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of their close connections with the caudate and lenticular nuclei, the subthalamic nucleus (nucleus of Luys) and the substantia nigra are usually considered as parts of the basal ganglia The claustrum and amygdaloid nuclear complex, because of their largely different connections and functions, are usually excluded For reasons indicated further on, some physiologists have expanded the list of basal ganglionic structures to include the red nucleus, the intralaminar thalamic nuclei, and the reticular formations of the upper brainstem These additional structures receive direct cortical projections and give rise to rubrospinal and reticulospinal tracts which run parallel to the corticospinal (pyramidal) ones; hence they were also once referred to as extrapyramidal However, these nonpyramidal linkages are structurally independent of the major extrapyramidal circuits that include the striatum, pallidum, thalamus, and premotor and supplementary motor cortices Insofar as the nal links in this circuit the premotor and supplementary motor cortices ultimately project onto the motor cortex, the aforementioned circuit is more aptly referred to as prepyramidal (Thach and Montgomery) Earlier views of basal ganglionic organization emphasized the serial connectivity of the basal ganglionic structures and the funneling of efferent projections to the ventrolateral thalamus and thence to the motor cortex (Fig 4-3) This concept was based largely on the classic experimental work of Whittier and Mettler and of Carpenter, in the late 1940s These investigators demonstrated, in monkeys, that a characteristic movement disorder, which they termed choreoid dyskinesia, could be brought about in the limbs of one side of the body by a lesion localized to the opposite subthalamic nucleus They also showed that for such a lesion to provoke dyskinesia, the adjacent pallidum and pallidofugal bers had to be preserved; that is, a secondary lesion placed in the medial segment of the pallidum, in the fasciculus lenticularis, or in the ventrolateral thalamic nuclear group abolished the dyski-.

Figure 4-2 Diagram of the basal ganglia in the coronal plane, illustrating the main interconnections (see text for details) The pallidothalamic connections are illustrated in Fig 4-3

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nesia This experimental hyperkinesia could also be abolished permanently by interruption of the lateral corticospinal tract but not by interruption of the other motor or sensory pathways in the spinal cord Caudate nucleus These observations were interpreted Putamen to mean that the subthalamic nucleus normally exerts an inhibitory or regulating in uence on the globus pallidus and ventral thalamus Removal of this in uence by selective destruction of the subthalamic nucleus is expressed physiologically by irregular choreoid activity, presumably arising from the intact pallidum and conveyed by pallidofugal Corpus bers to the ventrolateral thalamic callosum nuclei, thence by thalamocortical Caudate nucleus bers to the ipsilateral premotor corLateral ventricle tex, and from there, by short assoPutamen ciation bers, to the motor cortex, Thalamus Globus pallidus: all in a serial manner New obserExternal segment Basal Internal Internal segment vations have made it apparent that ganglia capsule there are instead a number of paralSubthalamic lel circuits Ultimately the abnormal Claustrum nucleus movement is expressed via the latSubstantia nigra Amygdala eral corticospinal tract, which in this instance functions as the nal executive pathway Figure 4-1 Overview of the components of the basal ganglia in coronal view The main nuclei of the basal A general principle that ganglia are represented in blue, as labeled on the right (Reproduced with permission from Kandel ER, Schwartz emerged and withstood the test of JH, Jessell TM: Principles of Neural Science, 4th ed New York: McGraw-Hill, 2000)

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