生物钟在蛋鸡排卵-产蛋过程中的调控作用

doi:10.3864/j.issn.0578-1752.2018.16.014

摘要/Abstract

摘要： 生物体内源性的昼夜节律使其能够预测周边环境周期性的变化，使机体的内在代谢和周边环境保持一致。在禽类卵泡的成熟、排卵和蛋的形成过程中，不同生理进程在时间上的吻合显示了机体自身以及机体与环境之间的协调统一。动物对营养物质的摄入、内分泌激素的生成、能量代谢等一系列的行为和生理过程都有生物钟参与调控。文章从光照和营养两种因素入手，综述了生物钟在神经内分泌、能量摄入和能量代谢中的调控作用，揭示了蛋鸡的排卵和产蛋机制。1.光信号通过调控生物钟影响下丘脑-垂体-性腺轴（HPG轴），从而调控机体的繁殖活动。在光信号刺激下，位于禽类视交叉上核（SCN）和松果体的中枢生物钟作用于下丘脑，使下丘脑定时性释放促性腺激素释放激素（GnRH）和促性腺激素抑制激素（GnIH），GnRH和GnIH继而作用于垂体调节释放促性腺激素-促黄体生成素（LH）和促卵泡激素（FSH），卵巢中存在的外周生物钟接受中枢的同步化信号来维持生物节律，促使禽类的卵泡成熟和定时排卵；2.除了受到HPG的神经内分泌调控之外，蛋鸡的排卵-产蛋过程还受到机体能量代谢的影响。中枢和外周的生物钟基因能够调控食欲调节系统，从而影响能量摄入；生物钟能够通过调控代谢过程中重要限速酶的表达、整合核受体和营养信号蛋白、调节代谢感受器和代谢物、影响肠道微生物等途径来调节能量代谢，影响卵黄前体物质的合成、转运和沉积；禽类松果体分泌的褪黑素可通过介导降钙素、甲状旁腺素（PTH）及雌激素分泌，节律性地调节体内钙代谢，影响蛋壳的形成。能量摄入的时间和行为、机体能量代谢和能量状态也可以通过腺苷酸活化蛋白激酶（AMPK）、过氧化物酶体增殖物激活受体α（PPARα）等一些与食欲调控和能量代谢相关的细胞因子反过来调控生物钟。营养-生物钟-能量代谢三者之间相互作用，使生物体适应环境的能力增强，能量利用达到最优。因此，通过调整进食时间和食物组分（如饲料能量水平和钙水平），能够改变能量代谢从而调节生物钟的功能。将环境（光照管理）和营养（饲喂时间、饲料配方）综合研究并加以运用，使机体生物钟成为连接外部环境信号和内部能量代谢的纽带，既能响应外界环境刺激，又能同时调控机体能量代谢进程，从而使各项生理功能得到更好地发挥，这将为蛋鸡的产蛋调控机制研究提供新的视角。

