This Article Full Text Full Text (PDF) Supplemental Data Alert me when this article is cited Alert me if a correction is posted Citation Map Services Email this article to a friend Similar articles in this journal Similar articles in PubMed Alert me to new issues of the journal Download to citation manager Request Copyright Permission Citing Articles Citing Articles via HighWire Citing Articles via Google Scholar Google Scholar Articles by Nakao, N. Articles by Yoshimura, T. Search for Related Content PubMed PubMed Citation Articles by Nakao, N. Articles by Yoshimura, T.

Circadian Clock Gene Regulation of Steroidogenic Acute Regulatory Protein Gene Expression in Preovulatory Ovarian Follicles

Division of Biomodeling (N.N., S.Y., A.N., T.Ya., T.W., T.A., S.E., T.Yo.), Graduate School of Bioagricultural Sciences, and Institute for Advanced Research (T.Yo.), Nagoya University, Chikusa-ku, Nagoya 464-8601, Japan; Department of Biophysics and Biochemistry (T.O., Y.F.), Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo 113-0033, Japan; and Roslin Institute (P.J.S.), Roslin, Midlothian EH25 9PS, United Kingdom

Address all correspondence and requests for reprints to: Takashi Yoshimura, Ph.D., Division of Biomodeling, Graduate School of Bioagricultural Sciences and Institute for Advanced Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya 464-8601, Japan. E-mail: takashiy{at}agr.nagoya-u.ac.jp.

It is now known that circadian clocks are localized not only in the central pacemaker but also in peripheral organs. An example of a clock-dependent peripheral organ is the ovary of domestic poultry in which ovulation is induced by the positive feedback action of ovarian progesterone on the neuroendocrine system to generate a preovulatory release of LH during a daily 6–10 h "open period" of the ovulatory cycle. It has been assumed previously that the timing of ovulation in poultry is controlled solely by a clock-dependent mechanism within the neuroendocrine system. Here, we question this assumption by demonstrating the expression of the clock genes, Per2 (Period 2) and Per3, Clock, and Bmal1 (brain and muscle Arnt-like protein 1), in preovulatory follicles in laying quail. Diurnal changes in Per2 and Per3 expression were seen in the largest preovulatory follicle (F1) but not in smaller follicles. We next sought to identify clock-driven genes in preovulatory follicles focusing on those involved in the synthesis of progesterone. One such gene was identified, encoding steroidogenic acute regulatory protein (StAR), which showed 24-h changes in expression in the F1 follicle coinciding with those of Per2. Evidence that StAR gene expression is clock driven was obtained by showing that its 5' flanking region contains E-box enhancers that bind to CLOCK/BMAL1 heterodimers to activate gene transcription. We also showed that LH administration increased the promoter activity of chicken StAR. We therefore suggest that the timing of ovulation in poultry involves an LH-responsive F1 follicular clock that is involved in the timing of the preovulatory release of progesterone.

This article has been cited by other articles:

				W. P. Williams III, E. M. Gibson, C. Wang, S. Tjho, N. Khattar, G. E. Bentley, K. Tsutsui, and L. J. Kriegsfeld Proximate mechanisms driving circadian control of neuroendocrine function: Lessons from the young and old Integr. Comp. Biol., November 1, 2009; 49(5): 519 - 537. [Abstract] [Full Text] [PDF]

				T. Yoshikawa, M. Sellix, P. Pezuk, and M. Menaker Timing of the Ovarian Circadian Clock Is Regulated by Gonadotropins Endocrinology, September 1, 2009; 150(9): 4338 - 4347. [Abstract] [Full Text] [PDF]

				I. K. Johnsen, R. Kappler, C. J. Auernhammer, and F. Beuschlein Bone Morphogenetic Proteins 2 and 5 Are Down-regulated in Adrenocortical Carcinoma and Modulate Adrenal Cell Proliferation and Steroidogenesis Cancer Res., July 15, 2009; 69(14): 5784 - 5792. [Abstract] [Full Text] [PDF]

				C. Torres-Farfan, L. Abarzua-Catalan, F. J. Valenzuela, N. Mendez, H. G. Richter, G. J. Valenzuela, and M. Seron-Ferre Cryptochrome 2 Expression Level Is Critical for Adrenocorticotropin Stimulation of Cortisol Production in the Capuchin Monkey Adrenal Endocrinology, June 1, 2009; 150(6): 2717 - 2722. [Abstract] [Full Text] [PDF]

				C. K. Ratajczak, K. L. Boehle, and L. J. Muglia Impaired Steroidogenesis and Implantation Failure in Bmal1-/- Mice Endocrinology, April 1, 2009; 150(4): 1879 - 1885. [Abstract] [Full Text] [PDF]

				P. Bebas, C. P. Goodall, M. Majewska, A. Neumann, J. M. Giebultowicz, and P. E. Chappell Circadian clock and output genes are rhythmically expressed in extratesticular ducts and accessory organs of mice FASEB J, February 1, 2009; 23(2): 523 - 533. [Abstract] [Full Text] [PDF]

				G. H. Son, S. Chung, H. K. Choe, H.-D. Kim, S.-M. Baik, H. Lee, H.-W. Lee, S. Choi, W. Sun, H. Kim, et al. Adrenal peripheral clock controls the autonomous circadian rhythm of glucocorticoid by causing rhythmic steroid production PNAS, December 30, 2008; 105(52): 20970 - 20975. [Abstract] [Full Text] [PDF]

				T. J. Nakamura, M. T. Sellix, M. Menaker, and G. D. Block Estrogen directly modulates circadian rhythms of PER2 expression in the uterus Am J Physiol Endocrinol Metab, November 1, 2008; 295(5): E1025 - E1031. [Abstract] [Full Text] [PDF]

				J.D. Alvarez, A. Hansen, T. Ord, P. Bebas, P. E. Chappell, J. M. Giebultowicz, C. Williams, S. Moss, and A. Sehgal The Circadian Clock Protein BMAL1 Is Necessary for Fertility and Proper Testosterone Production in Mice J Biol Rhythms, February 1, 2008; 23(1): 26 - 36. [Abstract] [PDF]

				G. F. Ball The Ovary Knows More than You Think! New Views on Clock Genes and the Positive Feedback Control of Luteinizing Hormone Endocrinology, July 1, 2007; 148(7): 3029 - 3030. [Full Text] [PDF]