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J. Biol. Chem., Vol. 278, Issue 42, 41519-41527, October 17, 2003
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From the aClock Cell Biology Research Group, Institute for Biological Resources and Functions, National Institute of Advanced Industrial Science and Technology, Central 6, 1-1-1 Higashi, Tsukuba, Ibaraki 305-8566, cComputational Biology Research Center, National Institute of Advanced Industrial Science and Technology, Aomi Frontier Building, 17F, 2-43 Aomi, Koto-Ku, Tokyo 135-0064, the dFirst Laboratory for Human Gene Research, Department of Human Gene Research, Kazusa DNA Research Institute, 2-6-7 Kazusa-Kamatari, Kisarazu, Chiba 292–0818, eClinical Molecular Biology, Faculty of Pharmaceutical Sciences, Teikyo University, 1091-1 Suarashi, Sagamiko, Tsukui, Kanagawa 199-0195, fDepartment of Surgery and Clinical Oncology, Graduate School of Medicine, Osaka University, 2-2 Yamadaoka, Suita, Osaka 565-0871, gDepartment of Hygiene and Public Health, School of Human Sciences, Waseda University, 2-579-15 Mikajima, Tokorozawa, Saitama 359-1192, hDepartment of Biomolecular Engineering, Tokyo Institute of Technology, Yokohama, Kanagawa 226-8501, iHorikoshi Gene Selector Project, Exploratory Research for Advanced Technology, Japan Science and Technology Corporation, 5-9-6 Tokodai, Tsukuba, Ibaraki 300-2635, jRadiation Biology Center, Kyoto University, Yoshidakonoe-Cho, Sakyo-Ku, Kyoto, Kyoto 606-8501, the kInstitute of Applied Biochemistry, University of Tsukuba, Tsukuba, Ibaraki 305-8502, and the mLaboratory of Developmental Biology, Institute of Molecular and Cellular Biosciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo-ku, Tokyo 113-0032, Japan
CLOCK is a positive component of a transcription/translation-based negative feedback loop of the central circadian oscillator in the suprachiasmatic nucleus in mammals. To examine CLOCK-regulated circadian transcription in peripheral tissues, we performed microarray analyses using liver RNA isolated from Clock mutant mice. We also compared expression profiles with those of Cryptochromes (Cry1 and Cry2) double knockout mice. We identified more than 100 genes that fluctuated from day to night and of which expression levels were decreased in Clock mutant mice. In Cry-deficient mice, the expression levels of most CLOCK-regulated genes were elevated to the upper range of normal oscillation. Most of the screened genes had a CLOCK/BMAL1 binding site (E box) in the 5'-flanking region. We found that CLOCK was absolutely concerned with the circadian transcription of one type of liver genes (such as DBP, TEF, and Usp2) and partially with another (such as mPer1, mPer2, mDec1, Nocturnin, P450 oxidoreductase, and FKBP51) because the latter were damped but remained rhythmic in the mutant mice. Our results showed that CLOCK and CRY proteins are involved in the transcriptional regulation of many circadian output genes in the mouse liver. In addition to being a core component of the negative feedback loop that drives the circadian oscillator, CLOCK also appears to be involved in various physiological functions such as cell cycle, lipid metabolism, immune functions, and proteolysis in peripheral tissues.
Received for publication, May 1, 2003 , and in revised form, June 24, 2003.
* This work was supported by Grant-in-aid for Young Scientists (B) 15700492 from the Ministry of Education, Culture, Sports, Science, and Technology (MEXT) (to K. O.), by a grant for Exploratory Research for Advanced Technology of the Japan Science and Technology Corporation (to M. H.), by a grant for the support of young researchers with a term from MEXT (to K. M.), and by the Industrial Technology Research grant program from the New Energy and Industrial Technology Development Organization of Japan (to K. M.). 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.
The Gene Expression Omnibus (GEO) accession numbers for the expression data of this work are GSM6792–7 (www.ncbi.nlm.nih.gov/geo).
b These authors contributed equally to this work.
l To whom correspondence should be addressed: Clock Cell Biology Group, National Institute of Advanced Industrial Science and Technology, Central 6, 1-1-1, Higashi, Tsukuba, Ibaraki 305-8566, Japan. Tel.: 81-29-861-6053; Fax: 81-29-861-9499; E-mail: n.ishida{at}aist.go.jp.
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