关键词: 生物钟, 蛋鸡, 产蛋, 光照, 能量

Abstract: The endogenous circadian rhythm enables the organisms to predict the changes of environmental cycle, which maintains consistency between body metabolism and the external environment. During the maturation of follicular, ovulation, and the formation of egg in birds, the coincidence of the different physiological processes in time shows the unity of the body itself and the coordination between the body and the environment. Biological clock participates in a series of behavior and physiological processes such as nutrition intake, the production of endocrine hormones and energy metabolism. In the present review, the role of biological clock in neuroendocrine, energy intake and energy metabolism has been discussed, from the points of light factor and nutrition factor, to reveal the potential regulating mechanism underlying ovulation and egg laying of hens. (1) Light signal acts on hypothalamic- pituitary-gonadal axis (HPG) by regulating the biological clock to influence reproductive activities. Under the stimulation of light, the central clocks in suprachiasmatic nucleus (SCN) and pineal act on hypothalamus, and make it to release gonadotropin releasing hormone (GnRH) and gonadotropin inhibitory hormones (GnIH) periodically. GnRH and GnIH then act on pituitary, and make it to release gonadotropin hormone, that is luteinizing hormone (LH) and follicle-stimulating hormone (FSH). Periphery clocks in ovary receive the central synchronization signal to maintain the biological rhythm, thereby regulating the maturation of follicles and ovulation. (2) In addition to being regulated by the neuroendocrine system of HPG axis, the ovulation-egg production process of laying hens is also affected by the body's energy metabolism. The central and peripheral clock genes regulate the appetite regulation system and thus affect energy intake; Biological clock can regulate the expression of key enzymes in the process of metabolism, integrate the nuclear receptors and nutrition signaling proteins, regulate metabolism sensors and metabolites, affect gut microbes to regulate energy metabolism, and affect the synthesis, transport and deposition of yolk precursor; Melatonin secreted by bird's pineal can regulate calcium metabolism rhythmically by mediating the secretion of calcitonin, parathyroid hormone (PTH) and estrogen, and influence the formation of egg shell. The time and the behavior of energy intake, the body energy metabolism and energy status can also modulate biological clock, through some appetite regulation and energy metabolism related cytokines such as AMP-activated protein kinase (AMPK), and peroxisome proliferator-activated receptors α (PPARα). There are interactions between nutrient, biological clock and energy metabolism, which accommodate organisms with the surrounding and optimize the energy utilization. Therefore, by adjusting the time of eating and the composition of feed (such as the energy level of feed and calcium level), energy metabolism can be changed to regulate the function of the biological clock. In conclusion, it will provide a new perspective for researching regulation mechanism of egg laying, if we make an integrated study on environment factor (light management) and nutrition (feeding time and feed formula) in which biological clock linked external factors and internal energy metabolism, that is, biological clock can both response to environmental stimuli, and regulate the body's energy metabolism process, to optimize the various physiological functions.

Key words: biological clock, laying hen, egg laying, light, energy

王晓鹃，刘磊，焦洪超，赵景鹏，林海. 生物钟在蛋鸡排卵-产蛋过程中的调控作用[J]. 中国农业科学, 2018, 51(16): 3181-3190.

WANG XiaoJuan, LIU Lei, JIAO HongChao, ZHAO JingPeng, LIN Hai. Regulation of Biological Clock in Ovulation-Laying of Laying Hens[J]. Scientia Agricultura Sinica, 2018, 51(16): 3181-3190.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2018.16.014

https://www.chinaagrisci.com/CN/Y2018/V51/I16/3181

参考文献

[1] 倪银华, 吴涛, 王露, 夏李群, 张丹萍, 傅正伟. 肾上腺糖皮质激素与生物钟基因表达调控的相关研究进展. 遗传, 2008, 30(2): 135-141.

NI Y H, WU T, WANG L, XIA L Q, ZHANG D P, FU Z W. Advances in interactions between glucocorticoid hormones and circadian gene expression. Hereditas (Beijing),2008, 30(2): 135-141. (in Chinese)

[2] WU T, JIN Y, NI Y, ZHANG D, KATO H, FU Z. Effects of light cues on re-entrainment of the food-dominated peripheral clocks in mammals. Gene, 2008, 419(1-2): 27-34.

[3] WU T, JIN Y, KATO H, FU Z. Light and food signals cooperate to entrain the rat pineal circadian system. Journal of Neuroscience Research, 2008, 86(14): 3246-3255.

[4] WU T, DONG Y, YANG Z, KATO H, NI Y, FU Z. Differential resetting process of circadian gene expression in rat pineal glands after the reversal of the light/dark cycle via a 24 h light or dark period transition. Chronobiology International, 2009, 26(5): 793-807.

[5] DONG Y, WU T, NI Y H, KATO H, FU Z W. Effect of fasting on the peripheral circadian gene expression in rats. Biological Rhythm Research, 2010, 41(1): 41-47.

