Emergence and combinatorial accumulation of jittering regimes in spiking oscillators with delayed feedback

Interaction via pulses is common in many natural systems, especially neuronal. In this article we study one of the simplest possible systems with pulse interaction: a phase oscillator with delayed pulsatile feedback. When the oscillator reaches a specific state, it emits a pulse, which returns after propagating through a delay line. The impact of an incoming pulse is described by the oscillator's phase reset curve (PRC). In such a system we discover an unexpected phenomenon: for a sufficiently steep slope of the PRC, a periodic regular spiking solution bifurcates with several multipliers crossing the unit circle at the same parameter value. The number of such critical multipliers increases linearly with the delay and thus may be arbitrary large. This bifurcation is accompanied by the emergence of numerous "jittering" regimes with non-equal interspike intervals (ISIs). Each of these regimes corresponds to a periodic solution of the system with a period roughly proportional to the delay. The number of different "jittering" solutions emerging at the bifurcation point increases exponentially with the delay. We describe the combinatorial mechanism that underlies the emergence of such a variety of solutions. In particular, we show how a periodic solution exhibiting several distinct ISIs can imply the existence of multiple other solutions obtained by rearranging of these ISIs. We show that the theoretical results for phase oscillators accurately predict the behavior of an experimentally implemented electronic oscillator with pulsatile feedback

Phase models and clustering in networks of oscillators with delayed coupling

We consider a general model for a network of oscillators with time delayed, circulant coupling. We use the theory of weakly coupled oscillators to reduce the system of delay differential equations to a phase model where the time delay enters as a phase shift. We use the phase model to study the existence and stability of cluster solutions. Cluster solutions are phase locked solutions where the oscillators separate into groups. Oscillators within a group are synchronized while those in different groups are phase-locked. We give model independent existence and stability results for symmetric cluster solutions. We show that the presence of the time delay can lead to the coexistence of multiple stable clustering solutions. We apply our analytical results to a network of Morris Lecar neurons and compare these results with numerical continuation and simulation studies

Synchronization of coupled neural oscillators with heterogeneous delays

We investigate the effects of heterogeneous delays in the coupling of two excitable neural systems. Depending upon the coupling strengths and the time delays in the mutual and self-coupling, the compound system exhibits different types of synchronized oscillations of variable period. We analyze this synchronization based on the interplay of the different time delays and support the numerical results by analytical findings. In addition, we elaborate on bursting-like dynamics with two competing timescales on the basis of the autocorrelation function.Comment: 18 pages, 14 figure

Emergence of global synchronization in directed excitatory networks of type I neurons

The collective behaviour of neural networks depends on the cellular and synaptic properties of the neurons. The phase-response curve (PRC) is an experimentally obtainable measure of cellular properties that quantifies the shift in the next spike time of a neuron as a function of the phase at which stimulus is delivered to that neuron. The neuronal PRCs can be classified as having either purely positive values (type I) or distinct positive and negative regions (type II). Networks of type 1 PRCs tend not to synchronize via mutual excitatory synaptic connections. We study the synchronization properties of identical type I and type II neurons, assuming unidirectional synapses. Performing the linear stability analysis and the numerical simulation of the extended Kuramoto model, we show that feedforward loop motifs favour synchronization of type I excitatory and inhibitory neurons, while feedback loop motifs destroy their synchronization tendency. Moreover, large directed networks, either without feedback motifs or with many of them, have been constructed from the same undirected backbones, and a high synchronization level is observed for directed acyclic graphs with type I neurons. It has been shown that, the synchronizability of type I neurons depends on both the directionality of the network connectivity and the topology of its undirected backbone. The abundance of feedforward motifs enhances the synchronizability of the directed acyclic graphs

Mathematical frameworks for oscillatory network dynamics in neuroscience

The tools of weakly coupled phase oscillator theory have had a profound impact on the neuroscience community, providing insight into a variety of network behaviours ranging from central pattern generation to synchronisation, as well as predicting novel network states such as chimeras. However, there are many instances where this theory is expected to break down, say in the presence of strong coupling, or must be carefully interpreted, as in the presence of stochastic forcing. There are also surprises in the dynamical complexity of the attractors that can robustly appear—for example, heteroclinic network attractors. In this review we present a set of mathemat- ical tools that are suitable for addressing the dynamics of oscillatory neural networks, broadening from a standard phase oscillator perspective to provide a practical frame- work for further successful applications of mathematics to understanding network dynamics in neuroscience

Behavior control in the sensorimotor loop with short-term synaptic dynamics induced by self-regulating neurons

The behavior and skills of living systems depend on the distributed control provided by specialized and highly recurrent neural networks. Learning and memory in these systems is mediated by a set of adaptation mechanisms, known collectively as neuronal plasticity. Translating principles of recurrent neural control and plasticity to artificial agents has seen major strides, but is usually hampered by the complex interactions between the agent's body and its environment. One of the important standing issues is for the agent to support multiple stable states of behavior, so that its behavioral repertoire matches the requirements imposed by these interactions. The agent also must have the capacity to switch between these states in time scales that are comparable to those by which sensory stimulation varies. Achieving this requires a mechanism of short-term memory that allows the neurocontroller to keep track of the recent history of its input, which finds its biological counterpart in short-term synaptic plasticity. This issue is approached here by deriving synaptic dynamics in recurrent neural networks. Neurons are introduced as self-regulating units with a rich repertoire of dynamics. They exhibit homeostatic properties for certain parameter domains, which result in a set of stable states and the required short-term memory. They can also operate as oscillators, which allow them to surpass the level of activity imposed by their homeostatic operation conditions. Neural systems endowed with the derived synaptic dynamics can be utilized for the neural behavior control of autonomous mobile agents. The resulting behavior depends also on the underlying network structure, which is either engineered or developed by evolutionary techniques. The effectiveness of these self-regulating units is demonstrated by controlling locomotion of a hexapod with 18 degrees of freedom, and obstacle-avoidance of a wheel-driven robot. © 2014 Toutounji and Pasemann

Short Conduction Delays Cause Inhibition Rather than Excitation to Favor Synchrony in Hybrid Neuronal Networks of the Entorhinal Cortex

How stable synchrony in neuronal networks is sustained in the presence of conduction delays is an open question. The Dynamic Clamp was used to measure phase resetting curves (PRCs) for entorhinal cortical cells, and then to construct networks of two such neurons. PRCs were in general Type I (all advances or all delays) or weakly type II with a small region at early phases with the opposite type of resetting. We used previously developed theoretical methods based on PRCs under the assumption of pulsatile coupling to predict the delays that synchronize these hybrid circuits. For excitatory coupling, synchrony was predicted and observed only with no delay and for delays greater than half a network period that cause each neuron to receive an input late in its firing cycle and almost immediately fire an action potential. Synchronization for these long delays was surprisingly tight and robust to the noise and heterogeneity inherent in a biological system. In contrast to excitatory coupling, inhibitory coupling led to antiphase for no delay, very short delays and delays close to a network period, but to near-synchrony for a wide range of relatively short delays. PRC-based methods show that conduction delays can stabilize synchrony in several ways, including neutralizing a discontinuity introduced by strong inhibition, favoring synchrony in the case of noisy bistability, and avoiding an initial destabilizing region of a weakly type II PRC. PRCs can identify optimal conduction delays favoring synchronization at a given frequency, and also predict robustness to noise and heterogeneity