INTRODUCTION
Exposure of cells to various environmental insults including
heat shock results in the induction of a small group of
conserved proteins known as the heat shock proteins (Hsps)
(reviewed by Morimoto et al., 1994). In addition, some of the
Hsps have been reported to be expressed in the absence of
stress during embryogenesis and development in the fruitfly
Drosophila melanogaster (Dm) (Sirotkin and Davidson, 1982).
In D. melanogaster, the low molecular mass Hsps (s-Hsps:
small heat shock proteins), Hsp27, Hsp26, Hsp23, and Hsp22,
are encoded by four of seven heat-inducible genes clustered at
chromosomal region 67B (Petersen et al., 1979; Ayme and
Tissières, 1985). In addition to being stress-inducible, the
synthesis of s-Hsps can be regulated by the molting hormone
20-hydroxyecdysone as shown in developing imaginal discs
and in cultured cells (Ireland et al., 1982; Sirotkin and
Davidson, 1982; Cheney and Shearn, 1983; Beaulieu et al.,
1989).
While the synthesis of the four s-Hsps of D. melanogaster
is coordinated during heat shock in cultured cells, their
expression during fly development is not (reviewed by Arrigo
and Tanguay, 1991; Arrigo and Landry, 1994; Michaud et al.,
1997). For example, ovarian cells of D. melanogaster contain
mRNAs encoding Hsp27 and Hsp26 that are stored in oocytes
(Graziosi et al., 1980; Zimmerman et al., 1983; Mason et al.,
1984). P-element transformation studies have revealed the
presence of multiple regulatory elements controlling the tran-
scription of these small Hsp genes (Cohen and Meselson, 1985;
Glaser and Lis, 1990). Using a hsp26-lacZ fusion gene, Glaser
et al. (1986) showed that the hsp26 gene was expressed in the
absence of stress in different tissues including gonads and neu-
rocytes. The expression of Hsp27, Hsp26 and Hsp23 in the
central nervous system and in germ lines of D. melanogaster
has also been reported (Pauli et al., 1990; Haass et al; 1990;
Marin et al., 1993, 1996; Marin and Tanguay, 1996).
In addition, each s-Hsp appears to be expressed in specific
cells of some organs. In gonads of Drosophila, Hsp26 mRNA
has been detected in nurse cells of developing ovaries, in
primary spermatocytes as well as in some spermatogonia
(Glaser et al., 1986). Hsp27 and Hsp26 polypeptides have been
detected in specific cells of the germ line and in some somatic
parts of the male reproductive system (Pauli et al., 1990; Marin
et al., 1993). Hsp23 has also been reported to be expressed in
1989 Journal of Cell Science 110, 1989-1997 (1997)
Printed in Great Britain © The Company of Biologists Limited 1997
JCS9612
The developmental and heat-shock-induced expression of
two small heat-shock proteins (Hsp23 and Hsp27) was
investigated during spermatogenesis in Drosophila
melanogaster. Both of these Hsps were expressed in
unstressed and stressed male gonads as shown by
immunoblotting. Immunostaining of whole-mount organs
and thin sections of testes showed that an anti-Hsp23
antibody specifically decorated cells of the somatic lineage,
such as the cyst cells and the epithelial cells of the testis and
of the seminal vesicle. Hsp27 was expressed in some somatic
cells (cyst cells and epithelial cells of the accessory glands)
and, in addition, was also visible in the maturing sperma-
tocytes of the germline. The same cell-specific pattern of
expression was observed after heat shock, and cells which
did not express Hsp23 and Hsp27 in the absence of stress
were similarly unable to mount a heat shock response for
these s-Hsps. However other Hsps such as Hsp70 and
Hsp22 were induced under heat-shock conditions in testes.
Actinomycin D prevented the heat-induced accumulation
of these Hsps indicating that the induction of Hsps was
regulated at the transcriptional level. The heat shock tran-
scriptional factor of Drosophila (DmHSF), present in sig-
nificantly lower amount in testes when compared to other
tissues such as the head, was shown to be required for the
heat activation of Hsp22 and Hsp70. Immunostaining
revealed that HSF expression was restricted to specific cells
such as cyst cells, epithelial pigment cells, spermatogonia
and spermatids but not the primary spermatocytes. These
data show that the expression and induction of the different
small Hsps is regulated in a cell-specific manner under both
normal and heat shock conditions and suggest that factors
other than the DmHSF are involved in this regulation in
male gonads.
Key words: Spermatogenesis, Hsp27, Hsp23, Heat shock,
Drosophila, Heat shock factor
SUMMARY
Cell-specific expression and heat-shock induction of Hsps during
spermatogenesis in Drosophila melanogaster
Sébastien Michaud
1
, Raquel Marin
1
, J. Timothy Westwood
2
and Robert M. Tanguay
1
1
CHUL Research Center (CHUQ) and Laboratory of Cellular and Developmental Genetics, RSVS, Université Laval, Ste-Foy,
Québec, Canada G1K 7P4
2
Department of Zoology, Erindale College, University of Toronto, Mississauga, Ontario, Canada L5L 1C6
*Author for correspondence (e-mail: Robert.Tanguay@rsvs.ulaval.ca)