Anterograde and retrograde transport methods were used to study the corticocortical connectivity of areas 3a, 3b, 1, 2, 5, 4 and 6 of the monkey cerebral cortex. Fields were identified by cytoarchitectonic features and by thalamic connectivity in the same brains. Area 3a was identified by first recording a short latency group I afferent evoked potential. Attempts were made to analyze the data in terms of: (1) routes whereby somatic sensory input might influence the performance of motor cortex neurons; (2) possible multiple representations of the body surface in the component fields of the first somatic sensory area (SI). Apart from vertical interlaminar connections, two types of intracortical connectivity are recognized. The first, regarded as "non-specific," consists of axons spreading out in layers I, III and V-VI from all sides of an injection of isotope; these cross architectonic borders indiscrimininately. They are not unique to the regions studied. The second is formed by axons entering the white matter and re-entering other fields. In these, they terminate in layers I-IV in one or more mediolaterally oriented strips of fairly constant width (0.5--1 mm) and separated by gaps of comparable size. Though there is a broadly systematic topography in these projections, the strips are probably best regarded as representing some feature other than receptive field position. Separate representations are nevertheless implied in area 3b, in areas 1 and 2 (together), in areas 3a and 4 (together) and in area 5; with, in each case, the representations of the digits pointed at the central sulcus. Area 3b is not connected with areas 3a or 4, but projects to a combined areas 1 and 2. Area 1 is reciprocally connected with area 3a and area 2 reciprocally with area 4. The connectivity of area 3a, as conventionally identified, is such that it is probably best regarded not as an entity, but as a part of area 4. Areas identified by others as area 3a should probably be regraded as parts of area 3b. Parts of area 5 that should be more properly considered as area 2, and other parts that receive thalamic input not from the ventrobasal complex but from the lateral nuclear complex and anterior pulvinar, are also interconnected with area 4. More posterior parts of area 5 are connected with laterally placed parts of area 6. A more medial part of area 6, the supplementary motor area, occupies a pivotal position in the sensory-motor cortex, for it receives fibers from areas 3a, 4, 1, 2 and 5 (all parts), and projects back to areas 3a, 4 and 5.

The thalamic connectivity of areas 3b, 1 and 2 of the first somatic sensory cortex (SI) and of the adjacent areas 4, 3a and 5 has been studied in monkeys with anterograde and retrograde labeling techniques.Anteroposterior sectors of the SI cortex are represented in the thalamic ventrobasal complex by curved lamellae of thalamocortical relay cells extending through the dorsoventral and anteroposterior dimensions of the ventrobasal complex. Within such a lamella there are clustered aggregations of cells each projecting to a punctate zone of SI. Such cortical zones are less than 1 mm in circumference and are interpreted as comparable to the "columns" of electrophysiological studies. Each clustered aggregation in the ventrobasal complex is of limited mediolateral and dorsoventral extent but extends through much of the anteroposterior dimension of the ventrobasal complex. Punctate zones lying adjacent to one another in the mediolateral dimension of the SI cortex are connected with aggregations of cells lying in adjacent lamellae of the ventrobasal complex. Punctate zones lying anterior or posterior to one another in the anteroposterior dimension of SI are connected with aggregations of cells lying ventral or dorsal to one another in a lamella of the ventrobasal complex. 'Collectively, these two nuclei, forming the part of the thalamus activated by somesthetic stimuli, are termed the "ventrobasal complex" (Rose and Mountcastle, '52, '54). The ventrobasal complex excludes the oral division of the ventroposterolateral nucleus, as defined by Olszewski ('52).

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The cells of origin of the corticostriatal projection have been identified in squirrel monkeys by the use of the retrograde horseradish peroxidase method. In the subfields of the somatic sensory, motor, parietal and frontal areas of the cortex, cells projecting to the ipsilateral striatum are relatively sparsely distributed and form a group of small- to medium-sized pyramidal cells with an average somal diameter from area to area of 14-16 mum. Such cells are found only in layer V of the cortex (mainly in the more superficial parts of the layer). Since they are consistently smaller than the pyramidal cells of layer V that project to the brainstem and spinal cord and since they lie outside layer VI which gives rise to corticothalamic axons, the corticostriatal axons are unlikely to be collaterals of axons projecting to other sites. The cells of origin of the crossed corticostriatal projection are also found in layer V and are pyramidal cells with somal diameters in the same range as above. They are found only in areas 4, 8, and 6. Studies with the anterograde, autoradiographic method in rhesus, cynomologous and squirrel monkeys, indicate that the somatic sensory areas project to most of the antero-posterior extent of the ipsilateral putamen. Subareas 3a, 3b, 1 and 2 of the somatic sensory cortex project to the same region and the projection overlaps similarly extensive projections from the motor and certain other areas of the cortex. However, in each case the pattern of terminal labeling is in the form of interrupted clusters, strips and bands. A single small injection of the cortex is associated with only one or two such clusters of terminal labeling. This seems to imply that individual corticostriatal fibers end in a very restricted manner and that the terminal ramifications of fibers from one cortical area may alternate in the putamen with those arising in other areas.

The retrograde axonal transport method has been employed to identify the cell bodies of cortical neurons projecting directly to the spinal cord in the monkey. The investigation has focused on aspects of the laminar, columnar, and somatotopic organization of corticospinal neurons within each of the cytoarchitectural and functional subdivisions of the sensorimotor cortex. The principle findings of these experiments are that: i) cortical regions containing cell bodies of corticospinal neurons are the first motor cortex (area 4), the first somatic sensory cortex (areas 3a, 3b, 1, and 2), and part of the immediately adjacent posterior parietal cortex (area 5), the second somatic sensory cortex, the supplementary motor cortex (the medial aspect of area 6), and the medial part of the posterior parietal cortex in a region termed the supplementary sensory area; ii) corticospinal neurons display a somatotopic organization within each of these functional subdivisions of the sensorimotor cortex; iii) all corticospinal neurons arise from layer V of the cortex; and iv) corticospinal neurons within the first motor and first somatic sensory cortex often occur in clusters, perhaps reflecting a columnar organization in the sensorimotor cortex. These findings demonstrate the origins of the corticospinal system to be more extensive than previously recognized and show that a number of common features characterize the organization of corticospinal neurons in all cortical areas. Across cortical subdivisions, however, major differences exist in the extent of spinal segmental representations, in the manner in which corticospinal neurons occur in groups, and in the numerical density and sizes of corticospinal neurons. These aspects of the organization of the corticospinal system presumably reflect specialization of the different cortical areas in spinal cord sensory and motor control.