coexist in a constant environment, can coexist in a limit cycle fashion when subjected to suitable periodic harvesting or removal rates.Research supported by National Science Foundation Grant No. MCS-7901307     

Global dynamics of an SIRS epidemic model with saturation incidence

In this paper, the dynamical behavior of an SIRS epidemic model with birth pulse, pulse vaccination, and saturation incidence is studied. By using a discrete map, the existence and stability of the infection-free periodic solution and the endemic periodic solution are investigated. The conditions required for the existence of supercritical bifurcation are derived. A threshold for a disease to be extinct or endemic is established. The Poincaré map and center manifold theorem are used to discuss flip bifurcation of the endemic periodic solution. Moreover, numerical simulations for bifurcation diagrams, phase portraits and periodic solutions, which are illustrated with an example, are in good agreement with the theoretical analysis.     

An approximate solution to the periodic bidomain equations in one dimension

An approximate, computationally tractable solution is proposed for the potentials in the bidomain model with periodic intracellular junctions (the periodic bidomain model). This new approach is based on the one-dimensional rigorous spectral method described previously by Trayanova and Pilkington (IEEE Trans. Biomed. Eng., May 1993). The total solution to the one-dimensional periodic bidomain problem is decomposed in the spectral domain into solutions to (1) the single-fiber classical bidomain problem in which the intracellular conductivity value incorporates the average contribution from cytoplasm and junction and (2) the “junctional” potential problem due to the presence of junctions at discrete locations alone. Solving for the junctional term rigorously requires most of the numerical effort in the solution for the periodic bidomain potentials. Here the junctional potential is found approximately with little numerical effort. A comparison between the rigorous and the approximate solutions serves as a justification for the proposed approximate solution procedure. The procedure outlined in this paper is applicable to higher spatial dimensions where both tissue anisotropy and junctional inhomogeneities play a role in establishing the transmembrane potential distribution.     

Theoretical and experimental investigations of the excitation of high-frequency oscillations in an ion collective accelerator model based on the Doppler effect

Results are presented from studies of a two-beam scheme of ion acceleration by a high-frequency field excited by an electron beam due to the instabilities associated with anomalous and normal Doppler effects. The dynamics of the excitation of eigenmodes in a periodic slow wave structure (SWS) by a relativistic electron beam via the anomalous Doppler effect is investigated theoretically. Mechanisms for the saturation of the instability are considered, analytical expressions for the maximum field amplitude and the efficiency with which the energy of beam electrons is converted into the energy of the excited wave are derived, and the results of numerical simulations of such excitation are presented. An experimental stand designed to test the principles and possibility of proton acceleration up to an energy of 8 MeV at a current up to 3 A is described. A double resonance (associated with anomalous and normal Doppler effects) occurring in the interaction of an electron beam with a helical SWS is studied experimentally. In this case, an increase in the efficiency with which the accelerating high-frequency field is excited is observed.     

Numerical bifurcation analysis of delay differential equations arising from physiological modeling

This paper has a dual purpose. First, we describe numerical methods for continuation and bifurcation analysis of steady state solutions and periodic solutions of systems of delay differential equations with an arbitrary number of fixed, discrete delays. Second, we demonstrate how these methods can be used to obtain insight into complex biological regulatory systems in which interactions occur with time delays: for this, we consider a system of two equations for the plasma glucose and insulin concentrations in a diabetic patient subject to a system of external assistance. The model has two delays: the technological delay of the external system, and the physiological delay of the patient's liver. We compute stability of the steady state solution as a function of two parameters, compare with analytical results and compute several branches of periodic solutions and their stability. These numerical results allow to infer two categories of diabetic patients for which the external system has different efficiency.     

