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Animals use stereo sampling of odor concentration to localize sources and follow odor trails. We analyze the dynamics of a bilateral model that depends on the simultaneous comparison between odor concentrations detected by left and right sensors. The general model consists of three differential equations for the positions in the plane and the heading. When the odor landscape is an infinite trail, then we reduce the dynamics to a planar system whose dynamics have just two fixed points. Using an integrable approximation (for short sensors) we estimate the basin of attraction. In the case of a radially symmetric landscape, we again can reduce the dynamics to a planar system, but the behavior is considerably richer with multi-stability, isolas, and limit cycles. As in the linear trail case, there is also an underlying integrable system when the sensors are short. In odor landscapes that consist of multiple spots and trail segments, we find periodic and chaotic dynamics and characterize the behavior on trails with gaps and that turn corners.

We review the theory of weakly coupled oscillators for smooth systems. We then examine situations where application of the standard theory falls short and illustrate how it can be extended. Specific examples are given to non-smooth systems with applications to the Izhikevich neuron. We then introduce the idea of isostable reduction to explore behaviors that the weak coupling paradigm cannot explain. In an additional example, we show how bifurcations that change the stability of phase locked solutions in a pair of identical coupled neurons can be understood using the notion of isostable reduction., Comment: 23 pages, 5 figures, review

Cross-frequency coupling (CFC) is thought to play an important role in communication across distant brain regions. However, neither the mechanism of its generation nor the influence on the underlying spiking dynamics is well understood. Here, we investigate the dynamics of two interacting distant neuronal modules coupled by inter-regional long-range connections. Each neuronal module comprises an excitatory and inhibitory population of quadratic integrate-and-fire neurons connected locally with conductance-based synapses. The two modules are coupled reciprocally with delays that represent the long-range conduction time. We applied the Ott-Antonsen ansatz to reduce the spiking dynamics to the corresponding mean field equations as a small set of delay differential equations. Bifurcation analysis on these mean field equations shows inter-regional conduction delay is sufficient to produce CFC via a torus bifurcation. Spike correlation analysis during the CFC revealed that several local clusters exhibit synchronized firing in gamma-band frequencies. These clusters exhibit locally decorrelated firings between the cluster pairs within the same population. In contrast, the clusters exhibit long-range gamma-band cross-covariance between the distant clusters that have similar firing frequency. The interactions of the different gamma frequencies produce a beat leading to population-level CFC. We analyzed spike counts in relation to the phases of the macroscopic fast and slow oscillations and found population spike counts vary with respect to macroscopic phases. Such firing phase preference accompanies a phase window with high spike count and low Fano factor, which is suitable for a population rate code. Our work suggests the inter-regional conduction delay plays a significant role in the emergence of CFC and the underlying spiking dynamics may support long-range communication and neural coding., Comment: 13 pages, 4 figures

In this paper, we study pairs of oscillators that are indirectly coupled via active (excitable) cells. We introduce a scalar phase model for coupled oscillators and excitable cells. We first show that one excitable and one oscillatory cell will exhibit phase locking at a variety of $m:n$ patterns. We next introduce a second oscillatory cell and show that the only attractor is synchrony between the oscillators. We will also study the robustness to heterogeneity when excitable cell fires or is quiescent. We next examine the dynamics when the oscillators are coupled via two excitable cells. In this case, the dynamics are very complicated with many forms of bistability and, in some cases, chaotic behavior. We also apply weak coupling analysis to this case and explain some of the degeneracies observed in the bifurcation diagram. We also look at pairs of oscillators coupled via long chains of excitable cells and show that small differences in the frequency of the oscillators makes their locking more robust. Finally, we demonstrate many of the same phenomena seen in the phase model for a gap-junction coupled system of Morris-Lecar neurons., Comment: 11 pages, 13 figures, accepted for publication in PRE

