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J. Biol. Chem., Vol. 279, Issue 33, 34411-34420, August 13, 2004

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The Direct Peroxisome Proliferator-activated Receptor Target Fasting-induced Adipose Factor (FIAF/PGAR/ANGPTL4) Is Present in Blood Plasma as a Truncated Protein That Is Increased by Fenofibrate Treatment^*

Stéphane Mandard, Fokko Zandbergen, Nguan Soon Tan, Pascal Escher¶, David Patsouris, Wolfgang Koenig||, Robert Kleemann**, Arjen Bakker, Frank Veenman, Walter Wahli, Michael Müller, and Sander Kersten

From the Nutrition, Metabolism and Genomics Group, Division of Human Nutrition, Wageningen University, 6700 EV, Wageningen, The Netherlands, Center for Integrative Genomics, Université de Lausanne, Lausanne CH-1015, Switzerland, ^¶Institute of Physiology, Pharmazentrum, University of Basel, Basel CH-4056, Switzerland, the ^||Department of Internal Medicine II-Cardiology, University of Ulm, Ulm D-89081, Germany, ^**Gaubius Laboratory, TNO-Prevention and Health, Leiden 2301 CE, The Netherlands, and the Department of Human Biology, Maastricht University, Maastricht 6200 MD, The Netherlands

The fasting-induced adipose factor (FIAF, ANGPTL4, PGAR, HFARP) was previously identified as a novel adipocytokine that was up-regulated by fasting, by peroxisome proliferator-activated receptor agonists, and by hypoxia. To further characterize FIAF, we studied regulation of FIAF mRNA and protein in liver and adipose cell lines as well as in human and mouse plasma. Expression of FIAF mRNA was up-regulated by peroxisome proliferator-activated receptor (PPAR) and PPAR/ agonists in rat and human hepatoma cell lines and by PPAR and PPAR/ agonists in mouse and human adipocytes. Transactivation, chromatin immunoprecipitation, and gel shift experiments identified a functional PPAR response element within intron 3 of the FIAF gene. At the protein level, in human and mouse blood plasma, FIAF was found to be present both as the native protein and in a truncated form. Differentiation of mouse 3T3-L1 adipocytes was associated with the production of truncated FIAF, whereas in human white adipose tissue and SGBS adipocytes, only native FIAF could be detected. Interestingly, truncated FIAF was produced by human liver. Treatment with fenofibrate, a potent PPAR agonist, markedly increased plasma levels of truncated FIAF, but not native FIAF, in humans. Levels of both truncated and native FIAF showed marked interindividual variation but were not associated with body mass index and were not influenced by prolonged semistarvation. Together, these data suggest that FIAF, similar to other adipocytokines such as adiponectin, may partially exert its function via a truncated form.

Received for publication, March 19, 2004 , and in revised form, June 8, 2004.

^* This work was supported by the Netherlands Organization for Scientific Research, with additional support by the Royal Netherlands Academy of Art and Sciences, the Dutch Diabetes Foundation, the Wageningen Center for Food Sciences, the Swiss National Science Foundation, and the Human Frontier Science Program. The costs of publication of this article were defrayed in part by the payment of page charges. This 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: Nutrition, Metabolism and Genomics Group, Division of Human Nutrition, Wageningen University, P. O. Box 8129, 6700 EV Wageningen, The Netherlands. Tel.: 31-317-48-57-87; Fax: 31-317-48-33-42; E-mail: sander.kersten{at}wur.nl.

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