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     

Periodic metabolic systems: Oscillations in multiple-loop negative feedback biochemical control networks

Summary For a general multiple loop feedback inhibition system in which the end product can inhibit any or all of the intermediate reactions it is shown that biologically significant behaviour is always confined to a bounded region of reaction space containing a unique equilibrium. By explicit construction of a Liapunov function for the general n dimensional differential equation it is shown that some values of reaction parameters cause the concentration vector to approach the equilibrium asymptotically for all physically realizable initial conditions. As the parameter values change, periodic solutions can appear within the bounded region. Some information about these periodic solutions can be obtained from the Hopf bifurcation theorem. Alternatively, if specific parameter values are known a numerical method can be used to find periodic solutions and determine their stability by locating a zero of the displacement map. The single loop Goodwin oscillator is analysed in detail. The methods are then used to treat an oscillator with two feedback loops and it is found that oscillations are possible even if both Hill coefficients are equal to one.     

A five-link 2D brachiating ape model with life-like zero-energy-cost motions

We have found periodic life-like brachiating motions of a rigid-body ape model that use no muscle or gravitational energy to move steadily forward. The most complicated of these models has 5 links (a body and two arms, each with 2 links) and 7 degrees of freedom in flight. The defining feature of all our periodic solutions is that all collisions are at zero relative velocity. These motions are found using numerical integration and root-finding that is sufficiently precise so as to imply that the solutions found correspond to mathematical solutions with exactly zero energy cost. The only actuation and control in the model is for maintaining contact with and releasing handholds which requires no mechanical work. The similarity of these energy-free simulations to the motions of apes suggests that muscle-use minimization at least partially characterizes the coordination strategies they use.     

The effect of the lateral reticular nucleus of the medulla oblongata on the cerebellar cortex

It was established as a result of a laminar analysis of evoked potentials (EP) in the paramedian lobe of the cerebellum of unanesthetized cats that in response to stimulation of the lateral reticular nucleus (LRN) excitation of granulosa cells develops which is not accompanied by excitation of Purkinje cells. Destruction of the lateral nucleus leads to a considerable decrease in the "diffuse" component of the EP which develop during stimulation of the somatic nerves. The results obtained show that the afferent pathway whose fibers innervate granulosa cells of a special type, which have been previously described, passes through the LRN. These cells have extensive receptive fields and do not directly affect the activity of Purkinje cells. A hypothesis is proposed on the basis of the results obtained and data in the literature that the extensive receptive fields of the LRN cells and of the granulosa cells innervated by them are due to interneuronal connections within the lateral nucleus.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Laboratory on the Use of Mathematical Methods in Biology, Moscow University. Translated from Neirofiziologiya, Vol. 2, No. 6, pp. 581–586, November–December, 1970.     

State-dependent impulsive models of integrated pest management (IPM) strategies and their dynamic consequences

A state-dependent impulsive model is proposed for integrated pest management (IPM). IPM involves combining biological, mechanical, and chemical tactics to reduce pest numbers to tolerable levels after a pest population has reached its economic threshold (ET). The complete expression of an orbitally asymptotically stable periodic solution to the model with a maximum value no larger than the given ET is presented, the existence of which implies that pests can be controlled at or below their ET levels. We also prove that there is no periodic solution with order larger than or equal to three, except for one special case, by using the properties of the LambertW function and Poincare map. Moreover, we show that the existence of an order two periodic solution implies the existence of an order one periodic solution. Various positive invariant sets and attractors of this impulsive semi-dynamical system are described and discussed. In particular, several horseshoe-like attractors, whose interiors can simultaneously contain stable order 1 periodic solutions and order 2 periodic solutions, are found and the interior structure of the horseshoe-like attractors is discussed. Finally, the largest invariant set and the sufficient conditions which guarantee the global orbital and asymptotic stability of the order 1 periodic solution in the meaningful domain for the system are given using the Lyapunov function. Our results show that, in theory, a pest can be controlled such that its population size is no larger than its ET by applying effects impulsively once, twice, or at most, a finite number of times, or according to a periodic regime. Moreover, our theoretical work suggests how IPM strategies could be used to alter the levels of the ET in the farmers' favour.