[6] MUKHERJI A, KOBIITA A, CHAMBON P. Shifting the feeding of mice to the rest phase creates metabolic alterations, which, on their own, shift the peripheral circadian clocks by 12 hours. Proceedings of the National Academy of Sciences, 2015, 112(48): E6683- E6690.

[7] IKEDA Y, SASAKI H, OHTSU T, SHIRAISHI T, TAHARA Y, SHIBATA S. Feeding and adrenal entrainment stimuli are both necessary for normal circadian oscillation of peripheral clocks in mice housed under different photoperiods. Chronobiology International, 2015, 32(2): 195-210.

[8] CHAIX A, ZARRINPAR A, MIU P, PANDA S. Time-restricted feeding is a preventative and therapeutic intervention against diverse nutritional challenges. Cell Metabolism, 2014, 20(6): 991-1005.

[9] LARRONDO L F, OLIVARES-YAÑEZ C, BAKER C L, LOROS J J, DUNLAP J C. Circadian rhythms. Decoupling circadian clock protein turnover from circadian period determination. Science, 2015, 347(6221): 1257277.

[10] GWINNER E, BRANDSTATTER R. Complex bird clocks. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 2001, 356(1415): 1801-1810.

[11] UNDERWOOD H, STEELE C T, ZIVKOVIC B. Circadian organization and the role of the pineal in birds. Microscopy Research and Technique, 2001, 53(1): 48-62.

[12] CASSONE V M. Avian Circadian Organization: A Chorus of Clocks. Frontiers in Neuroendocrinology, 2014, 35(1): 76-88.

[13] RUSSO K A. Circadian and Metabolic Influences on the Reproductive Axis. 2016.

[14] SMITH M J JIENNES L, WISE P M. Localization of the VIP2 receptor protein on GnRH neurons in the female rat. Endocrinology, 2000, 141(11): 4317-4320.

[15] KYRIACOU C P. The molecular ethology of the period gene in Drosophila. Behavior Genetics, 1990, 20(2): 191-211.

[16] BERSON D M, DUNN F A, TAKAO M. Phototransduction by retinal ganglion cells that set the circadian clock. Science, 2002, 295(5557): 1070-1073.

[17] SELLIX M T, MENAKER M. Circadian clocks in the ovary. Trends in Endocrinology & Metabolism, 2010, 21(10): 628-636.

[18] 张志超. 时钟基因在母鸡生殖系统中的节律性表达及颗粒细胞中时控基因的筛选[D]. 四川: 四川农业大学, 2016.

ZHANG Z C. A study of circadian clock gene rhythmic expression in hens’ reproductive system and the screening of clock controlled genes in granulose cells [D]. Sichuan: Sichuan Agricultural University, 2016. (in Chinese)

[19] CHEN H, ZHAO L, CHU G, KITO G, YAMAUCHI N, SHIGEYOSHI Y, HASHIMOTO S, HATTORI M A. FSH induces the development of circadian clockwork in rat granulosa cells via a gap junction protein Cx43-dependent pathway. American Journal of Physiology Endocrinology and Metabolism, 2013, 304(6): E566-E575.

[20] CHU G, MISAWA I, CHEN H, YAMAUCHI N, SHIGEYOSHI Y, HASHIMOTO S, HATTORI M A. Contribution of FSH and triiodothyronine to the development of circadian clocks during granulosa cell maturation. American Journal of Physiology Endocrinology and Metabolism, 2012, 302(6): E645-E653.

[21] TISCHKAU S A, HOWELL R E, HICKOK J R, KRAGER S L, BAHR J M. The luteinizing hormone surge regulates circadian clock gene expression in the chicken ovary. Chronobiology International, 2011, 28(1): 10-20.

