Laminar pattern of termination of the ipsilateral cortical projection from SII to SI in cats

The present light and electron microscopic experiments were carried out on the first somatic sensory area (SI) of cats to determine the laminar distribution of axon terminals from the ipsilateral second somatic sensory area (SII) and to identify the types of synapses between these terminals and the neuronal elements of SI. Phaseolus vulgaris-leucoagglutinin (PHA-L) was iontophoretically injected into multiple sites and at different cortical depths of the forepaw representation zone of SII. Fixed brain blocks containing the injected SII and ipsilateral SI were cut into slices and processed immunocytochemically to stain PHA-L-filled fibers and terminals. Light microscopic examination of SI revealed patches of anterograde labeling in the forepaw representation zone, concentrated mainly in supragranular layers. In these layers, thin immunolabeled fibers branched extensively and formed a dense plexus that was more prominent in layers II and I. Conversely, the infragranular layers contained fragments of vertically oriented thick fibers that rarely emitted axon collaterals. PHA-L-labeled axons had numerous swellings along their course, interpreted as boutons en passant, and stalked boutons. Of 19,661 labeled terminals (17,833 beads and 1,828 stalked boutons), 84.74% were observed in supragranular layers, with the highest concentration in layer II (33.15%) and lower in layers I (26.27%) and III (25.30%). The proportion of terminals was lower in layers IV (6.49%) and V (5.45%) and lowest in layer VI (3.32%). These counts also showed that boutons en passant were the majority (90.70%) and stalked boutons, the minority (9.30%). The ratio of these two types of presynaptic specializations was similar (9:1) in all six layers. Electron microscopic examination of the labeled regions of SI showed that both axon swellings and stalked boutons formed synapses of the asymmetric type with SI neuronal elements. The majority (85.37%) of a sample of 130 labeled terminals synapsed on SI neurons in layers I-III. The identified postsynaptic profiles were dendritic spines (61.11%) or medium-sized and small dendrites (38.89%). These results are discussed in relation to those of a companion study on the laminar pattern of the projection from SI to SII of cats (P. Barbaresi, A. Minelli, and T. Manzoni, 1994, J. Comp. Neurol. 343:582-596). Based on the anatomical organization of these reciprocal connections, there seems to be no clear hierarchicalal relationship between SI and SII in cats.