The mesencephalic–hypoglossal nuclei loop as a possible central pattern generator for rhythmical whisking in rats

It has been previously demonstrated that the Me5 nucleus is involved in the genesis of reflex activities at whisker pad level. Specific Me5 neurons, which provide sensory innervation of the macrovibrissae, are monosynaptically connected with small hypoglossal neurons innervating the extrinsic muscles that control macrovibrissal movements. Artificial whisking, induced by the electrical stimulation of the peripheral stump of the facial nerve and the electrical stimulation of the XII nucleus or the infraorbital nerve, induced evoked responses in the whisker pad extrinsic motor units, along with a significant increase in the electromyographic activity of the extrinsic pad muscles (Mameli et al. in Acta Oto-Laryngol 126:1334–1338, 2006; in Pfűgers Arch Eur J Physiol 456:1189–1198, 2008; in Brain Res 1283:34–40, 2009; in Exp Brain Res 234:753–761, 2016). In anaesthetized rats, we evaluated the possible involvement of this Me5–XII loop in the genesis of rhythmical whisking. The anatomical findings showed that in addition to the ipsilateral, even the contralateral Me5 nucleus could be retrogradely labeled by the Dil tracer injected into the whisker pad of one side, they, furthermore, showed labeled axons extending across the midline between the two nuclei. The electrophysiological findings agreed with the neuroanatomical results, since the mechanical or artificially induced deflection of the whiskers of one side, evoked in the Me5 contralateral nucleus different patterns of responses. The hypothesis that the Me5–XII loops, along with their cross-linked relationship, could act as a “central generator” responsible for the stereotyped symmetrical pattern of macrovibrissal movements such as rhythmical whisking has been discussed.