Central Actions of GHS-25/Bailey et al. Vol. 14, No. 1 15
Author to whom all correspondence and reprint requests should be
addressed: Alex Bailey, Eurosterare Administrator, Centre for Reproductive
Biology, Edinburgh EH3 9ET. E-mail: alex.bailey@obg93.obg.ed.ac.uk
Endocrine, vol. 14, no. 1, 15–19, February 2001 0969–711X/01/14:15–19/$11.25 © 2001 by Humana Press Inc. All rights of any nature whatsoever reserved.
15
Growth hormone secretagogues (GHSs) increase the
activity of hypothalamic arcuate nucleus neurons
thought to be involved in controlling the release of
growth hormone (GH). The GHS receptor is also found
in hypothalamic regions not associated with the
release of GH, suggesting that GHSs may influence
other hypothalamic systems. This study utilized
double-labeling immunocytochemical techniques to
examine the hypothalamic actions of a novel non-
peptide GHS, GHS-25. In common with other GHSs,
GHS-25 induced significant amounts of Fos immunore-
activity in the arcuate nucleus of conscious male rats.
However, unlike other GHSs, GHS-25 also induced
Fos immunoreactivity in the supraoptic nucleus.
Double labeling revealed that approx 66% of supraop-
tic nucleus cells that were Fos positive after the admin-
istration of GHS-25 were also immunoreactive for
oxytocin. Thus, in addition to its actions on the GH
axis, GHS-25 may influence the release of neurohypo-
physeal hormone.
Key Words: Hypothalamus; arcuate nucleus; supraop-
tic nucleus; oxytocin; vasopressin.
Introduction
Growth hormone (GH) release can be elicited directly
from the anterior pituitary gland by growth hormone secre-
tagogues (GHSs), such as GHRP-6 and MK-0677 (1,2). In
addition to inducing GH release directly from the pituitary,
GHSs act centrally to increase Fos protein expression in the
hypothalamic arcuate nucleus and induce a prolonged in-
crease in the electrical activity of neurosecretory arcuate
nucleus neurons (3,4). Within the arcuate nucleus, the main
populations of GHS-responsive cells are thought to include
both GH-releasing hormone (GHRH) and neuropeptide Y
(NPY) cells (5). Indeed, GHS mRNA has recently been
localized to GHRH and NPY arcuate nucleus neurons (6,7).
However, GHS receptor mRNA is also found in several
other hypothalamic nuclei (e.g., the supraoptic nucleus
[SON] and suprachiasmatic nucleus [SCN]), and in other
discrete regions of the brain, including the CA2 and CA3
regions of the hippocampus, the dentate gyrus, and sub-
stantia nigra (8). This had led to speculation that the GHS
receptor (GHS-R), and therefore its endogenous ligand,
may play a role in processes that are not generally associ-
ated with the control of GH release.
The cloning of the GHS-R has provided a context in
which agonist/receptor interactions can be investigated sys-
tematically. This has revealed that several structurally
diverse compounds are capable of binding to the receptor
and that different classes of GHSs can bind to different
regions of the ligand-binding pocket of the receptor (9).
This, in turn, has led to the generation of several novel
GHSs that may activate intracellular pathways different
from those by conventional GHSs, possibly leading to a
broader range of physiologic actions.
Recent studies have demonstrated that some GHSs are
capable of eliciting the release of vasopressin from hypo-
thalamic explants (10), and vasopressin release is thought
to, at least partially, underlie the hypothalamo-pituitary-
adrenal axis activation induced by GHSs (11). Cell bodies
of vasopressin neurons are located in the SON, SCN, and
paraventricular nucleus (PVN) of the hypothalamus (12),
and all these nuclei express GHS-R mRNA (8). However,
previous studies have shown that GH-releasing peptide-6
(GHRP-6) and MK-0677 do not elicit Fos immunoreactivity
within these regions (3,4). As within the hypothalamic-
neurohypophyseal system, the use of Fos immunocyto-
chemistry as a marker of neuronal activity is considered a
reliable and intensity-dependent stimulus (13); it appears
that GHRP-6 and MK-0677 do not activate cells in the SON
or PVN. However, because novel GHSs may interact with
the GHS-R in a different manner from GHRP-6 or MK-
0677, the apparent disparities between these studies may
reflect activation of different regions of the GHS-R.
Therefore, in the present study, we employed the immu-
nocytochemical detection of Fos protein in the rat to com-
Central Actions of the Nonpeptide Growth Hormone
Secretagogue GHS-25
Alex R. T. Bailey,
1
Louise Gilliver,
2
Gareth Leng,
2
and Roy G. Smith
1
1
Huffington Center on Aging, Baylor College of Medicine, Houston, TX;
2
Department of Biomedical Sciences, University
Medical School, Edinburgh, EH8 9AG, UK.