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Originally published In Press as doi:10.1074/jbc.M203997200 on May 6, 2002

J. Biol. Chem., Vol. 277, Issue 29, 26089-26097, July 19, 2002
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Fatty Acid Homeostasis and Induction of Lipid Regulatory Genes in Skeletal Muscles of Peroxisome Proliferator-activated Receptor (PPAR) alpha  Knock-out Mice
EVIDENCE FOR COMPENSATORY REGULATION BY PPARdelta *

Deborah M. MuoioDagger §, Paul S. MacLean, David B. Lang, Shi LiDagger , Joseph A. Houmard, James M. Way||, Deborah A. Winegar||, J. Christopher Corton**, G. Lynis Dohm, and William E. KrausDagger

From the Dagger  Departments of Medicine and Cell Biology, Duke University Medical Center, Durham, North Carolina 27710, the  Department of Biochemistry and the Human Performance Laboratory, East Carolina University, Greenville, North Carolina 27858, the || Departments of Metabolic Diseases and Nuclear Receptor Biology, GlaxoSmithKline, Research Triangle Park, North Carolina 27709, and the ** Chemical Industry Institute of Toxicology Centers for Health Research, Research Triangle Park, North Carolina 27709

Ablation of peroxisome proliferator activated receptor (PPAR) alpha , a lipid-activated transcription factor that regulates expression of beta -oxidative genes, results in profound metabolic abnormalities in liver and heart. In the present study we used PPARalpha knockout (KO) mice to determine whether this transcription factor is essential for regulating fuel metabolism in skeletal muscle. When animals were challenged with exhaustive exercise or starvation, KO mice exhibited lower serum levels of glucose, lactate, and ketones and higher nonesterified fatty acids than wild type (WT) littermates. During exercise, KO mice exhausted earlier than WT and exhibited greater rates of glycogen depletion in liver but not skeletal muscle. Fatty acid oxidative capacity was similar between muscles of WT and KO when animals were fed and only 28% lower in KO muscles when animals were starved. Exercise-induced regulation and starvation-induced regulation of pyruvate-dehydrogenase kinase 4 and uncoupling protein 3, two classical and robustly responsive PPARalpha target genes, were similar between WT and KO in skeletal muscle but markedly different between genotypes in heart. Real time quantitative PCR analyses showed that unlike in liver and heart, in mouse skeletal muscle PPARdelta is severalfold more abundant than either PPARalpha or PPARgamma . In both human and rodent myocytes, the highly selective PPARdelta agonist GW742 increased fatty acid oxidation about 2-fold and induced expression of several lipid regulatory genes, including pyruvate-dehydrogenase kinase 4 and uncoupling protein 3, responses that were similar to those elicited by the PPARalpha agonist GW647. These results show redundancy in the functions of PPARs alpha  and delta  as transcriptional regulators of fatty acid homeostasis and suggest that in skeletal muscle high levels of the delta -subtype can compensate for deficiency of PPARalpha .

* This work was supported by National Institutes of Health Grants 46121-06 (to G. L. D.), F32DK 10017-01 (to D. M. M.) and HL57354 (to W. E. K.) and a grant from the North Carolina Institute of Nutrition (to D. M. M. and J. A. H.).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: P.O. Box 3327, Duke University Medical Center, Durham, NC 27710. Tel.: 919-684-3644; Fax: 919-684-8907; E-mail: muoio@duke.edu.

Copyright © 2002 by The American Society for Biochemistry and Molecular Biology, Inc.
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