[22] OLANREWAJU H A, THAXTON J P, DOZIER W A, PURSWELL J. ROUSH W B, BRANTON S L. A review of lighting programs for broiler production. International Journal of Poultry Science, 2006, 5(4): 301-308.

[23] 杨利国. 动物繁殖学. 北京: 中国农业出版社, 2003.

YANG L G. Animal Reproduction. Beijing: China Agriculture Press, 2003. (in Chinese)

[24] HAHN T P, BALL G F. Changes in brain GnRH associated with photorefractoriness in house sparrows (Passer domesticus). General and Comparative Endocrinology, 1995, 99(3): 349-363.

[25] RANI S, SINGH S, MISRA M, KUMAR V. The influence of light wavelength on reproductive photorefractoriness in migratory blackheaded bunting (Emberiza melanocephala). Reproduction Nutrition Development, 2001, 41(4): 277-284.

[26] JOHNSTON J D. Photoperiodic regulation of prolactin secretion: changes in intra-pituitary signalling and lactotroph heterogeneity. Journal of Endocrinology, 2004. 180(3): 351-356.

[27] SHARP P J, BLACHE D. A neuroendocrine model for prolactin as the key mediator of seasonal breeding in birds under long- and short-day photoperiods. Canadian Journal of Physiology and Pharmacology, 2003, 81(4): 350-358.

[28] WANG X J, LI Y, SONG Q Q, GUO Y Y, JIAO H C, SONG Z G, LIN H. Corticosterone regulation of ovarian follicular development is dependent on the energy status of laying hens. The Journal of Lipid Research, 2013, 54(7): 1860-1876.

[29] TAO Z, SONG W, ZHU C, XU W, LIU H, ZHANG S, HUIFANG L. Comparative transcriptomic analysis of high and low egg-producing duck ovaries. Poultry Science, 2017, 96(12): 4378-4388.

[30] ZHANG Z C, WANG Y G, LI L, YIN H D, LI D Y, WANG Y, ZHAO X L, LIU Y P, ZHU Q. Circadian clock genes are rhythmically expressed in specific segments of the hen oviduct. Poultry Science, 2016, 95(7): 1653-1659.

[31] BUYSE J, ADELSOHN D S, DECUYPERE E, SCANES C G. Diurnal-nocturnal changes in food intake, gut storage of ingesta, food transit time and metabolism in growing broiler chickens: a model for temporal control of energy balance. British Poultry Science, 1993, 34(4): 699-709.

[32] VAN DER POL C W, MOLENAAR R, BUITINK C J, VAN ROOVERT-REIJRINK I A, MAATJENS C M, VAN DEN BRAND H, KEMP B. Lighting schedule and dimming period in early life: consequences for broiler chicken leg bone development. Poultry Science, 2015, 94(12): 2980-2988.

[33] FICK L J, FICK G H, BELSHAM D D. Rhythmic clock and neuropeptide gene expression in hypothalamic mHypoE-44 neurons. Molecular and Cellular Endocrinology, 2010, 323(2): 298-306.

[34] KETTNER N M, MAYO S A, HUA J, LEE C, MOORE D D, FU L. Circadian dysfunction induces leptin resistance in mice. Cell Metabolism, 2015, 22(3): 448-459.

[35] YOSHIDA K, MCCORMACK S, ESPA R A, CROCKER A, SCAMMELL T E. Afferents to the orexin neurons of the rat brain. Journal of Comparative Neurology, 2006, 494(5): 845-861.

[36] ESTABROOKE I V, MCCARTHY M T, KO E, CHOU TC, CHEMELLI R M, YANAGISAWA M, SAPER C B, SCAMMELL T E. Fos expression in orexin neurons varies with behavioral state. Journal of Neuroscience, 2001, 21(5): 1656-1662.

[37] KALSBEEK A, YI C X, LA FLEUR S E, FLIERS E. The hypothalamic clock and its control of glucose homeostasis. Trends in Endocrinology and Metabolism, 2010, 21(7): 402-410.

