~ Pergamon
0306-4522(95)00595-1
Neuroscience Vol. 72, No. 4, pp. 1023-1036, 1996
Elsevier ScienceLtd
Copyright © 1996 IBRO
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REDUCTION IN ENKEPHALIN AND SUBSTANCE P
MESSENGER RNA IN THE STRIATUM OF EARLY
GRADE HUNTINGTON'S DISEASE: A DETAILED
CELLULAR IN SITU HYBRIDIZATION STUDY
S. J. AUGOOD,*f R. L. M. FAULL,~ D. R. LOVE§ and P. C. EMSON*
*MRC Molecular Neuroscience Group, Department of Neurobiology, The Babraham Institute,
Babraham, Cambridge CB2 4AT, U.K.
:~Department of Anatomy and §School of Biological Science, University of Auckland, Private Bag 92019,
Auckland, New Zealand
Abstract--The expression of enkephalin and substance P messenger RNAs was examined in the
caudate-putamen of human post mortem tissue from control and Huntington's disease tissue using in situ
hybridization techniques and human specific enkephalin and substance P [35S]oligonucleotides. Macro-
scopic and microscopic quantification of enkephalin and substance P gene expression was carried out using
computer-assisted image analysis. Tissue was collected from six control cases with no sign of neurological
disease and six Huntington's disease cases ranging from grades 0 to 3 as determined by neuropathological
evaluation. The clinical and pathological diagnosis of Huntington's disease was confirmed unequivocally
by genetic analysis of the CAG repeat length in both copies of IT15, the Huntington's disease gene. A
marked reduction in both enkephalin and substance P messenger RNAs was detected in all regions of the
caudate nucleus and putamen in Huntington's disease grades 2/3 when compared to controls; in the dorsal
caudate few enkephalin or substance P messenger RNA-positive cells were detected. For the early grade
(0/1) Huntington's disease cases, a heterogeneous reduction in both enkephalin and substance P messenger
RNAs was noted; for enkephalin messenger RNA the striatal autoradiograms displayed a conspicuous
patchy appearance. Detailed cellular analysis of the dorsal caudate revealed a striking reduction in the
number of enkephalin and substance P messenger RNA-positive cells detected and in the intensity of
hybridization signal/cell.
These data suggest that both the "indirect" GABA/enkephalin and "direct" GABA/substance P
pathways are perturbed very early in the course of the disease and that these early changes in chemical
signalling may possibly underlie the onset of clinical symptoms.
Key words: basal ganglia, striatal compartments, in situ hybridization, Huntington's disease, neuropeptide
gene expression.
Expansion of a trinucleotide (CAG) glutamine repeat
sequence in excess of 40 repeats within one copy of
the human IT15 gene is now recognized as being the
genetic basis of Huntington's disease (HD), 29 a pro-
gressive neurological disorder characterized clinically
by cognitive impairment, choreiform movements and
akathesia. 34 How expansion of this polyglutamine
sequence, located within the coding region of the
gene, results in this neurological disorder is still
unknown. Pathologically, HD is characterized by a
loss of striatal medium-sized spiny GABA projection
neurons; the earliest signs of cell loss appear in the
dorsomedial caudate and dorsal putamen. 49 These
striatal GABA projection cells comprise at least two
tTo whom correspondence should be addressed.
Abbreviations: ENK, enkephalin; GFAP, glial fibrillary
acidic protein; GPe, globus pallidus externus; GPi,
globus pallidus internus; HD, Huntington's disease;
NADPH, reduced nicotinamide adenine dinucleotide
phosphate; SP, substance P.
chemical phenotypes: one phenotype expresses the
neuropeptide enkephalin (ENK) and the other the
neuropeptides substance P (SP) and dynorphin. TM In
rodents, combined retrograde tracing and in situ
hybridization studies have shown that each of these
two neuropeptide systems predominantly innervates
different target nuclei: GABA/ENK cells innervate
the external pallidal segment (GPe), whilst GABA/SP
cells innervate the entopeduncular nucleus (rat hom-
ologue of the primate internal pallidal segment, GPi)
and the substantia nigra. 18 In non-human primates,
a similar topography of target innervation has
been established, ~5,46 suggesting that the underlying
chemical circuitry of the primate basal ganglia may
be similar, although more complex.
Initial biochemical 7,12 and immunocytochemical
studies of advanced HD cases found a marked re-
duction in GABA in the striatum 47 and in ENK- and
SP-like immunoreactivity ~6in the GPe and substantia
nigra (respectively), providing strong evidence for
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