We extend the theory of weakly coupled oscillators to incorporate slowly varying inputs and parameters. We employ a combination of regular perturbation and an adiabatic approximation to derive equations for the phase-difference between a pair of oscillators. We apply this to the simple Hopf oscillator and then to a biophysical model. The latter represents the behavior of a neuron that is subject to slow modulation of a muscarinic current such as would occur during transient attention through cholinergic activation. Our method extends and simplifies the recent work of Kurebayashi (Physical Review Letters, 111, 214101, 2013) to include coupling. We apply the method to an all-to-all network and show that there is a waxing and waning of synchrony of modulated neurons., Comment: 28 pages, 11 figures

We consider the existence of non-synchronized fixed points to the Kuramoto model defined on sparse networks: specifically, networks where each vertex has degree exactly three. We show that "most" such networks support multiple attracting phase-locked solutions that are not synchronized, and study the depth and width of the basins of attraction of these phase-locked solutions. We also show that it is common in "large enough" graphs to find phase-locked solutions where one or more of the links has angle difference greater than $\pi/2$.

In this paper we provide two representations for stochastic ion channel kinetics, and compare the performance of exact simulation with a commonly used numerical approximation strategy. The first representation we present is a random time change representation, popularized by Thomas Kurtz, with the second being analogous to a "Gillespie" representation. Exact stochastic algorithms are provided for the different representations, which are preferable to either (a) fixed time step or (b) piecewise constant propensity algorithms, which still appear in the literature. As examples, we provide versions of the exact algorithms for the Morris-Lecar conductance based model, and detail the error induced, both in a weak and a strong sense, by the use of approximate algorithms on this model. We include ready-to-use implementations of the random time change algorithm in both XPP and Matlab. Finally, through the consideration of parametric sensitivity analysis, we show how the representations presented here are useful in the development of further computational methods. The general representations and simulation strategies provided here are known in other parts of the sciences, but less so in the present setting., Comment: 39 pages, 6 figures, appendix with XPP and Matlab code

A neural correlate of parametric working memory is a stimulus specific rise in neuron firing rate that persists long after the stimulus is removed. Network models with local excitation and broad inhibition support persistent neural activity, linking network architecture and parametric working memory. Cortical neurons receive noisy input fluctuations which causes persistent activity to diffusively wander about the network, degrading memory over time. We explore how cortical architecture that supports parametric working memory affects the diffusion of persistent neural activity. Studying both a spiking network and a simplified potential well model, we show that spatially heterogeneous excitatory coupling stabilizes a discrete number of persistent states, reducing the diffusion of persistent activity over the network. However, heterogeneous coupling also coarse-grains the stimulus representation space, limiting the capacity of parametric working memory. The storage errors due to coarse-graining and diffusion tradeoff so that information transfer between the initial and recalled stimulus is optimized at a fixed network heterogeneity. For sufficiently long delay times, the optimal number of attractors is less than the number of possible stimuli, suggesting that memory networks can under-represent stimulus space to optimize performance. Our results clearly demonstrate the effects of network architecture and stochastic fluctuations on parametric memory storage., Comment: 42 pages, 9 figures

We study the effects of noise on stationary pulse solutions (bumps) in spatially extended neural fields. The dynamics of a neural field is described by an integrodifferential equation whose integral term characterizes synaptic interactions between neurons in different spatial locations of the network. Translationally symmetric neural fields support a continuum of stationary bump solutions, which may be centered at any spatial location. Random fluctuations are introduced by modeling the system as a spatially extended Langevin equation whose noise term we take to be multiplicative or additive. For nonzero noise, these bumps are shown to wander about the domain in a purely diffusive way. We can approximate the effective diffusion coefficient using a small noise expansion. Upon breaking the (continuous) translation symmetry of the system using a spatially heterogeneous inputs or synapses, bumps in the stochastic neural field can become temporarily pinned to a finite number of locations in the network. In the case of spatially heterogeneous synaptic weights, as the modulation frequency of this heterogeneity increases, the effective diffusion of bumps in the network approaches that of the network with spatially homogeneous weights., Comment: 40 pages, 11 figures

