An interneuronal relay for group I and II muscle afferents in the midlumbar segments of the cat spinal cord.

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Abstract
1. The properties of interneurons located in the 4th lumbar segment of the cat spinal cord (L4 interneurones) have been investigated by intracellular and extracellular recording from individual neurones. The study focused on interneurones projecting to hind-limb motor nuclei and/or interposed in pathways from group II muscle afferents. The projection of motor nuclei was assessed from antidromic activation of the neurones by stimuli applied in the motor nuclei of the 7th lumbar (L7) segment. 2. Interneurones which projected to gastrocnemius-soleus or hamstring motor nuclei were found in laminae VI and VII and at the border between laminae VII and VIII. The dominant peripheral input to most of them was from group II muscle afferents, but they were also influenced by group I muscle afferents and by afferents in cutaneous, joint and interosseous nerves. Both excitatory post-synaptic potentials (e.p.s.p.s) and inhibitory post-synaptic potentials (i.p.s.p.s.) were evoked from all of these fibre systems. 3. The same kind of multimodal input was also found in other interneurons in laminae VI and VII. However, their axonal projections were not identified and they might have included neurones projecting to motor nuclei (though outside the areas which were stimulated) as well as neurones with more local actions. 4. Interneurones located in laminae IV and V of the dorsal horn appeared to constitute a separate functional population since both their projections and their input differed from those of the more ventrally located interneurones; none of the dorsal horn interneurones were found to project to motor nuclei and none had input from group I afferents, although they were influenced by group II muscle afferents and by afferents in cutaneous, joint and interosseous nerves. 5. Many of the excitatory actions from group I and II afferents upon L4 interneurones were found to be evoked monosynaptically. A high proportion of L4 neurones synapsing upon motoneurones would thus be interposed in disynaptic reflex pathways from these afferents. In comparison to actions evoked via interneurones of the caudal lumbar segments, any post-synaptic potentials (p.s.p.s.) evoked via L4 interneurones would be delayed. These delays would amount to 0.4-0.9 ms for p.s.p.s. from group I afferents and by 0.5-2.5 ms for group II p.s.p.s. 6. In many interneurones, particularly those located ventrally, i.p.s.p.s. were evoked by group I and II muscle afferents at latencies which indicated that they were evoked disynaptically. They may therefore reflect inhibitory interactions between subpopulations of L4 interneurones. 7. The main input to L4 interneurones appears to be from group II afferents from the quadriceps and sartorius muscles. They are therefore likely to be particularly strongly activated during extension of the hip joint, when quadriceps and sartorius are stretched. The possibility is thus considered that L4 interneurones play a key role in reflexes determined by limb position.