Bifurcation mechanisms involved in the generation of action potentials (spikes) by neurons are reviewed here. We show how the type of bifurcation determines the neuro-computational properties of the cells. For example, when the rest state is near a saddle-node bifurcation, the cell can fire all-or-none spikes with an arbitrary low frequency, it has a well-defined threshold manifold, and it acts as an integrator; i.e. the higher the frequency of incoming pulses, the sooner it fires. In contrast, when the rest state is near an Andronov–Hopf bifurcation, the cell fires in a certain frequency range, its spikes are not all-or-none, it does not have a well-defined threshold manifold, it can fire in response to an inhibitory pulse, and it acts as a resonator; i.e. it responds preferentially to a certain (resonant) frequency of the input. Increasing the input frequency may actually delay or terminate its firing. We also describe the phenomenon of neural bursting, and we use geometric bifurcation theory to extend the existing classification of bursters, including many new types. We discuss how the type of burster defines its neuro-computational properties, and we show that different bursters can interact, synchronize and process information differently.

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Neural excitability, spiking and bursting

A.S. Perelson, D.E. Kirschner and R. De Boer (Math. Biosci. 114 (1993) 81) proposed an ODE model of cell-free viral spread of human immunodeficiency virus (HIV) in a well-mixed compartment such as the bloodstream. Their model consists of four components: uninfected healthy CD4(+) T-cells, latently infected CD4(+) T-cells, actively infected CD4(+) T-cells, and free virus. This model has been important in the field of mathematical modeling of HIV infection and many other models have been proposed which take the model of Perelson, Kirschner and De Boer as their inspiration, so to speak (see a recent survey paper by A.S. Perelson and P.W. Nelson (SIAM Rev. 41 (1999) 3-44)). We first simplify their model into one consisting of only three components: the healthy CD4(+) T-cells, infected CD4(+) T-cells, and free virus and discuss the existence and stability of the infected steady state. Then, we introduce a discrete time delay to the model to describe the time between infection of a CD4(+) T-cell and the emission of viral particles on a cellular level (see A.V.M. Herz, S. Bonhoeffer, R.M. Anderson, R.M. May, M.A. Nowak [Proc. Nat. Acad. Sci. USA 93 (1996) 7247]). We study the effect of the time delay on the stability of the endemically infected equilibrium, criteria are given to ensure that the infected equilibrium is asymptotically stable for all delay. Numerical simulations are presented to illustrate the results.

A delay-differential equation model of HIV infection of CD4(+) T-cells.

We consider quasistatic motion and stability of a single degree of freedom elastic system undergoing frictional slip. The system is represented by a block (slider) slipping at speed V and connected by a spring of stiffness k to a point at which motion is enforced at speed V 0 We adopt rate and state dependent frictional constitutive relations for the slider which describe approximately experimental results of Dieterich and Ruina over a range of slip speeds V . In the simplest relation the friction stress depends additively on a term A In V and a state variable θ; the state variable θ evolves, with a characteristic slip distance, to the value − B In V , where the constants A, B are assumed to satisfy B > A > 0. Limited results are presented based on a similar friction law using two state variables. Linearized stability analysis predicts constant slip rate motion at V 0 to change from stable to unstable with a decrease in the spring stiffness k below a critical value k cr . At neutral stability oscillations in slip rate are predicted. A nonlinear analysis of slip motions given here uses the Hopf bifurcation technique, direct determination of phase plane trajectories, Liapunov methods and numerical integration of the equations of motion. Small but finite amplitude limit cycles exist for one value of k , if one state variable is used. With two state variables oscillations exist for a small range of k which undergo period doubling and then lead to apparently chaotic motions as k is decreased. Perturbations from steady sliding are imposed by step changes in the imposed load point motion. Three cases are considered: (1) the load point speed V 0 is suddenly increased; (2) the load point is stopped for some time and then moved again at a constant rate; and (3) the load point displacement suddenly jumps and then stops. In all cases, for all values of k :, sufficiently large perturbations lead to instability. Primary conclusions are: (1) ‘stick-slip’ instability is possible in systems for which steady sliding is stable, and (2) physical manifestation of quasistatic oscillations is sensitive to material properties, stiffness, and the nature and magnitude of load perturbations.

http://esag.harvard.edu/rice/111_GuRiRuinaTse_RateStateStab_JMPS84.pdf

Slip motion and stability of a single degree of freedom elastic system with rate and state dependent friction

The original Hilbert's 16th problem can be split into four parts consisting of Problems A–D. In this paper, the progress of study on Hilbert's 16th problem is presented, and the relationship between Hilbert's 16th problem and bifurcations of planar vector fields is discussed. The material is presented in eight sections. Section 1: Introduction: what is Hilbert's 16th problem? Section 2: The first part of Hilbert's 16th problem. Section 3: The second part of Hilbert's 16th problem: introduction. Section 4: Focal values, saddle values and finite cyclicity in a fine focus, closed orbit and homoclinic loop. Section 5: Finiteness problem. Section 6: The weakened Hilbert's 16th problem. Section 7: Global and local bifurcations of Zq–equivariant vector fields. Section 8: The rate of growth of Hilbert number H(n) with n.

Hilbert's 16th problem and bifurcations of planar polynomial vector fields

Spiral waves in diverse excitable media exhibit strikingly variegated behavior. Mechanistic interpretations of excitability in laboratory systems are commonly tested by comparing the wavelength, period, and meander patterns of the model’s spiral waves with laboratory observations, but models seem seldom to be rejected by such tests. The reason may be that almost any excitable medium behaves in many respects like almost any other, if its parameters are properly adjusted within a reasonable range. What generalizations can be made about ‘‘excitable media’’ in the absence of more specifications? It would be useful to distinguish such generic features from idiosyncrasies of specific models. The range of behavioral flexibility of the FitzHugh–Nagumo excitable medium is explored by varying two of its parameters and comparing the results with other excitable media to suggest a generic pattern of parameter dependence. The results exhibit the remarkable diversity of rotor behavior in a single model and provide a database for quantitative testing of mathematical generalizations.

Varieties of spiral wave behavior: An experimentalist's approach to the theory of excitable media.

1. The Hopf Bifurcation Theorum 2. Applications: Ordinary Differential Equations (by hand) 3. Numerical Evaluation of Hopf Bifurcation Formulae 4. Applications: Differential-Difference and Integro-differential Equations (by hand) 5. Applications: Partial Differential Equations (by hand).

Nicholas D. Kazarinoff

Papers

Theory and applications of Hopf bifurcation