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ELSEVIER Neuroscience Letters 187 (1995) 119-122
Galanin is contained in GABAergic neurons in the rat spinal dorsal horn
D.R. Simmons, R.C. Spike, A.J. Todd*
Laboratory of Human Anatomy, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK
Received 11 January 1995; revisedversionreceived25 January 1995; accepted27 January 1995
Abstract
In order to determine which types of neuron in laminae I-III of the rat spinal dorsal horn contain the peptide galanin, pre-embedding
immunocytochemistry with antiserum to galanin was combined with post-embedding detection of GABA- and glycine-like
immunoreactivities. Sixty-eight galanin immunoreactive neurons in laminae I-III selected from four rats were examined, and in each
case semi-thin sections through the cell body were tested with a monoclonal antibody to GABA and an antiserum to glycine. All of the
68 galanin-immunoreactive neurons tested were GABA-immunoreactive, while only one of them (in lamina III) was glycine-
immunoreactive. This suggests that galanin is contained in inhibitory intemeurons, and that (like enkephalin, neuropeptide Y and
thyrotropin-releasing hormone) it is mainly restricted to GABAergic neurons which do not use glycine as a co-transmitter.
Keywords: Immunocytochemistry; Spinal cord; Glycine; Peptides; Transmitter co-localisation
Galanin is a peptide containing 29 amino acids, which
was originally isolated from pig intestine [14], and has
since been found in various parts of the central nervous
system [1,7,8,10]. Many galanin-immunoreactive axons
are present in laminae I and II of the rat spinal dorsal horn
[1,7,8,10,21,23,24], and these are thought to be derived
from both dorsal root ganglion cells and local interneu-
rons [1,5,11,17,21,23,24]. Galanin-immunoreactive neu-
rons in the superficial laminae of the dorsal horn were
initially observed after intrathecal administration of col-
chicine [1,7,8], however with more sensitive immunocy-
tochemical techniques they can be detected without appli-
cation of colchicine [23,24]. At present, relatively little is
known about the types of dorsal horn neuron that contain
galanin. Attempts have been made to classify neurons in
the superficial dorsal horn on both morphological and
neurochemical grounds (for review, see Ref. [17]). Within
lamina II, two main types of neuron have been identified:
stalked and islet cells [4]. Stalked cells have their cell
bodies in the dorsal part of the lamina and dendrites that
pass ventrally, while islet cells have long, rostrocaudally
orientated dendrites. There is a relationship between mor-
phology and transmitter content, since large islet cells in
lamina II are GABA-immunoreactive, whereas stalked
* Corresponding author. Tel.: +44 41 3398855; Fax: +44 41 3304299.
cells and a population of small islet cells are not [15,17].
Glycine-like immunoreactivity (-LI) is also observed in
cell bodies in laminae I-III, but is virtually restricted to
cells that are GABA-immunoreactive, and it has been
suggested that these neurons release both inhibitory
transmitters [20]. Zhang et al. [24] have recently reported
that galanin-immunoreactive neurons in lamina II of rat
dorsal horn resemble islet cells, and since many of these
are thought to be inhibitory interneurons which release
GABA (with or without glycine), the present study was
carried out to determine whether galanin co-exists with
GABA or glycine in cells of the superficial dorsal horn.
Pre-embedding immunocytochemistry to detect gala-
nin in cell bodies was combined with post-embedding
detection of GABA- and glycine-LI as described previ-
ously [3,9]. Four adult Albino-Swiss rats (either sex; 220-
280 g) were deeply anaesthetised and perfused with a
fixative containing 1% glutaraldehyde/l% formaldehyde
in 0.1 M phosphate buffer (PB). Segments of mid-lumbar
spinal cord were removed, stored in the same fixative
overnight and cut into 70-/tm transverse sections with a
Vibratome. The sections were treated in 1% sodium bo-
rohydride for 30 min, rinsed extensively in phosphate
buffered saline (PBS), incubated for 1 h in 10% goat se-
rum in PBS and then overnight in rabbit antiserum to rat
galanin (Peninsula) diluted 1:10000, and processed by
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