[38] FROY O. Metabolism and circadian rhythms-implications for obesity. Endocrine Reviews, 2010, 31(1): 1-24.

[39] YAN A, ZHANG L, TANG Z, ZHANG Y, QIN C, LI B, LI W, LIN H. Orange-spotted grouper (Epinephelus coioides) orexin: Molecular cloning, tissue expression, ontogeny, daily rhythm and regulation of NPY gene expression. Peptides, 2011, 32(7):1363-1370.

[40] KALSBEEK A, PALM I F, LA FLEUR S E, SCHEER F A, PERREAU-LENZ S, RUITER M, KREIER F, CAILOTTO C, BUIJS R M. SCN outputs and the hypothalamic balance of life. Journal of Biological Rhythms, 2006, 21(6): 458-469.

[41] KENNAWAY D J, VARCOE T J, VOULTSIOS A, BODEN M J. Global loss of bmal1 expression alters adipose tissue hormones, gene expression and glucose metabolism. PLoS One, 2013, 8(6): e65255.

[42] ADAMANTIDIS A, DE LECEA L. The hypocretins as sensors for metabolism and arousal. The Journal of Physiology, 2009, 587(Pt 1): 33-40.

[43] SAPER C B, CHOU T C, ELMQUIST J K. The need to feed: homeostatic and hedonic control of eating. Neuron, 2002, 36(2): 199-211.

[44] ASHER G, SASSONE-CORSI P. Time for Food: The Intimate Interplay between Nutrition, Metabolism, and the Circadian Clock. Cell, 2015, 161(1): 84-92.

[45] VETTER C, SCHEER F A J L.Circadian Biology: Uncoupling Human Body Clocks by Food Timing. Current Biology, 2017, 27(13): R656-R658.

[46] WEHRENS S M T, CHRISTOU S, ISHERWOOD C, MIDDLETON B, GIBBS M A, ARCHER S N, SJENE D J, JOHNSTON J D. Meal timing regulates the human circadian system. Current Biology, 2017, 27(12): 1768-1775.

[47] ZVONIC S, PTITSYN A A, CONRAD S A, SCOTT L K, FLOYD Z E, KILROY G, WU X, GOH B C, MYNATT R L, GIMBLE J M. Characterization of peripheral circadian clocks in adipose tissues. Diabetes, 2006, 55(4): 962-970.

[48] BRAY M S, RATCLIFFE W F, GRENETT M H, BREWER R A, GAMBLE K L, YOUNG M E. Quantitative analysis of light-phase restricted feeding reveals metabolic dyssynchrony in mice. International Journal of Obesity, 2013, 37(6): 843-852.

[49] VOLLMERS C, GILL S, DITACCHIO L, PULIVARTHY S R, LE H D, PANDA S. Time of feeding and the intrinsic circadian clock drive rhythms in hepatic gene expression. Proceedings of the National Academy of Sciences, 2009, 106(50): 21453-21458.

[50] NIELSEN B L, LITHERLAND M, NODDEGAARD F. Effects of qualitative and quantitative feed restriction on the activity of broiler chickens. Applied Animal Behaviour Science, 2003. 83(4): 309-323.

[51] LAMIA K A, SACHDEVA U M, DITACCHIO L, WILLIAMS E C, ALVAREZ J G, EGAN D F, VASQUEZ D S, JUGUILON H, PANDA S, SHAW R J, THOMPSON C B, EVANS R M. AMPK regulates the circadian clock by cryptochrome phosphorylation and degradation. Science, 2009, 326(5951): 437-440.

[52] ANDO H, KUMAZAKI M, MOTOSUGI Y, USHIJIMA K, MAEKAWA T, ISHIKAWA E, FUJIMURA A. Impairment of peripheral circadian clocks precedes metabolic abnormalities in ob/ob mice. Endocrinology, 2011, 152(4): 1347-1354.

