Localization and Connectivity in Spinal Interneuronal Networks: The Adduction–Caudal Extension–Flexion Rhythm in the Frog

Abstract
We have previously reported that focal intraspinal N -methyl-d-aspartate (NMDA) iontophoresis in the frog elicits a motor output, which is organized in terms of its constituent isometric force directions at the ipsilateral ankle and its topography. Furthermore, the associated EMG patterns can be reconstructed as the linear combinations of seven muscle synergies, labeled A to G. We now focus on one of the most common NMDA-elicited outputs, the adduction–caudal extension–flexion rhythm, and examine the relationship between the different force phases in terms of synergies and topography. Two distinct EMG patterns produce caudal extensions, and only one of the two patterns is used at most sites. The key synergy combinations for the two patterns are B + e and D + c (strongest synergies capitalized). These two patterns map at distinct locations in the lumbar cord. Within individual sites rhythms, we find linkages among the synergies used to produce adductions, the onsets of flexions after caudal extensions, and the synergy pattern producing the caudal extensions. For example, the synergy composition of adductions at B + e caudal extension sites is dominated by E + b and at D + c caudal extension sites by C + d. The two types of adductions map at distinct locations, situated between the two caudal extension regions. Specifically the linked patterns of caudal extension–adduction interleave rostrocaudally in a CE2-ADD1-ADD2-CE1 sequence, where 1 and 2 refer to the two pattern types. The implications of this topography and connectivity with respect to motor systems organization and behaviors are discussed.

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