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J Biol Chem, Vol. 273, Issue 47, 31191-31194, November 20, 1998
From the Department of Biochemistry, Molecular Biology, and Cell
Biology, Northwestern University, Evanston, Illinois 60208
The mammalian genome encodes a family of lactate
dehydrogenase (LDH) isozymes. Two of these, ldha and
ldhb, are expressed ubiquitously. The ldhc gene
is active only in the germinal epithelium during spermatogenesis. In
our analysis of ldhc gene regulation, we found that a
60-base pair promoter sequence was sufficient for testis-specific
expression in an in vitro transcription assay. To confirm
these findings, a genomic fragment containing 100 base pairs
overlapping the transcription start site was isolated and linked to the
Escherichia coli lacZ gene. We report that this genomic
fragment drives testis-specific expression in transgenic mice. We
conclude that transcription of the transgene and possibly of the
endogenous ldhc gene is restricted to leptotene/pachytene primary spermatocytes.
Spermatogenesis is a complex process requiring the coordinate
expression of a number of genes. One of these, ldhc, encodes the testis-specific isozyme of lactate dehydrogenase
(LDH-C4).1 The
kinetic properties and enzyme-substrate interactions (1) suggest that
LDH-C4 is uniquely suited for satisfying the metabolic requirements of differentiating germ cells and functional spermatozoa. Ldhc is one of several genes that are expressed at various
stages of spermatogenesis and either encode unique proteins or produce mRNAs specific to male germ cells (2). A clone containing the 5'-flanking region of the murine ldhc gene was isolated from
a mouse genomic library. There are no permanent cell lines available for promoter analysis in male germ cells, and reliable methods for
transfecting primary germ cells have not been developed. We have been
able to circumvent this problem successfully with an in
vitro transcription assay. Promoter activity was demonstrated within a 720-bp fragment by in vitro transcription assays in
testis nuclear extract. Liver nuclear extracts significantly repressed ldhc promoter activity in this assay system. Analysis of a
series of deletion mutants revealed that a 60-bp core promoter sequence was sufficient to direct basal, testis-specific transcription (3). To
confirm these findings, a genomic fragment containing approximately 100 bp immediately upstream from the transcription start site was isolated
and linked to the Escherichia coli lacZ reporter gene. We
report that this genomic fragment drives testis-specific Generation of Transgenic Mice--
Two constructs containing
murine ldhc promoter sequences were designed to direct
expression of lacZ. A 720-bp mouse ldhc fragment (
These two plasmid DNAs were digested with KpnI and
HindIII. The 4-kilobase fragments that contain a 100-bp
promoter linked to the lacZ gene were purified from agarose
following electrophoresis and injected into the pronucleus of
fertilized single-cell stage CD-1 eggs. Integration of the transgene
was determined by Southern blot hybridization using a 1.4-kilobase
fragment from the Analysis of
The enzyme activity for Immunohistochemical Analysis--
Testes were removed from adult
mice and decapsulated prior to fixation in Bouin's solution. Fixed
tissues were embedded in paraffin, sectioned, and stained with a rabbit
anti- To determine the pattern of expression of the ldhc gene
in transgenic mice, a 100-bp genomic fragment in pNAss Four founder 100H
Transgenic Mice Demonstrate a Testis-specific Promoter for
Lactate Dehydrogenase, LDHC*
ABSTRACT
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Abstract
Introduction
Procedures
Results
Discussion
References
INTRODUCTION
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Abstract
Introduction
Procedures
Results
Discussion
References
-galactosidase expression in transgenic mice. We conclude that transcription of the transgene is restricted to
leptotene-pachytene-stage primary spermatocytes, even though endogenous
LDH-C4 increases in concentration during spermatogenesis.
