Callosal projections from area SII to SI in monkeys: Anatomical organization and comparison with association projections

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Abstract
The present research was aimed at ascertaining in the macaque monkey the reciprocity of the heterotopical callosal connections between SI and SII, with particular regard to the connectivity of the hand representation, and at comparing the topographical and laminar pattern of these callosal connections with those of association connections entertained by these areas. Horseradish peroxidase (HRP) was unilaterally injected into area SI in five monkeys. The sites of HRP delivery included the trunk and the hand zones preliminarily identified by recording multi‐unit responses to peripheral stimulation by means of microelectrodes. Anterograde and retrograde labelling was studied in SII of both sides. The results showed the complete reciprocity of the heterotopical callosal connections between SI and SII. In the latter area both callosal axon terminals and neurones were found, which were labelled from either the trunk or the hand zone of contralateral SI. Labelling of callosal axon terminals occurred mainly in layer IV and in the lowermost part of layer III. Labelled callosal neurones were mainly in the lower half of layer III, whereas few occurred in infragranular layers. Topographically, the distribution of callosal terminals and cell bodies duplicated the distribution of association terminals and cell bodies labelled in SII on the side ipsilateral to HRP injection. The laminar pattern of termination of association fibres from SI was similar to that of callosal fibres. However, the distribution of association‐projecting neurones in SII showed a striking difference from that of callosal‐projecting neurones. Unlike the latter neurones, which were mainly located in supragranular layers, association cell bodies overwhelmingly dwelt in layers V and VI and were less numerous in layers II and III. This laminar pattern of association SII‐SI cells corresponds to the “feed‐backward” model and fits the laminar pattern of their axon terminations (Friedman: Brain Res. 273: 147–151, '83). The association and callosal inputs and outputs of area SII are discussed in relation to the function of the forward and backward type of reciprocal connections entertained with SI in the ipsilateral hemisphere and to the function of SII in the interhemispheric exchange of somatosensory information.