GHS-R Family Members and Ligands/Smith et al. Vol. 14, No. 1 9
Author to whom all correspondence and reprint requests should be
addressed: Dr. Roy G. Smith, Huffington Center on Aging, Baylor College
of Medicine, M320, One Baylor Plaza, Houston TX 77030-3498. E-mail:
rsmith@bcm.tmc.edu
Endocrine, vol. 14, no. 1, 9–14, February 2001 0969–711X/01/14:9–14/$11.50 © 2001 by Humana Press Inc. All rights of any nature whatsoever reserved.
9
We have previously reported the cloning and charac-
terization of a new orphan G-protein-coupled recep-
tor (GPC-R), the growth hormone secretagogue
receptor (GHS-R), and shown that this receptor medi-
ates the activity of the growth hormone–releasing
peptides (GHRPs) and nonpeptide ligands such as
L-692,429 and MK-0677. Because the GHS-R obvi-
ously does not belong to any of the known GPC-R
subfamilies, we searched for GHS-R family members
by screening a human genomic library using low-strin-
gency hybridization and screening a Pufferfish
genomic library. The Pufferfish was selected because
of its compact genome. From the human genomic
library, a homolog, GPR38, with 52% identity to the
GHS-R was isolated. From the Pufferfish library, three
family members were isolated. The Pufferfish gene
having 58% identity to the GHS-R, on expression in
HEK293 cells, was activated with GHRP-6 and MK-0677.
These results indicate that the GHS-R has been con-
served for at least 400 million years and that the
Pufferfish genome is appropriate for isolation of GHS-
R family members. In our search for endogenous
ligands for the orphan receptors GHS-R and GPR38,
we showed that adenosine is a partial agonist of the
GHS-R and that motilin is the endogenous ligand for
GPR38. We also confirmed that the endogenous ligand
ghrelin is a full agonist of the GHS-R.
Key Words: Growth hormone secretagogue receptor;
motilin; adenosine.
Introduction
There remain many important issues to be resolved con-
cerning the biologic role of the growth hormone secreta-
gogue receptors (GHS-Rs) and the identity of endogenous
ligands for these receptors. Because the GHS-R appears to
be the first of a new family of G-protein-coupled receptors
(GPC-Rs) (1), we wished to identify GHS-R family mem-
bers and investigate the molecular evolution of the GHS-R.
We initially focused on the isolation of GHS-R family
members by searching expressed sequence tag (EST) data-
bases and screening a human genomic library. It soon became
evident that identification of GHS-R family members by
the latter approach was an enormous task. We reported the
isolation of three GHS-R homologs (GPR38, GPR39, and
FM3) from a human genomic PAC library (2,3). However,
screening this library provided positive hybridization sig-
nals for 325 PAC clones (2). The relatively low percentage
of coding sequences in the human genome complicates
characterization of GHS-R family members from this num-
ber of positive signals. To simplify the process, we evalu-
ated the potential of using the Pufferfish, a vertebrate
organism with a compact genome. The Pufferfish was
selected because it is evolutionarily distant from human
and homologous genes are likely conserved because of their
functional importance. Our objectives were first to deter-
mine whether the orphan GHS-R had been evolutionarily
conserved from Pufferfish to humans, and, second, to deter-
mine whether this vertebrate species would provide a short-
cut for identifying new GHS-R family members. To identify
natural ligands for the GHS-R and family members, we
developed high-volume screening assays. The assays were
developed such that they would be robust enough for iden-
tifying compounds in crude tissue extracts.
In this review we describe the identification of human
GHS-R homologs in the Pufferfish. We also report the iden-
tification of adenosine as an endogenous ligand of the GHS-
R and motilin (MTL) as the ligand for the closest GHS-R
family member, GPR38.
Human GHS-R Homolog in Pufferfish
Teleost fish have the specialized functions of higher
vertebrates; hence, genes necessary for complex functions
are present in their genome. The genome of the Japanese
Pufferfish (Fugu rubripes) had been proposed by Brenner
et al. (4) to be a compact model genome for vertebrates. The
Pufferfish genome (400 Mb) has approximately the same
Growth Hormone Secretagogue Receptor Family Members
and Ligands
Roy G. Smith,
1,2
Reid Leonard,
2
Alex R. T. Bailey,
1
Oksana Palyha,
2
Scott Feighner,
2
Carina Tan,
2
Karen Kulju McKee,
2
Sheng-Shung Pong,
2
Pat Griffin,
2
and Andrew Howard
2
1
Huffington Center on Aging and Department of Molecular and Cellular Biology, Baylor College of Medicine, Houston, TX;
and
2
Merck Research Laboratories, Rahway NJ