Abstract

Networks in the spinal cord, which are responsible for the generation of rhythmic movements, commonly known as central pattern generators (CPGs), have remained elusive for decades. Although it is well-known that many spinal neurons are rhythmically active, little attention has been given to the distribution of firing rates across the population. Here, we argue that firing rate distributions can provide an important clue to the organization of the CPGs. The data that can be gleaned from the sparse literature indicate a firing rate distribution, which is skewed toward zero with a long tail, akin to a normal distribution on a log-scale, i.e., a “log-normal” distribution. Importantly, such a shape is difficult to unite with the widespread assumption of modules composed of recurrently connected excitatory neurons. Spinal modules with recurrent excitation has the propensity to quickly escalate their firing rate and reach the maximum, hence equalizing the spiking activity across the population. The population distribution of firing rates hence would consist of a narrow peak near the maximum. This is incompatible with experiments, that show wide distributions and a peak close to zero. A way to resolve this puzzle is to include recurrent inhibition internally in each CPG modules. Hence, we investigate the impact of recurrent inhibition in a model and find that the firing rate distributions are closer to the experimentally observed. We therefore propose that recurrent inhibition is a crucial element in motor circuits, and suggest that future models of motor circuits should include recurrent inhibition as a mandatory element.

Highlights

  • It is known that the core neural elements of rhythmic movement, the central pattern generators (CPGs), are located in the spinal cord and the medulla, the neuronal architecture of these networks has remained perplexing

  • Using optogenetics and light activation or inhibition of spatially restricted regions in the spinal cord it was possible to Firing-Rate Distributions and Motor Control exclusively activate either flexor or extensor rhythms suggesting a modular organization (Hägglund et al, 2013)

  • The idea of the “log-normal” brain is inspired by the skewed shape of the firing rate distributions (Mizuseki and Buzsáki, 2013; Buzsáki and Mizuseki, 2014)

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Summary

INTRODUCTION

It is known that the core neural elements of rhythmic movement, the central pattern generators (CPGs), are located in the spinal cord and the medulla, the neuronal architecture of these networks has remained perplexing. A common theme in the literature is the half-center organization inspired by Brown (1914), where two rhythm generating modules, which have recurrent excitation, are coupled reciprocally via inhibitory populations to ensure an alternating flexor and extensor activity (McLean and Dougherty, 2015; Kiehn, 2016; Grillner and El Manira, 2020). It has been widely suggested that the rhythm generating modules are composed of a recurrently connected excitatory networks, where the rhythm is generated by a subset of neurons with pacemaker properties (Grillner and El Manira, 2020). What decides if a network is stable and would a network composed purely of recurrent excitation be sensible for generation of motor activity?

RECURRENT NETWORKS
DYNAMICS OF A RECURRENT EXCITATORY NETWORK
EXPERIMENTALLY OBSERVED FIRING RATE DISTRIBUTIONS
COULD RECIPROCAL INHIBITION HELP?
INCLUSION OF RECURRENT INHIBITION
BLOCKING INHIBITION MAKES THE NETWORK UNSTABLE
DISCUSSION
METHODS
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