[53] CATON P W, KIESWICH J, YAQOOB M M, HOLNESS M J, SUGDEN M C. Metformin opposes impaired AMPK and SIRT1 function and deleterious changes in core clock protein expression in white adipose tissue of genetically-obese db/db mice. Diabetes, Obesity and Metabolism, 2011, 13(12): 1097-1104.

[54] MCCARTHY J J, ANDREWS J L, MCDEARMON E L, CAMPBELL K S, BARBER B K, MILLER B H, WALKER J R, HOGENESCH J B, TAKAHASHI J S, ESSER K A. Identification of the circadian transcriptome in adult mouse skeletal muscle. Physiological Genomics, 2007, 31(1): 86-95.

[55] KALSBEEK A, RUITER M, LA FLEUR S E, CAILOTTO C, KREIER F, BUIJS R M. The hypothalamic clock and its control of glucose homeostasis. Progress in Brain Research, 2006, 153: 283-307.

[56] FROY O. The relationship between nutrition and circadian rhythms in mammals. Frontiers in Neuroendocrinology, 2007, 28(2-3): 61-71.

[57] YANG X, DOWNES M, YU RT, BOOKOUT AL, HE W, STRAUME M, MANGELSDORF D J, EVANS R M. Nuclear receptor expression links the circadian clock to metabolism. Cell, 2006, 126(4): 801-810.

[58] HERICHOVA I, ZEMAN M, JURANI M, LAMOS?OVA D. Daily rhythms of melatonin and selected biochemical parameters in plasma of Japanese quail. Avian and Poultry Biology Reviews, 2004, 15(3-4): 205-210.

[59] SHOSTAK A, Meyer-Kovac J, Oster H. Circadian regulation of lipid mobilization in white adipose tissues. Diabetes, 2013, 62(7): 2195-2203.

[60] TUREK F W, JOSHU C, KOHSAKA A, LIN E, IVANOVA G, MCDEARMON E, LAPOSKY A, LOSEE-OLSON S, EASTON A, JENSEN D R, ECKEL R H, TAKAHASHI J S, BASS J. Obesity and metabolic syndrome in circadian Clock mutant mice. Science, 2005, 308(5724): 1043-1045.

[61] PASCHOS G K, IBRAHIM S, SONG W L, KUNIEDA T, GRANT G, REYES T M, BRADFIELD C A, VAUGHAN C H, EIDEN M, MASOODI M, GRIFFIN J L, WANG F, LAWSON J A, FITZGERALD G A. Obesity in mice with adipocyte-specific deletion of clock component Arntl. Nature Medicine, 2012, 18(12): 1768-1777.

[62] SHIMBA S, ISHII N, OHTA Y, OHNO T, WATABE Y, HAYASHI M, WADA T, AOYAGI T, TEZUKA M. Brain and muscle Arnt-like protein-1 (BMAL1), a component of the molecular clock, regulates adipogenesis. Proceedings of the National Academy of Sciences, 2005, 102(34): 12071-12076.

[63] LAU P, NIXON S J, PARTON R G, MUSCAT G E. RORalpha regulates the expression of genes involved in lipid homeostasis in skeletal muscle cells: Caveolin-3 and CPT-1 are direct targets of ROR. The Journal of Biological Chemistry, 2004, 279(35): 36828-36840.

[64] GARBARINO-PICO E, CARPENTIERI A R, CASTAGNET P I, PASQUARE S J, GIUSTO N M, CAPUTTO B L, GUIDO M E. Synthesis of retinal ganglion cell phospholipids is under control of an endogenous circadian clock: daily variations in phospholipid- synthesizing enzyme activities. Journal of Neuroscience Research, 2004, 76(5): 642-652.

