Pergamon
0306-4522(94)00629-6
Neuroscience Vol. 66, No. 4, pp. 959-982, 1995
Elsevier ScienceLtd
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NEUROPEPTIDE Y AND SOMATOSTATIN-LIKE
IMMUNOREACTIVITY IN NEURONS OF THE MONKEY
AMYGDALA
A. J. MCDONALD,* F. MASCAGNI and J. R. AUGUSTINE
Department of Cell Biology and Neuroscience, University of South Carolina School of Medicine,
Columbia, SC 29208, U.S.A.
Akstraet--Neurons in the monkey amygdala exhibiting neuropeptide Y-like immunoreactivity and
somatostatin-like immunoreactivity were identified using an avidin-biotin immunohistochemical tech-
nique. Differential co-existence of the two peptides was demonstrated using two-color immunoperoxidase
and adjacent section methods. Numerous neuropeptide Y-positive neurons were observed in the
basolateral and superficial amygdaloid nuclei. A moderate number of neuropeptide Y-positive neurons
was seen in the medial subdivision of the central nucleus, but only a few neurons were observed in the
lateral subdivision. Numerous somatostatin-positive neurons were stained in all major amygdaloid nuclei
and always outnumbered neuropeptide Y-positive cells. All amygdaloid nuclei contained numerous
peptide-positive fibers whose density varied depending on the nucleus. Approximately 90% of neuro-
peptide Y-positive neurons also exhibited somatostatin-like immunoreactivity. The percentage of
somatostatin-positive neurons that exhibited neuropeptide-Y immunoreactivity varied in different nuclei.
In the superficial amygdaloid nuclei, medial subdivision of the central nucleus and most portions of the
basolateral nuclei the predominant cell type stained with both the neuropeptide Y and somatostatin
antibodies was a spine-sparse non-pyramidal neuron. In the dorsal portion of the lateral nucleus, however,
most peptide-positive neurons had spiny dendrites. Only the cell bodies and proximal dendrites of
somatostatin-positive neurons in the lateral subdivision of the central nucleus were immunostained.
This study demonstrates that specific cell populations in the primate amygdala contain neuropeptide
Y, somatostatin or both peptides. Most peptide-positive neurons in the basolateral and superficial
amygdaloid nuclei appear to be local circuit neurons that contribute to the dense plexus of peptide-positive
axons in these regions. The finding of neurons with spiny dendrites in the dorsal part of the lateral nucleus
suggests that these cells may be functionally different from peptide-positive neurons in other portions of
the basolateral amygdala. The lateral subdivision of the central nucleus is distinguished from other
amygdaloid nuclei by containing a large population of somatostatin-positive neurons that do not exhibit
neuropeptide Y immunoreactivity.
Neuropeptide Y (NPY) is a 36-amino acid peptide
that appears to act as a neuromodulator in the central
and peripheral nervous systems. 14'15'31'37'57'59 It is the
most abundant peptide in the brain and is especially
plentiful in the amygdala, 1'4 suggesting that NPY may
play an important role in modulating the activity of
amygdaloid neurons involved in behavioral, affective
and mnemonic functions. Immunohistochemical
studies in the rat have shown that the amygdala
contains a dense plexus of NPY-containing
axons and a moderate number of NPY-containing
perikaryaJ 3'28 Although mapping studies of the fore-
brain in the squirrel monkey 66 and human 73 have
demonstrated that there are NPY-containing cells
*To whom correspondence should be addressed.
Abbreviations: BDHC, benzidine dihydrochloride; CL,
lateral subdivision of the central nucleus; CM, medial
subdivision of the central nucleus; DAB, 3,3"-di-
aminobenzidine; Ld, dorsal portion of the lateral
nucleus; Lv, ventral portion of the lateral nucleus;
NADPH, reduced nicotinamide adenine dinucleotide
phosphate; NPY, neuropeptide Y; PBS, phosphate-
buffered saline.
and fibers in the primate amygdala, there have been
no comprehensive, detailed investigations of these
neuronal elements.
The peptide neuromodulator somatostatin was
first isolated from ovine hypothalamus and shown
to be a cyclic peptide consisting of 14 amino acids
(somatostatin-14). 8 Subsequent studies revealed the
existence of an N-terminally extended form of
somatostatin consisting of 28 amino acids (somato-
statin-28) and a related peptide consisting of the first
12 amino acids of the N-terminal of somatostatin-28
(somatostatin-28~ 12). 6'21 Previous studies of somato-
statin immunoreactivity in the amygdala of non-
human primates have utilized antibodies to somato-
statin-28~ 12 and the N-terminal segment of
somatostatin-28. 2'j7 There have been no studies using
antibodies that recognize somatostatin-14, the main
form of somatostatin present in the mammalian
brain. 2°
Immunohistochemical co-localization studies
have demonstrated that virtually all NPY-containing
neurons in the rat amygdala also contain somato-
statin, but the percentage of somatostatin neurons
959