Model of the early development of thalamo-cortical connections and area patterning via signaling molecules

The mammalian cortex is divided into architectonic and functionally distinct areas. There is growing experimental evidence that their emergence and development is controlled by both epigenetic and genetic factors. The latter were recently implicated as dominating the early cortical area specification. In this paper, we present a theoretical model that explicitly considers the genetic factors and that is able to explain several sets of experiments on cortical area regulation involving transcription factors Emx2 and Pax6, and fibroblast growth factor FGF8. The model consists of the dynamics of thalamo- cortical connections modulated by signaling molecules that are regulated genetically, and by axonal competition for neocortical space. The model can make predictions and provides a basic mathematical framework for the early development of the thalamo-cortical connections and area patterning that can be further refined as more experimental facts become known.Comment: brain, model, neural development, cortical area patterning, signaling molecule

Formation of antiwaves in gap-junction-coupled chains of neurons

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

Chimera States for Coupled Oscillators

Arrays of identical oscillators can display a remarkable spatiotemporal pattern in which phase-locked oscillators coexist with drifting ones. Discovered two years ago, such "chimera states" are believed to be impossible for locally or globally coupled systems; they are peculiar to the intermediate case of nonlocal coupling. Here we present an exact solution for this state, for a ring of phase oscillators coupled by a cosine kernel. We show that the stable chimera state bifurcates from a spatially modulated drift state, and dies in a saddle-node bifurcation with an unstable chimera.Comment: 4 pages, 4 figure

Phase Response Curves of Coupled Oscillators

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 figur

Limits and dynamics of stochastic neuronal networks with random heterogeneous delays

Realistic networks display heterogeneous transmission delays. We analyze here the limits of large stochastic multi-populations networks with stochastic coupling and random interconnection delays. We show that depending on the nature of the delays distributions, a quenched or averaged propagation of chaos takes place in these networks, and that the network equations converge towards a delayed McKean-Vlasov equation with distributed delays. Our approach is mostly fitted to neuroscience applications. We instantiate in particular a classical neuronal model, the Wilson and Cowan system, and show that the obtained limit equations have Gaussian solutions whose mean and standard deviation satisfy a closed set of coupled delay differential equations in which the distribution of delays and the noise levels appear as parameters. This allows to uncover precisely the effects of noise, delays and coupling on the dynamics of such heterogeneous networks, in particular their role in the emergence of synchronized oscillations. We show in several examples that not only the averaged delay, but also the dispersion, govern the dynamics of such networks.Comment: Corrected misprint (useless stopping time) in proof of Lemma 1 and clarified a regularity hypothesis (remark 1

Effective phase description of noise-perturbed and noise-induced oscillations

An effective description of a general class of stochastic phase oscillators is presented. For this, the effective phase velocity is defined either by invariant probability density or via first passage times. While the first approach exhibits correct frequency and distribution density, the second one yields proper phase resetting curves. Their discrepancy is most pronounced for noise-induced oscillations and is related to non-monotonicity of the phase fluctuations

Synchronization Transition in the Kuramoto Model with Colored Noise

We present a linear stability analysis of the incoherent state in a system of globally coupled, identical phase oscillators subject to colored noise. In that we succeed to bridge the extreme time scales between the formerly studied and analytically solvable cases of white noise and quenched random frequencies.Comment: 4 pages, 2 figure

Collective synchronization in spatially extended systems of coupled oscillators with random frequencies

We study collective behavior of locally coupled limit-cycle oscillators with random intrinsic frequencies, spatially extended over $d$-dimensional hypercubic lattices. Phase synchronization as well as frequency entrainment are explored analytically in the linear (strong-coupling) regime and numerically in the nonlinear (weak-coupling) regime. Our analysis shows that the oscillator phases are always desynchronized up to $d=4$, which implies the lower critical dimension $d_{l}^{P}=4$ for phase synchronization. On the other hand, the oscillators behave collectively in frequency (phase velocity) even in three dimensions ($d=3$), indicating that the lower critical dimension for frequency entrainment is $d_{l}^{F}=2$. Nonlinear effects due to periodic nature of limit-cycle oscillators are found to become significant in the weak-coupling regime: So-called {\em runaway oscillators} destroy the synchronized (ordered) phase and there emerges a fully random (disordered) phase. Critical behavior near the synchronization transition into the fully random phase is unveiled via numerical investigation. Collective behavior of globally-coupled oscillators is also examined and compared with that of locally coupled oscillators.Comment: 18 pages, 18 figure

Correlations, fluctuations and stability of a finite-size network of coupled oscillators

The incoherent state of the Kuramoto model of coupled oscillators exhibits marginal modes in mean field theory. We demonstrate that corrections due to finite size effects render these modes stable in the subcritical case, i.e. when the population is not synchronous. This demonstration is facilitated by the construction of a non-equilibrium statistical field theoretic formulation of a generic model of coupled oscillators. This theory is consistent with previous results. In the all-to-all case, the fluctuations in this theory are due completely to finite size corrections, which can be calculated in an expansion in 1/N, where N is the number of oscillators. The N -> infinity limit of this theory is what is traditionally called mean field theory for the Kuramoto model.Comment: 25 pages (2 column), 12 figures, modifications for resubmissio

Synchronization of Excitatory Neurons with Strongly Heterogeneous Phase Responses

In many real-world oscillator systems, the phase response curves are highly heterogeneous. However, dynamics of heterogeneous oscillator networks has not been seriously addressed. We propose a theoretical framework to analyze such a system by dealing explicitly with the heterogeneous phase response curves. We develop a novel method to solve the self-consistent equations for order parameters by using formal complex-valued phase variables, and apply our theory to networks of in vitro cortical neurons. We find a novel state transition that is not observed in previous oscillator network models.Comment: 4 pages, 3 figure