[65] HATORI M, HIROTA T, IITSUKA M, KURABAYASHI N, HARAGUCHI S, KOKAME K, SATO R, NAKAI A, MIYATA T, TSUTSUI K, FUKADA Y. Light-dependent and circadian clock-regulated activation of sterol regulatory element-binding protein, X-box-binding protein 1, and heat shock factor pathways. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(12): 4864-4869.

[66] DEMIERRE M F, HIGGINS P D R, GRUBER S B, HAWK E, LIPPMAN S M. Statins and cancer prevention. Nature Reviews Cancer, 2005, 5(12): 930-942.

[67] CRETENET G, LE CLECH M, GACHON F. Circadian clock-coordinated 12 hr period rhythmic activation of the IRE1α pathway controls lipid metabolism in mouse liver. Cell Metabolism, 2010, 11(1): 47-57.

[68] CANAPLE L, RAMBAUD J, DKHISSI-BENYAHYA O, RAYET B, TAN N S, MICHALIK L, DELAUNAY F, WAHLI W, LAUDET V. Reciprocal regulation of brain and muscle Arnt-like protein 1 and peroxisome proliferator-activated receptor α defines a novel positive feedback loop in the rodent liver circadian clock. Journal of Molecular Endocrinology, 2006, 20(8): 1715-1727.

[69] SCHMUTZ I, RIPPERGER JA, BAERISWYL-AEBISCHER S, ALBRECHT U. The mammalian clock component PERIOD2 coordinates circadian output by interaction with nuclear receptors. Genes & Development, 2010, 24(4): 345-357.

[70] COSTA MJ, SO A Y, KAASIK K, KRUEGER K C, PILLSBURY M L, FU Y H, PTACEK L J, YAMAMOTO K R, FELDMAN B J. Circadian rhythm gene period 3 is an inhibitor of the adipocyte cell fate. The Journal of Biological Chemistry, 2011, 286(11): 9063-9070.

[71] MINAMI Y, KASUKAWA T, KAKAZU Y, IIGO M, SUGIMOTO M, IKEDA S, YASUI A, VAN DER HORST G T, SOGA T, UEDA H R. Measurement of internal body time by blood metabolomics. Proceedings of the National Academy of Sciences, 2009, 106(24): 9890-9895.

[72] UM J H, PENDERGAST J S, SPRINGER D A, FORETZ M, VIOLLET B, BROWN A, KIM M K, YAMAZAKI S, CHUNG J H. AMPK regulates circadian rhythms in a tissue- and isoform-specific manner. PLoS One, 2011, 6(3): e18450.

[73] RODGERS J T, LERIN C, HAAS W, GYGI S P, SPIEGELMAN B M, PUIGSERVER P. Nutrient control of glucose homeostasis through a complex of PGC-1α and SIRT1. Nature, 2005, 434(7029): 113-118.

[74] UM J H, YANG S, YAMAZAKI S, KANG H, VIOLLET B, FORETZ M, CHUNG J H. Activation of 5'-AMP-activated kinase with diabetes drug metformin induces casein kinase Iε (CKIε)-dependent degradation of clock protein mPer2. The Journal of Biological Chemistry, 2007, 282(29): 20794-20798.

[75] NAKAHATA Y, SAHAR S, ASTARITA G, KALUZOVA M, SASSONE-CORSI P. Circadian control of the NAD+ salvage pathway by CLOCK-SIRT1. Science, 2009, 324(5927): 654-657.

[76] RAMSEY K M, YOSHINO J, BRACE C S, ABRASSART D, KOBAYASHI Y, MARCHEVA B, HONG H K, CHONG J L, BUHR E D, LEE C, TAKAHASHI J S, IMAI S, BASS J. Circadian clock feedback cycle through NAMPT-mediated NAD+ biosynthesis. Science, 2009, 324(5927): 651-654.

[77] O'NEILL J S, MAYWOOD E S, CHESHAM J E, TAKAHASHI J S, HASTINGS M H. cAMP-dependent signaling as a core component of the mammalian circadian pacemaker. Science, 2008, 320(5878): 949-953.