EXPERIMENTAL PROCEDURES
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Abstract
Introduction
Procedures
Results
Discussion
References
710 to +12) for construct 100H was amplified by PCR as described previously (3). Similarly, the 700-bp fragment for construct 100
H
was amplified as described in Zhou and Goldberg (4). These two PCR
products were subcloned into the blunted EcoRI site of the
pNAss vector from CLONTECH (GenBank accession
number U02433) to create constructs: 100H (wild type promoter), and
100H (mutant promoter).
-galactosidase gene as a probe. Subsequently,
screening was done by PCR on genomic DNA isolated from mouse tails
using two oligonucleotides derived from the lacZ gene.
-Galactosidase--
Tissues were dissected and
stained for expression of -galactosidase using the method described
by Langford et al. (5). Briefly, the tissues were fixed for
2 h at 4 °C in 0.1 M sodium phosphate (pH 7.3)
containing 2% paraformaldehyde, 0.01% sodium deoxycholate, 0.02%
Nonidet P-40, and 0.2% glutaraldehyde. After three washes in 0.1 M sodium phosphate (pH 7.3), 2 mM
MgCl2, 0.01% sodium deoxycholate, and 0.02% Nonidet P-40,
the tissues were incubated overnight in X-gal staining solution,
including 0.1 M sodium phosphate, 1.3 mM
MgCl2, 3 mM K3Fe(CN)6, 3 mM K4Fe(CN)6, and 1 mg/ml X-gal. To
examine the distribution of the transgene histologically, testes were
frozen rapidly and mounted onto optimal cutting temperature compound.
20 µm sections were cut on a Cryostat and fixed briefly in 2%
paraformaldehyde solution. After incubation in X-gal, the sections were
counterstained with hematoxylin.
-galactosidase was measured using the
Galacto-light kit (Tropix Inc). The tissues were homogenized in 0.1 M potassium phosphate (pH 7.8) containing 0.2% Triton
X-100, 1 mM dithiothreitol, 0.2 mM
phenylmethylsulfonyl fluoride, and 5 µg/ml leupeptin. The tissue
debris were removed by brief centrifugation. The concentration of the
protein was determined by the Bio-Rad assay; equal amounts of protein
from various tissues were used to detect enzyme activity.
-galactosidase antibody (1:500) or anti-mouse
LDH-C4 antibody (1:1000) using the Histostain-SP Kit
according to the manufacturer (Zymed Laboratories Inc.) instructions.
RESULTS
Top
Abstract
Introduction
Procedures
Results
Discussion
References
was purified for microinjection into the pronuclei of single cell fertilized CD-1
mouse eggs. This genomic fragment contains the core promoter sequence,
including the 31-bp palindrome that was found to be necessary for
testis-specific expression (3). The transgene constructs are
illustrated in Fig. 1A. The
100 H construct contains a nearly 100-bp fragment (83/+12) linked
to the lacZ gene. Another construct, 100H was made by
removing most of the palindromic sequence (Fig. 1B).
View larger version (21K):
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Fig. 1.
Diagram of constructs for generating
transgenic mice. A, plasmid 100 H contains E. coli lacZ gene fused to 720-bp ldhc promoter.
HindIII were used to release the 4-kilobase fragment, which
includes only about 100 bp of ldhc promoter and
lacZ gene, and KpnI for microinjection.
100H is similar to 100 H except that most of the palindromic
sequence was deleted. The 31-bp palindromic sequence is shown in
panel B. Most of the 3' nucleotides were removed from the
palindrome in 100H (mutant). The transcription
initiation site is indicated by a dot.
transgenic lines (407, 279, 280, and 281) were
identified by Southern blot and by PCR of tail DNA. Two lines (282 and
283) integrated construct 100H. Expression of the transgene was
detected initially by soaking tissues in solutions containing the
-galactosidase substrate, X-gal. Three of four lines from construct
100H and one of two lines from 100H expressed the
lacZ gene only in the testis, where blue staining was
observed in the seminiferous tubules of the transgenic mice, not in
nontransgenic littermates (Fig. 2). None
of the other tissues from transgenic mice showed -gal activity in
this assay. The expression levels differ by three orders of magnitude
with line 407 showing the most abundant activity with a nearly uniform
staining pattern, whereas line 283 had the lowest level of -gal
activity with a spotty staining pattern (Fig. 2C). Because
line 407 showed the highest expression of the transgene, it was
analyzed in greater detail.