Using network models consisting of gap junction coupled Wang-Buszaki neurons, we demonstrate that it is possible to obtain not only synchronous activity between neurons but also a variety of constant phase shifts between 0 and \pi. We call these phase shifts intermediate stable phaselocked states. These phase shifts can produce a large variety of wave-like activity patterns in one-dimensional chains and two-dimensional arrays of neurons, which can be studied by reducing the system of equations to a phase model. The 2\pi periodic coupling functions of these models are characterized by prominent higher order terms in their Fourier expansion, which can be varied by changing model parameters. We study how the relative contribution of the odd and even terms affect what solutions are possible, the basin of attraction of those solutions and their stability. These models may be applicable to the spinal central pattern generators of the dogfish and also to the developing neocortex of the neonatal rat.

In the absence of synaptic coupling, two or more neural oscillators may become synchronized by virtue of the statistical correlations in their noisy input streams. Recent work has shown that the degree of correlation transfer from input currents to output spikes depends not only on intrinsic oscillator dynamics, but also depends on the length of the observation window over which the correlation is calculated. In this paper we use stochastic phase reduction and regular perturbations to derive the correlation of the total phase elapsed over long time scales, a quantity which provides a convenient proxy for the spike count correlation. Over short time scales, we derive the spike count correlation directly using straightforward probabilistic reasoning applied to the density of the phase difference. Our approximations show that output correlation scales with the autocorrelation of the phase resetting curve over long time scales. We also find a concise expression for the influence of the shape of the phase resetting curve on the initial slope of the output correlation over short time scales. These analytic results together with numerical simulations provide new intuitions for the recent counterintuitive finding that type I oscillators transfer correlations more faithfully than do type II over long time scales, while the reverse holds true for the better understood case of short time scales., Comment: 9 pages, 7 figures, submitted to Physical Review E

We review a recent approach to the mean-field limits in neural networks that takes into account the stochastic nature of input current and the uncertainty in synaptic coupling. This approach was proved to be a rigorous limit of the network equations in a general setting, and we express here the results in a more customary and simpler framework. We propose a heuristic argument to derive these equations providing a more intuitive understanding of their origin. These equations are characterized by a strong coupling between the different moments of the solutions. We analyse the equations, present an algorithm to simulate the solutions of these mean-field equations, and investigate numerically the equations. In particular, we build a bridge between these equations and Sompolinsky and collaborators approach (1988, 1990), and show how the coupling between the mean and the covariance function deviates from customary approaches.

The phase-resetting curve (PRC) describes the response of a neural oscillator to small perturbations in membrane potential. Its usefulness for predicting the dynamics of weakly coupled deterministic networks has been well characterized. However, the inputs to real neurons may often be more accurately described as barrages of synaptic noise. Effective connectivity between cells may thus arise in the form of correlations between the noisy input streams. We use constrained optimization and perturbation methods to prove that PRC shape determines susceptibility to synchrony among otherwise uncoupled noise-driven neural oscillators. PRCs can be placed into two general categories: Type I PRCs are non-negative while Type II PRCs have a large negative region. Here we show that oscillators with Type II PRCs receiving common noisy input sychronize more readily than those with Type I PRCs., Comment: 10 pages, 4 figures, submitted to Physical Review E

Many real oscillators are coupled to other oscillators and the coupling can affect the response of the oscillators to stimuli. We investigate phase response curves (PRCs) of coupled oscillators. The PRCs for two weakly coupled phase-locked oscillators are analytically obtained in terms of the PRC for uncoupled oscillators and the coupling function of the system. Through simulation and analytic methods, the PRCs for globally coupled oscillators are also discussed., Comment: 5 pages 4 figure

We investigate coupled identical phase oscillators with scale-free distribution of coupling strength. It is shown that partially locked states can occur due to the inhomogeneity in coupling and some properties of the coupling function. Various quantities of the partially locked states are computed through a self-consistency argument and the values show good agreement with simulation results., Comment: 4 figures, 7 pages