Search

Three morphological types of axons have been recognized in previous studies of the serotoninergic (5-HT) innervation of the cat cerebral cortex (Mulligan and Törk, 1988; DeFelipe et al., 1991): thick, nonvaricose axons, fine axons with small varicosities, and beaded axons with large, spherical varicosities. In the present study, the laminar density and distribution of the three 5-HT fiber types in area 17 are characterized. In both coronal and sagittal immunostained sections, 5-HT axons exhibit an overall gradient of decreasing density from layer I to the white matter. The three 5-HT axon types exhibit distinctive innervation patterns. (1) Fine axons with small varicosities comprise the greatest number of fibers in each layer (56-98%). They usually have oblique or radial trajectories, but some horizontally oriented fibers run through layers I, III, VI, and the white matter. (2) Non-varicose axons, the preterminal portions of the beaded axons with the large varicosities (Mulligan and Törk, 1988), comprise only about 7% of the total 5-HT fiber population in area 17, and are found mainly in layer I and in the white matter where they form a horizontally oriented plexus. (3) Large varicose axons ("beaded" axons) are rare (about 3% of the total 5-HT population) and are restricted to layers I and V. Although large varicose axons often form elaborate pericellular basketlike arbors around the soma and dendrites of neurons in other parts of the cat cerebral cortex (Mulligan and Törk, 1987, 1988), such arrays are rare in area 17. When observed in area 17, pericellular arrays are typically simple structures with a few large varicosities apparently contacting only the somata of a small population of layer I neurons. Comparison of these results with reports of 5-HT innervation of area 17 in other species suggests that the 5-HT innervation of the cortex is highly species specific.

Serotonergic axons in the posterior cerebral cortex of the cat were demonstrated immunohistochemically using a monoclonal antibody to serotonin (5-HT). This technique reveals the presence of a dense serotonergic innervation of single cortical neurons at the light microscopic level. 5-HT axons with large varicosities (1-6 microns in diameter) form distinct, basket-like arrays around counterstained somata principally in layer I. In each basket one or more axons encircle and make repeated contact with the soma. Some axons extend from the soma and apparently climb along the dendrites of the target neuron. The climbing 5-HT axons form a stellate or horizontal pattern suggesting that the target cells are non-pyramidal neurons of the supragranular layers.

Serotoninergic axons in the cat cerebral cortex were demonstrated immunohistochemically with a monoclonal antibody to serotonin (5-HT). Three types of 5-HT axons are distinguished at the light microscopic level by differences in their morphology. Small varicose axons are fine (less than 0.5 micron) and bear fusiform varicosities that are generally less than 1 micron in diameter. These axons extend throughout the width of the cortex and branch frequently, giving rise to widely spreading collaterals. Nonvaricose axons are smooth, show a relatively large and constant caliber (about 1 micron), travel in straight, horizontal trajectories, and branch infrequently. Large varicose axons are distinguished by large round or oval varicosities (1 micron or more in diameter) borne on fine-caliber fibers. These axons often form basket-like arbors around the somata of single neurons. In the simplest basket-like arbors, several large, round varicosities from a small number of axons contact the soma. In complex baskets intertwining collaterals contact the soma and apparently climb along and outline the cell's major dendrites. The patterns revealed by the climbing axons suggest that a variety of nonpyramidal cell types selectively receive dense 5-HT innervation. Serial reconstructions of the 5-HT axons within the cortex show that the large varicose axons arise as infrequent collaterals from the nonvaricose axons. A single nonvaricose parent axon gives rise to several large varicose axon collaterals that may contribute to different basket-like arbors. Conversely, a single basket-like arbor may be formed by large varicose axon collaterals from more than one nonvaricose parent axon. The small varicose axons do not appear to be related within the cortex to either the nonvaricose or large varicose axon types. The results support the hypothesis that the 5-HT projection to the cortex is organized into two subsystems, one of which may exert widespread influence in the cortex via highly divergent branches, while the other, with a more restricted distribution, acts on specific classes of cortical neurons.