BRAIN
RESEARCH
ELSEVIER
Brain Research 686 (1995) 150-159
Research report
Sleep permissive components within the dorsal raphe nucleus in the rat
B. E1 Karl *, L. Leger, S. Seguin, M. Jouvet, R. Cespuglio
Department of Experimental Medicine, 1NSERM U 52, CNRS-URA 1195, Claude Bernard University, 69373 Lyon Cedex 08, France
Accepted 14 March 1995
Abstract
Two peptides known for their hypnogenic properties, CLIP (corticotropin-like intermediate lobe peptide or ACTH 18-39) or VIP
(vasoactive intestinal polypeptide), were injected locally into the nucleus raphe dorsalis (nRD) of rats pretreated with p-chlorophenyl-
alanine (PCPA). During the dark period, the PCPA insomnia was primarily associated with a reduction in paradoxical sleep (PS), whereas
both slow wave sleep (SWS) and PS were decreased during the light period. Immunohistochemistry of serotonin in PCPA-pretreated
animals indicated a clear disappearance of 5-HT fibers in the basal hypothalamus and the nRD as compared to control animals. Local
injections of CLIP or VIP in the nRD restored PS and SWS. The positive injection sites corresponded to the anatomical distribution of
either CLIP or VIP fibers, i.e., the entire nRD for VIP and the antero-dorsal part of this nucleus for CLIP. The sleep effects obtained in
PCPA-pretreated rats involve a non-5-HT sleep permissive component within the nRD upon which these injected peptides act.
Keywords: Rat; Sleep; Wake; Nucleus raphe dorsalis; Local injection; Corticotropin-like intermediate lobe peptide; Vasoactive intestinal polypeptide;
p-Chlorophenylalanine
1. Introduction
That certain neuropeptides play a role in sleep regula-
tory processes is now generally accepted since they are
sleep inducing when administered by intraperitoneal (i.p.),
intravenous (i.v.) or intracerebroventricular (i.c.v.) routes.
Among these peptides, it is reported that i.c.v, administra-
tion of vasoactive intestinal polypeptide (VIP) at a 10 ng
dose produces an increase in slow wave sleep (SWS),
whereas at 100 ng paradoxical sleep (PS) is also enhanced
[16,39]. The administration of corticotropin-like intermedi-
ate lobe peptide (CLIP or ACTH18-39) during either the
dark [11] or light periods [47] also induces a PS increase.
Despite the common sleep inducing property of both of
the above peptides, their anatomical distribution within the
brain is very different. With respect to VIP, it is now
known that 4 major neuronal VIP systems are present in
brain whose perikarya are contained within the cortex, the
limbic system, the hypothalamus (suprachiasmatic nucleus)
and the rhombencephalic area (central grey, raphe nuclei,
medulla). From these different groups of perikarya, the
axonal processes produce a VIP innervation of almost the
* Corresponding author. Fax: (33) 7877-7172.
0006-8993/95/$09.50 © 1995 Elsevier Science B.V. All rights reserved
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entire brain. It must be emphasized that the nucleus raphe
dorsalis (nRD), an area of interest for the present report,
contains VIP perikarya in its medio-dorsal part and
widespread VIP axonal processes throughout the entire
nucleus. These anatomical aspects have been demonstrated
either by immunochemistry [18,42] or by VIP radio-im-
muno-assay [17].
The anatomical distribution of the proopiomelanocortin
(POMC) perikarya is more limited. These perikarya are
mainly located in the basal hypothalamus (within and
around the arcuate nuleus). Only one additional small
group has been described in the rat within the nucleus of
the solitary tract [28]. In contrast to VIP, the CLIP axonal
processes emerging from both of these groups of perikarya
are less numerous and do not reach the cortex. Instead,
they remain limited to the hypothalamus, thalamus, limbic
system, mesenchephalon and rhombencephalon [25,35].
The anterior and dorsal part of the nRD receives a strong
POMC innervation arising exclusively from the basal hy-
pothalamic group of perikarya [48]. Some of the axonal
POMC nerve endings have been demonstrated to be in
contact with nRD neurons [27].
Taking into account the above anatomical data and that
the nRD is thought to be a sleep permissive structure
gating phasic sleep events through its serotoninergic com-
ponent [23] and tonic sleep phenomena through a non