ORIGINAL ARTICLE
Immunolocalization of steroidogenic enzymes and their
expression during the breeding season in the testes of
wild raccoon dogs (Nyctereutes procyonoides)
Qiang WENG,
1,4
Toshio TSUBOTA,
3
Mingdao DAI,
1
Jiaju WENG,
2
Yang TIAN,
1
Meiyu XU,
1
Gen WATANABE
4,5
and Kazuyoshi TAYA
4,5
1
College of Biological Science and Technology, Beijing Forestry University and
2
School of Basic Medical Sciences,
Peking University, Beijing, China;
3
Laboratory of Wildlife Biology, Graduate School of Veterinary Medicine,
Hokkaido University, Kita, Sapporo,
4
Laboratory of Veterinary Physiology, Department of Veterinary Medicine,
Faculty of Agriculture, Tokyo University of Agriculture and Technology, Fuchu, Tokyo and
5
United Graduate
School of Veterinary Sciences, Gifu University, Gifu, Japan
ABSTRACT
The objective of this study was to investigate immunolocalization of steroidogenic enzymes 3bHSD, P450c17 and
P450arom and their expression during the breeding season in wild male raccoon dogs. The testicular weight, size and
seminiferous tubule diameters were measured, and histological and immunohistochemical observations of testes were
performed. The messenger RNA expression (mRNA) of 3bHSD, P450c17 and P450arom was measured in the testes during
the breeding season. 3bHSD was found in Leydig cells during the breeding and non-breeding seasons with more intense
staining in the breeding season. P450c17 was identified in Leydig cells and spermatids in the breeding season, whereas
it was present only in Leydig cells in the non-breeding season. The localization of P450arom changed seasonally: no
immunostaining in the non-breeding season; more extensive immunostaining in Leydig cells, Sertoli cells and elongating
spermatids in the breeding season. In addition, 3bHSD, P450c17 and P450arom mRNA were also expressed in the testes
during the breeding season. These results suggested that seasonal changes in testicular weight, size and seminiferous
tubule diameter in the wild raccoon dog were correlated with spermatogenesis and immunoreactivity of steroidogenic
enzymes and that steroidogenic enzymes may play an important role in the spermatogenesis and testicular recrudescence
and regression process.
Key words: immunohistochemistry, mRNA, raccoon dog, steroidogenic enzymes, testes.
INTRODUCTION
Reproductive strategies of seasonal breeders are adap-
tations to annual changes in the environment and
they minimize the animals’ energetic efforts for repro-
duction. Mature males show synchronized cycles of
testicular growth and involution between breeding
and non-breeding periods (Bronson & Heidemann
1994). Seasonal changes in spermatogenesis, mor-
phology of the testis and peripheral testosterone con-
centrations have been reported in numerous species
(Komatsu et al. 1997; Tsubota et al. 1997, 2001; Hay-
akawa et al. 2010; Zhang et al. 2010). In these animals,
immunohistochemical studies on testicular tissues
have demonstrated that the distributions of immu-
noreactivities for steroidogenic enzymes in testicular
tissue are associated with seasons and species. Steroid
biosynthesis is accomplished by the actions of two
major families of enzymes. The first are the hydroxy-
lase enzymes, encoded by genes belonging to the
cytochrome P450 superfamily. The second family,
the steroid dehydrogenase enzymes, belong to one
of two distinct groups, the shortchain alcohol
dehydrogenase/reductase family or the aldo-keto
reductase superfamily. Together, the P450 and dehy-
drogenase enzymes are necessary for steroidogensis
in the gonads, placenta and adrenals (Hinshelwood
1998).
Correspondence: Qiang Weng, Laboratory of Animal
Physiology, College of Biological Science and Technology,
Beijing Forestry University, Beijing 100083, China. (Email:
qiangweng@bjfu.edu.cn)
Received 20 April 2011; accepted for publication 26 Septem-
ber 2011.
Animal Science Journal (2012) doi: 10.1111/j.1740-0929.2011.00990.x
© 2012 The Authors
Animal Science Journal © 2012 Japanese Society of Animal Science