The raphe nuclei of the cat brain stem: A topographical atlas of their efferent projections as revealed by autoradiography

Abstract
Stereotaxic injections of [14C]leucine were made in nuclei raphe´centralis superior, raphe´dorsalis, raphe´magnus and raphe´pontis of the cat. The organization of the regional connections was outlined in a stereotaxic atlas using the autoradiographic tracing method: the majority of the ascending pathways from the rostral raphe´nuclei are directed mainly through a ventrolateral bundle via the ventral tegmental area of Tsai, with some lateral extensions to the substantia nigra, and then through the fields of Forel and the zona incerta. More rostrally the fibers are joined to the medial forebrain bundle through the hypothalamic region up to the preoptic area or the diagonal band of Broca. Multiple divisions leave this tract towards the epithalamic or the intralaminar thalamic nuclei, the stria terminalis, the septum, the capsula interna and the ansa lenticularis. The bulk of the rostral projections terminates in the frontal lobe, while some labeling is scarcely distributed throughout the rest of the neocortex. The projections of nucleus (n.) raphe´centralis superior are specifically associated with the n. interpeduncularis, the mammillary bodies and the hippocampal formation while the n. raphe´dorsalis innervates selectively the lateral geniculate bodies, striatum, piriform lobes, olfactory bulb and amygdala. The rest of the ascending fibers form the centrolateral or the dorsal ascending tracts radiating either in the reticular mesencephalic formation or in the periventricular gray matter. On the contrary there are heavy descending projections from n. raphe´centralis superior which distribute to the main nuclei of the brain stem, the central gray matter and the cerebellum. The ascending projections from the caudal raphe´nuclei are much less dense. They disseminate mainly in the colliculus superior, the pretectum, the nucleus of the posterior commissure, the preoculomotor complex and the intralaminar nuclei of the thalamus. From n. raphe´pontis, a dense labeling is selectively localized at the n. paraventricularis hypothalami with some rostral extensions to limbic areas. Diffuse caudal and rostral projections from both nuclei are observed in the mesencephalic, pontobulbar reticular formation and the cerebellum. The main differences come from the specific localization of their descending bulbospinal tracts inside the lateroventral funiculus of the spinal cervical cord.