Abstract

In plastic neuronal networks, the synaptic strengths are adapted to the neuronal activity. Specifically, spike-timing-dependent plasticity (STDP) is a fundamental mechanism that modifies the synaptic strengths based on the relative timing of pre- and postsynaptic spikes, taking into account the spikes' temporal order. In many studies, propagation delays were neglected to avoid additional dynamic complexity or computational costs. So far, networks equipped with a classic STDP rule typically rule out bidirectional couplings (i.e., either loops or uncoupled states) and are, hence, not able to reproduce fundamental experimental findings. In this review paper, we consider additional features, e.g., extensions of the classic STDP rule or additional aspects like noise, in order to overcome the contradictions between theory and experiment. In addition, we review in detail recent studies showing that a classic STDP rule combined with realistic propagation patterns is able to capture relevant experimental findings. In two coupled oscillatory neurons with propagation delays, bidirectional synapses can be preserved and potentiated. This result also holds for large networks of type-II phase oscillators. In addition, not only the mean of the initial distribution of synaptic weights, but also its standard deviation crucially determines the emergent structural connectivity, i.e., the mean final synaptic weight, the number of two-neuron loops, and the symmetry of the final connectivity pattern. The latter is affected by the firing rates, where more symmetric synaptic configurations emerge at higher firing rates. Finally, we discuss these findings in the context of the computational neuroscience-based development of desynchronizing brain stimulation techniques.

Highlights

  • In living systems, communication between different units typically does not occur instantaneously

  • Taking into account propagation delays is important for understanding the dynamics of neuronal networks and may help to further improve therapeutic brain stimulation techniques which aim at modulating plastic neuronal networks in diseased brains

  • Dendritic delays may range from sub-milliseconds to a few milliseconds,[4,9] whereas axonal delays may vary even more, up to tens of milliseconds in cortico-thalamic circuits.[10]. Comparing these experimentally observed ranges of dendritic and axonal propagation delays with the time scale of pairwise interactions between neurons indicates that propagation delays may have a significant impact on the nonlinear neuronal dynamics, especially on the performance of learning rules that modify the synaptic weight dynamics based on the timing of spiking neurons

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Summary

INTRODUCTION

Communication between different units typically does not occur instantaneously. When the presynaptic spike is temporally followed by a postsynaptic spike, a potentiation of the corresponding synaptic strength is induced by the STDP rule, whereas the synaptic strength is depressed when the ordering of pre- and postsynaptic spikes is reversed.[50] Due to this asymmetric protocol, in the absence of propagation delays, merely unidirectional connections can emerge between two reciprocally coupled spiking neurons under the influence of classic pair-based STDP.[42,43,44,45,46,47] This feature of STDP is in accordance with the existence of feedforward networks,[43,53,54] but it is in contradiction with the abundance of recurrent connections in cortical circuits.[60,63,64] as shown recently, incorporating dendritic and axonal propagation delays in the model can change the time lag between pre-post spike pairs so that different connectivity patterns may emerge in a system of plastic neuronal oscillators mediated by STDP.[34,35] Based on the range of dendritic and axonal propagation delays, symmetric connections, i.e., either two-neuron bidirectional loops or a decoupled motif can be stabilized.[34,35] In a special case where the dendritic and axonal propagation delays are identical, unidirectional connections can be observed. We discuss these findings in the context of the computation-based development of therapeutic brain stimulation techniques

Shortcomings of pair-based STDP
Variations of pair-based STDP
Emergence of symmetric connections
Multistability of final coupling regimes
CONCLUSIONS
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