Evidence That Mid-lumbar Neurones in Reflex Pathways from Group II Afferents Are Involved in Locomotion in the Cat

1. A group of interneurons in the mid-lumbar segments of the cat spinal cord which mediate disynaptic excitation or inhibition of motoneurones from group II muscle afferents have recently been described. To test the possibility that the activity of these interneurones is related to the activity in the neuronal networks which subserve locomotion we have investigated whether they are influenced by two procedures which can induce locomotion. These procedures were electrical stimulation within the cuneiform nucleus (the 'mesencephalic locomotor region') in anaesthetized preparations and systemic administration of 3,4-dihydroxyphenylalanine (DOPA) in decerebrate, spinalized, unanaesthetized preparations. The interneurones we have tested were located in the fourth lumbar (L4) segment and were excited by group II muscle afferents; more than half of them were antidromically activated from the hindlimb motor nuclei. 2. Stimuli applied in the cuneiform nucleus evoked excitatory postsynaptic potentials (EPSPs) in a high proportion of these interneurones. The stimuli also evoked distinct extracellular field potentials in the ventral horn of the L4 segment. The properties and latencies of both the intra- and extracellularly recorded potentials show that they were evoked disynaptically, via supraspinally located relay neurones and a fast-conducting descending tract. 3. Stimulation of the cortico- and rubrospinal tracts excited or inhibited some of the L4 neurones, often at latencies suggesting mono- or disynaptic coupling. The neurones which appeared to be monosynaptically excited from the cortico- and rubrospinal tracts tended to be located dorsal to the neurones which were activated from the cuneiform nucleus. 4. Systemic administration of DOPA depressed the responses evoked by stimulation of group II afferents of L4 interneurones which projected to motor nuclei. DOPA also depressed extracellular field potentials evoked by group II afferents in the intermediate zone and in the ventral horn (at the location of the interneurones) but hardly affected those in the dorsal horn. 5. By showing that both stimulation in the cuneiform nucleus and the administration of DOPA influence activity of L4 interneurones which are excited by group II afferents and which project to motor nuclei, the results of this study support the hypothesis that these neurones are in some way involved in locomotion. However, the opposing effects of DOPA administration and of stimulation in the cuneiform nucleus make the interpretation of their role in locomotion rather difficult before it is known to what extent they are active throughout the step cycle.