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J. Biol. Chem., Vol. 276, Issue 19, 15898-15904, May 11, 2001
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From the Departments of Biology and Kinesiology and Health Science,
York University, Toronto, Ontario M3J 1P3, Canada
Contractile activity induces adaptations in
the expression of genes encoding skeletal muscle mitochondrial
proteins; however, the putative signals responsible for these
adaptations remain unknown. We used electrical stimulation (5 Hz, 65 V)
of C2C12 muscle cells in culture to define some of the mechanisms
involved in contractile activity-induced changes in cytochrome
c gene expression. Chronic contractile activity (4 days, 3 h/day) augmented cytochrome c mRNA by 1.6-fold above
control cells. This was likely mediated by increases in transcriptional
activation, because cells transfected with full-length (
Contractile Activity-induced Transcriptional Activation of
Cytochrome c Involves Sp1 and Is Proportional to
Mitochondrial ATP Synthesis in C2C12 Muscle Cells*
726 base
pairs) or minimal (66 base pairs) cytochrome c
promoter/chloramphenicol acetyltransferase reporter constructs
demonstrated contractile activity-induced 1.5-1.7-fold increases in
the absence of contractile activity-induced increases in mRNA
stability. Transcriptional activation of the 726 promoter was
abolished when muscle contraction was inhibited at various subcellular
locations by pretreatment with either the Na+ channel
blocker tetrodotoxin, the intracellular Ca2+ chelator
1,2-bis(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid tetra(acetoxymethyl) ester, or the myosin ATPase inhibitor 2,3-butanedione monoxime. It was further reduced in unstimulated cells
when mitochondrial ATP synthesis was impaired using the uncoupler
2,4-dinitrophenol. Because the contractile activity-induced response
was evident within the minimal promoter, electromobility shift assays
performed within the first intron (+75 to +104 base pairs) containing
Sp1 sites revealed an elevated DNA binding in response to contractile
activity. This was paralleled by increases in Sp1 protein levels. Sp1
overexpression studies also led to increases in cytochrome
c transactivation and mRNA levels. These data suggest
that variations in the rate of mitochondrial ATP synthesis are
important in determining cytochrome c gene expression in
muscle cells and that this is mediated, in part, by Sp1-induced increases in cytochrome c transcription.
*
This work was supported by a grant from the Natural Sciences
and Engineering Council of Canada.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: Dept. of Biology, York
University, North York, ON M3J 1P3, Canada. Tel.: 416-736-2100, Ext. 66640; Fax: 416-736-5698; E-mail: dhood@yorku.ca.
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