View larger version (62K):
[in a new window]
Fig. 2.
Whole mount X-gal staining of mouse
testis. Mouse testes were removed, fixed, and then soaked
overnight in X-gal solution. A, nontransgenic mouse;
B, 100 H founder 281; C, 100H founder
283.
The tissue specificity of the reporter gene is shown in Fig.
3A by a -galactosidase
enzyme activity assay. A high level of enzyme activity was detected
only in testis homogenate, not in tissue extracts of liver, kidney,
spleen, heart, and brain from line 407 nor from a nontransgenic mouse.
The enzyme activity assays were performed also on line 283 transgenic
for 100H. Compared with nontransgenic controls, the testis showed
2-fold higher activity of the reporter gene, with no activity
detectable in somatic tissue extracts (Fig. 3B).
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To determine whether the transgene expression under the control of the
100-bp promoter followed the endogenous ldhc expression pattern, testis cross-sections from line 407 were analyzed
histochemically. X-gal staining was evident largely in those cells
close to the margin of the tubule but not near the lumen nor in
epididymal spermatozoa (Fig. 4). Specific
localization of transgene expression was accomplished by probing fixed,
paraffin-embedded sections with antibody to -galactosidase. There
was nonspecific staining of the acrosome and tails of the spermatids in
the nontransgenic testis. A strong signal was observed in pachytene
primary spermatocytes (Fig.
5A) of the transgenic testis.
Immunohistochemical localization of endogenous LDH-C4 was
visualized in pre- and post-meiotic stages of spermatogenesis in
sections incubated with specific antibody (Fig. 5B).
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and 
,
respectively. B, immunochemical localization of
LDH-C4 in a nontransgenic mouse testis by antiserum to
LDH-C4. The LDH-C4 level is barely detectable
in primary spermatocytes and increases as spermatogenesis
progresses.
Ldhc mRNA can be detected first in preleptotene
spermatocytes (6). The transcription product reaches a high level in
pachytene spermatocytes and round spermatids and decreases in
elongating spermatids. To determine the timing of transgene expression,
testes from 9- and 12-day old mice were fixed and sectioned. The
positive signal for anti--galactosidase could be seen only in day 12 testis, which is coincident with the appearance of pachytene
spermatocytes (data not shown). X-gal staining of cross-sections of
testis revealed a similar expression pattern in which the reporter gene
was transcribed in the cells along the base of the tubule where,
however, only a few individual cells stained blue.
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DISCUSSION |
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Discussion
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Testis-specific gene expression provides a challenge as well as a unique opportunity to investigate transcriptional regulation. The lack of a reliable and appropriate germ cell line that can be transfected limits the use of conventional techniques to investigate transcriptional regulation in such cells. Transgenic constructs have characterized transcriptional regulation and testis specificity of Prm-1(7), histone H1t (8), Tcp10 (9), proacrosin (10), testis ACE (5), proenkephalin (11), and c-kit (12). Transgenes developed for ectopic overexpression and mis-expression of regulatory proteins, including c-mos (13) and Bcl2 (14, 15), have disrupted spermatogenesis. We have used co-cultures of immortalized Sertoli, peritubular, and Leydig cell lines to support the survival of pachytene spermatocytes in vitro (16), and we have assayed transcription in testis nuclear extracts (4). Mouse and human ldhc genes have been isolated and sequenced, and functional promoter assays using CAT and lacZ reporters have been performed (4, 16). The results indicate that both the mouse and human genes contain several ubiquitous cis-acting elements, but not the same ones. For example, the mouse ldhc gene contains a functional TATA box, but the human gene may not. On the other hand, the human gene, but not the mouse gene, contains six potential Sp1 sites in this region. Nevertheless, the human ldhc promoter sequence is functional as a transgene in the mouse testis and retains stage and tissue specificity (17).
