European Journal of Neuroscience, Vol. 10, pp. 3257–3268, 1998 © European Neuroscience Association
The murine Y
1
receptor 5' upstream sequence directs
cell-specific and developmentally regulated LacZ
expression in transgenic mice CNS
Alessandra Oberto, Emanuela Tolosano,
1
Rossella Brusa, Fiorella Altruda,
1
Giancarlo Panzica
2
and Carola Eva
Section of Pharmacology, Department of Anatomy, Pharmacology and Forensic Medicine,
1
Department of Genetics, Biology
and Biochemistry, and
2
Section of Anatomy, Laboratory of Neuroendocrinology, Department of Anatomy, Pharmacology and
Forensic Medicine, University of Torino, Italy
Keywords: β-galactosidase, brain, neuropeptide Y, transgenic mouse, Y
1
receptor
Abstract
The Y
1
receptor for neuropeptide Y (NPY) is highly expressed in mammalian CNS where it mediates the
activation of several neurobiological functions. We have previously demonstrated that a 1.3-kb fragment
upstream of the transcription initiation sites of the murine Y
1
receptor gene is able to direct specific expression
of reporter genes in neuronal cell cultures. In the present study transgenic mice harbouring this putative
promoter region linked to the LacZ reporter gene were generated and analysed for temporal and spatial
distribution. Ten transgenic lines expressed β-galactosidase in the CNS but not in other organs such as heart,
liver and kidney. Histochemical analysis of brain from adult transgenic mice showed specific expression of the
transgene in specific brain regions with little variation. Four transgenic lines showed characteristic patterns of β-
galactosidase activity in the brain that are consistent with the expression of the endogenous gene. Prominent
LacZ activity was present in several telencephalic and diencephalic structures, including deeper layers of
cerebral cortex, amygdaloid complex, hippocampus, preoptic area, several thalamic and hypothalamic nuclei and
habenula. The ontogeny analysis indicates that the LacZ expression agrees with the temporal expression pattern
of rat Y
1
receptor mRNA. These data demonstrate that the 1.3-kb upstream region of the murine Y
1
receptor
gene contains the cis acting elements required for establishing a CNS-restricted and developmental stage-
specific pattern of expression in vivo. Moreover they provide further information on the distribution of this NPY
subtype receptor in mammalian brain.
Introduction
Neuropeptide Y (NPY) is a 36-amino-acid polypeptide with potent
central as well as peripheral biological effects, including stimulation
of food intake, anxiolysis, memory effects and cardiovascular effects
(Wahlestedt & Reis, 1993). NPY receptors are G-protein-coupled
receptors, all of which are associated with inhibition of adenylate
cyclase. At least six NPY receptor subtypes have been described on
the basis of their ability to bind NPY analogues, termed Y
1
,Y
2
,Y
3
,
Y
4
/PP, Y
5
and y
6
(Blomqvist & Herzog, 1997). Except for the Y
3
receptors, molecular cloning of cDNAs encoding corresponding
receptor proteins has been carried out (Eva et al., 1990; Larhammar
et al., 1992; Bard et al., 1995; Gerald et al., 1995, 1996; Rose et al.,
1995; Weinberg et al., 1996). They all belong to the G-protein-
coupled receptor superfamily and are coupled to more than one
second-messenger system. The functional y
6
receptor protein is only
expressed in mouse and rabbit and its role still remains to be
determined (Blomqvist & Herzog, 1997).
In the peripheral nervous system the Y
1
receptor is found predomi-
nantly at the sympathetic postjunctional sites in blood vessels, where
Correspondence: Dr Carola Eva, Sezione di Farmacologia, Dipartimento di Anatomia, Farmacologia e Medicina Legale, Universita ` di Torino, Via P. Giuria,
13-10125 Torino, Italy. E mail: eva@medfarm.unito.it
Received 12 February 1998, revised 28 May 1998, accepted 2 June 1998
it mediates the contractile response to NPY of vascular smooth
muscle, both directly and indirectly by potentiating the action of
other pressure agents, such as noradrenaline (Wahlestedt et al. 1986;
Grundemar et al., 1992; Malmstro ¨m, 1997).
In the rat central nervous system the distribution of Y
1
receptors
has been investigated using autoradiographical, immunohistochemical
and in situ hybridization techniques. Specific Y
1
binding sites are
mostly located in various cortical areas, olfactory anterior nucleus,
molecular layer of the dentate gyrus, claustrum and most of thalamic
nuclei (Dumont et al., 1990; Larsen et al., 1993; Dumont et al., 1996).
High levels of Y
1
receptors have been also identified in the arcuate
nucleus, and in the basket cells and Purkinje cells of cerebellum by
immunohistochemical studies (Zhang et al., 1994a; Fuxe et al., 1997;
Kopp et al., 1997). This distribution is not fully confirmed by in situ
hybridization, suggesting also a possible presynaptic role of the Y
1
receptor (Eva et al., 1990; Mikkelsen & Larsen, 1992; Larsen et al.,
1993). In addition Y
1
receptor and its mRNA were found in dorsal
horn of lumbar spinal cord and in dorsal root ganglia (Jazin et al.,
1993; Ji et al., 1994; Zhang et al., 1994a, b).