Ascending inputs to, and internal organization of, cortical motor areas

Modern anatomical studies show that, contrary to the long-held dogma, there appears to be essentially no convergence of lemniscal, cerebellar, pallidal, or substantial nigral afferents in the thalamus. Each afferent stream defines its own thalamic territory and, through the projection of these thalamic territories to separate cortical territories, the independence of the projections of subcortical motor nuclei upon the cortex is preserved. Only the spinothalamic system appears to gain access to both sensory and motor cortex. A further principle of organization in the sensorimotor thalamus is the presence of individual anatomically and physiologically defined channels, composed of separate afferent inputs and groups of neurons relaying to the cortex. In the somatic sensory relay nuclei the dissociation of cutaneous, deep slowly and rapidly adapting channels is clear-cut in the thalamus and at the input level of the cortex. In the motor system, inputs from each of the deep cerebellar nuclei appear to be dissociated from one another in the thalamus and these in turn from the vestibular and spinothalamic systems. Just as pallidal, nigral and cerebellar pathways are in position to control separate premotor and motor areas of the cortex, so separate channels leading through VLp appear to be in a position to control separate functional units in area 4. Within the cortex itself the absence of corticocortical connections passing from areas 3a and 3b to area 4 appears to indicate that information flow out of these areas is back to areas 1 and 2 for further processing before transmission to area 4 with all the consequences that entails for sensory convergence. Presumably, this route is sufficiently rapid for sensory inputs to reach area 4 at short latency. Although many data are beginning to accrue on the intrinsic structure and connectivity of the sensorimotor cortex, we are still distant from a complete wiring diagram. Circuitry involving thalamic afferents is becoming known slowly and the nature of the cells that are present and their transmitter characteristics are becoming evident from morphological and immunocytochemical studies, along with information on the patterns of axonal ramification of specific cell types, especially of GABAergic cells and of excitatory corticocortical cells.