Bard Ermentrout

Type i membranes, phase resetting curves, and synchrony

Type I membrane oscillators such as the Connor model (Connor et al. 1977) and the Morris-Lecar model (Morris and Lecar 1981) admit very low frequency oscillations near the critical applied current. Hansel et al. (1995) have numerically shown that synchrony is difficult to achieve with these models and that the phase resetting curve is strictly positive. We use singular perturbation methods and averaging to show that this is a general property of Type I membrane models. We show in a limited sense that so called Type II resetting occurs with models that obtain rhythmicity via a Hopf bifurcation. We also show the differences between synapses that act rapidly and those that act slowly and derive a canonical form for the phase interactions.

Simulation of networks of spiking neurons: A review of tools and strategies

We review different aspects of the simulation of spiking neural networks. We start by reviewing the different types of simulation strategies and algorithms that are currently implemented. We next review the precision of those simulation strategies, in particular in cases where plasticity depends on the exact timing of the spikes. We overview different simulators and simulation environments presently available (restricted to those freely available, open source and documented). For each simulation tool, its advantages and pitfalls are reviewed, with an aim to allow the reader to identify which simulator is appropriate for a given task. Finally, we provide a series of benchmark simulations of different types of networks of spiking neurons, including Hodgkin–Huxley type, integrate-and-fire models, interacting with current-based or conductance-based synapses, using clock-driven or event-driven integration strategies. The same set of models are implemented on the different simulators, and the codes are made available. The ultimate goal of this review is to provide a resource to facilitate identifying the appropriate integration strategy and simulation tool to use for a given modeling problem related to spiking neural networks.

Neural networks as spatio-temporal pattern-forming systems

Models of neural networks are developed from a biological point of view. Small networks are analysed using techniques from dynamical systems. The behaviour of spatially and temporally organized neural fields is then discussed from the point of view of pattern formation. Bifurcation methods, analytic solutions and perturbation methods are applied to these models.

On the human sensorimotor-cortex beta rhythm: sources and modeling.

Cortical oscillations in the beta band (13-35 Hz) are known to be modulated by the GABAergic agonist benzodiazepine. To investigate the mechanisms generating the approximately 20-Hz oscillations in the human cortex, we administered benzodiazepines to healthy adults and monitored cortical oscillatory activity by means of magnetoencephalography. Benzodiazepine increased the power and decreased the frequency of beta oscillations over rolandic areas. Minimum current estimates indicated the effect to take place around the hand area of the primary sensorimotor cortex. Given that previous research has identified sources of the beta rhythm in the motor cortex, our results suggest that these same motor-cortex beta sources are modulated by benzodiazepine. To explore the mechanisms underlying the increase in beta power with GABAergic inhibition, we simulated a conductance-based neuronal network comprising excitatory and inhibitory neurons. The model accounts for the increase in the beta power, the widening of the spectral peak, and the slowing down of the rhythms with benzodiazepines, implemented as an increase in GABAergic conductance. We found that an increase in IPSCs onto inhibitory neurons was more important for generating neuronal synchronization in the beta band than an increase in IPSCs onto excitatory pyramidal cells.

The Effects of Spike Frequency Adaptation and Negative Feedback on the Synchronization of Neural Oscillators

There are several different biophysical mechanisms for spike frequency adaptation observed in recordings from cortical neurons. The two most commonly used in modeling studies are a calcium-dependent potassium current Iahp and a slow voltage-dependent potassium current, Im. We show that both of these have strong effects on the synchronization properties of excitatorily coupled neurons. Furthermore, we show that the reasons for these effects are different. We show through an analysis of some standard models, that the M-current adaptation alters the mechanism for repetitive firing, while the after hyperpolarization adaptation works via shunting the incoming synapses. This latter mechanism applies with a network that has recurrent inhibition. The shunting behavior is captured in a simple two-variable reduced model that arises near certain types of bifurcations. A one-dimensional map is derived from the simplified model.

Linearization of F-1 curves by adaptation

We show that negative feedback to highly nonlinear frequency-current (F-I) curves results in an effective linearization. (By highly nonlinear we mean that the slope at threshold is infinite or very steep.) We then apply this to a specific model for spiking neurons and show that the details of the adaptation mechanism do not affect the results. The crucial points are that the adaptation is slow compared to other processes and the unadapted F-I curve is highly nonlinear.

Synchrony, stability, and firing patterns in pulse-coupled oscillators

We study non-trivial firing patterns in small assemblies of pulse-coupled oscillatory maps. We find conditions for the existence of waves in rings of coupled maps that are coupled bi-directionally. We also find conditions for stable synchrony in general all-to-all coupled oscillators. Surprisingly, we find that for maps that are derived from physiological data, the stability of synchrony depends on the number of oscillators. We describe rotating waves in two-dimensional lattices of maps and reduce their existence to a reduced system of algebraic equations which are solved numerically.

An adaptive model for synchrony in the firefly Pteroptyx malaccae

We describe a new model for synchronization of neuronal oscillators that is based on the observation that certain species of fireflies are able to alter their free-running period. We show that by adding adaptation to standard oscillator models it is possible to observe the frequency alteration. One consequence of this is the perfect synchrony between coupled oscillators. Stability and some analytic results are included along with numerical simulations.

Reduction of conductance-based models with slow synapses to neural nets

The method of averaging and a detailed bifurcation calculation are used to reduce a system of synaptically coupled neurons to a Hopfield type continuous time neural network. Due to some special properties of the bifurcation, explicit averaging is not required and the reduction becomes a simple algebraic problem. The resultant calculations show one how to derive a new type of squashing function whose properties are directly related to the detailed ionic mechanisms of the membrane. Frequency encoding as opposed to amplitude encoding emerges in a natural fashion from the theory. The full system and the reduced system are numerically compared.

Stripes or spots ? Nonlinear effects in bifurcation of reaction-diffusion equations on the square

Bifurcation to spatial patterns in a two-dimensional reaction—diffusion medium is considered. The selection of stripes versus spots is shown to depend on the nonlinear terms and cannot be discerned from the linearized model. The absence of quadratic terms leads to stripes but in most common models quadratic terms will lead to spot patterns. Examples that include neural nets and more general pattern formation equations are considered.