Neural substrates of two different rhythmical vibrissal movements in the rat

Normal rats show two types of rhythmical vibrissal movement, one synchronized precisely with alpha waves (about 9 Hz) of the thalamocortical system, and the other often synchronized with theta waves (about 7 Hz) of the septohippocampal system. The alpha-synchronized vibrissal movements appear while the rat stands still with a slow respiratory pattern, and are of small amplitude (a fine tremor). The theta-synchronized vibrissal movements appear during exploratory sniffing behavior, and are of large amplitude. Thus, a group of facial motoneurons which constitute the final common pathway for vibrissal movement apparently receive input convergently from these two neural systems. In the present study, we observed the following: (1) the two types of movement rarely, if ever, appeared simultaneously and the same was true of the two brain wave patterns. Topographically, the predominant appearance of the alpha waves was in the frontal (sensorimotor) cortex, whereas that of the theta waves was in the occipital cortex and hippocampus. (2) Bilateral ablation of either the entire neocortex or just its anterior (but not posterior) half eliminated the vibrissal alpha-tremor movement while leaving vibrissal theta-sniffing movement normal. In anterior decorticate rats, the tremor movements started to recover by 1-4 months, and were abolished again by the removal of the remaining posterior cortex. (3) Lesions of the medial septum or the fornix eliminated hippocampal theta waves, but had no effect on vibrissal sniffing movement or alpha wave-vibrissal tremor. (4) Cerebellectomy and, to a lesser extent, pharmacological lesions of the inferior olive slowed, but did not block, alpha waves. In addition, vibrissal tremor movement became intermittent and less vigorous. The same manipulations, however, did not affect theta wave-vibrissal sniffing movement. (5) Harmaline (30 mg/kg, i.p.) did not induce alpha-tremor, which, in combination with the results with cerebellectomy and inferior olive lesions, indicates that alpha-tremor is generated by a neural mechanism that is different from that for harmaline-induced generalized tremor of 10 Hz. These findings confirmed that there exist two patterns of synchrony between vibrissal movement and rhythmical brain activity in the rat, i.e. alpha wave-vibrissal tremor movement and theta wave-vibrissal sniffing movement, and suggest that the two patterns reflect a rhythmical mode of functioning of two different neural systems, probably the thalamocortical and the septohippocampal system, respectively.