Ldhc promoter sequences specific for mouse testis expression have begun to be identified. Data have been obtained to support the regulatory role of a palindromic sequence overlapping the TATA box and extending 3' to the transcription initiation site. A 103-kDa protein in testis nuclear extract and a 65-kDa protein in liver nuclear extract binds to this palindromic sequence within the 60-bp core promoter region (3). From mutational analyses, we have established the palindrome as a dual function regulatory region necessary for promoter activity in testis nuclear extract and as responsible for repression of the promoter in liver nuclear extract (4). Mutation of the TATA box within the palindromic sequence eliminates in vitro transcription in testis nuclear extract, as does mutation of the CCTGG sequence at the center of the palindrome (Fig. 1B) (4). Mutation of a sequence in the left arm of the palindrome has no effect on testis-specific transcription but allows a low level of transcription in liver nuclear extract (4). Although these results are convincing, a more formal proof of promoter activity and developmental specificity required the analyses of the transgenic animals constructed in the present study.
These data confirm our in vitro results demonstrating that
the ldhc promoter sequence as short as 100 bp is able to
direct testis-specific expression. Conclusions concerning the role of the palindromic sequence in conferring cell-type specificity must be
deferred because only a single mutant transgenic line expressed -galactosidase. Nevertheless, in that one transgenic line, promoter activity was testis-specific, suggesting that more of the promoter sequence than the palindrome is involved in cell-type specificity. For
example, Yang and Thomas (18) and Bonny et al. (19)
demonstrated enhancer activity and possible tissue specificity from
binding interactions of the transcription factor Sp1 within the GC box of the mouse and human ldhc promoters, respectively. This is
consistent with the finding described above that the human
ldhc promoter is functional in the context of the murine
germ cell (17).
An unexpected result in the present study was that the transgene
product was clearly confined to pachytene primary spermatocytes. This
distribution is consistent with the staining pattern seen in Fig. 2,
where tubules contain breaks in the deposition of X-gal reaction
product. This can be explained by the wave of spermatogenesis along a
tubule, where not all stages of the cycle will contain pachytene cells.
The immunohistochemical localization of endogenous LDH-C4
reveals a barely detectable level of the protein in pre-meiotic spermatocytes, with an increased intensity of staining as progression through the spermatogenic process continues. Using a combination of
Northern blot analyses and in situ hybridization techniques, Thomas et al. (20) reported that the ldhc gene
was expressed exclusively during meiosis and spermiogenesis, beginning
in leptotene/zygotene spermatocytes and continuing through to the
elongated spermatids. The most advanced stage at which our transgene
product can be detected is pachytene (Fig. 5). This difference in
expression of the transgene and endogenous ldhc may indicate
that the regulatory elements for postmeiotic expression involve
additional upstream sequence beyond the 100-bp fragment. Alternatively,
the endogenous mRNA may be more stable than that of the transgene,
and therefore may persist post-meiotically. Murine ldhc
mRNA has been shown to be stable during spermatogenesis (21).
Additional transgenic constructs are being prepared to answer these
questions and to elucidate the role of cis-regulation of
ldhc gene expression, in particular, and testis-specific
gene expression, in general, during spermatogenesis.
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FOOTNOTES |
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* This research was supported in part by NICHD, National Institutes of Health Grant HD05863.The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
To whom correspondence should be addressed: Department BMBCB,
Northwestern University, 2153 N. Campus Dr., Evanston, IL 60208. Tel.:
847-491-5416; Fax: 847-467-1380.
The abbreviations used are:
LDH, lactate
dehydrogenase; X-gal, 5-bromo-4-chloro-3-indolyl--D-galactopyranoside; PCR, polymerase chain reaction; bp, base pair(s).
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