[78] SAKKOU M, WIEDMER P, ANLAG K, HAMM A, SEUNTJENS E, ETTWILLER L, TSCHÖP M H, TREIER M. A role for brain-specific homeobox factor Bsx in the control of hyperphagia and locomotory behavior. Cell Metabolism, 2007, 5(6): 450-463.

[79] KOHSAKA A, LAPOSKY A D, RAMSEY K M, ESTRADA C, JOSHU C, KOBAYASHI Y, TUREK F W, BASS J. High-fat diet disrupts behavioral and molecular circadian rhythms in mice. Cell Metabolism, 2007, 6(5): 414-421.

[80] KUDO T, NAKAYAMA E, SUZUKI S, AKIYAMA M, SHIBATA S. Cholesterol diet enhances daily rhythm of Pai-1 mRNA in the mouse liver. American Journal of Physiology. Endocrinology and metabolism, 2004, 287(4): E644- E651.

[81] NIELSEN B L, LITHERLAND M, NODDEGAARD F. Effects of qualitative and quantitative feed restriction on the activity of broiler chickens. Applied Animal Behaviour Science, 2003. 83(4): 309-323.

[82] ASHER G, SASSONECORSI P. Time for Food: The Intimate interplay between nutrition, metabolism, and the circadian clock. Cell, 2015, 161 (1): 84-92.

[83] SUZUKI T. Regulation of intestinal epithelial permeability by tight junctions. Cellular and Molecular Life Sciences, 2013, 70 (4): 631-659.

[84] LIANG X, BUSHMAN F D, FITZGERALD G A. Rhythmicity of the intestinal microbiota is regulated by gender and the host circadian clock. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112 (33): 10479-10484.

[85] THAISS C A, ZEEVI D, LEVY M, SEGAL E, ELINAV E. A day in the life of the meta-organism: diurnal rhythms of the intestinal microbiome and its host. Gut Microbes, 2015, 6(2): 137-142.

[86] VOIGT R M, FORSYTH C B, GREEN S J, MUTLU E, ENGEN P, VITATERNA M H, TUREK F W, KESHAVARZIAN A. Circadian disorganization alters intestinal microbiota. PloS One, 2014, 9(5): e97500.

[87] TRINDER M, BISANZ J E, BURTON J P, REID G. Bacteria Need “Sleep” too?: microbiome circadian rhythmicity, metabolic disease, and beyond. University of Toronto Medical Journal, 2015, 92(3): 52-55.

[88] LIANG X, BUSHMAN F D, FITZGERALD G A. Time in motion: the molecular clock meets the microbiome. Cell, 2014, 159(3): 469-470.

[89] ROSSELOT A E, HONG C I, MOORE S R. Rhythm and bugs: Circadian clocks, gut microbiota, and enteric infections. Current Opinion in Gastroenterology, 2016, 32(1): 7-11.

[90] THAISS C A, ZEEVI D, LEVY M, ZILBERMAN-SCHAPIRA G, SUEZ J, TENGELER A C, ABRAMSON L, KATZ M N, KOREM T, ZMORA N, KUPERMAN Y, BITON I, GILAD S, HARMELIN A, SHAPIRO H, HALPERN Z, SEGAL E, ELINAV E. Transkingdom control of microbiota diurnal oscillations promotes metabolic homeostasis. Cell, 2014, 159(3): 514-529.

[91] ZARRINPAR A, CHAIX A, YOOSEPH S, PANDA S. Diet and feeding pattern affect the diurnal dynamics of the gut microbiome. Cell Metabolism, 2014, 20(6): 1006-1017.

[92] THAISS C A, LEVY M, KOREM T, DOHNALOVA L, SHAPIRO H, JAITIN D A, DAVID E, WINTER D R, GURY-BENARI M, TATIROVSKY E, TUGANBAEV T, FEDERICI S, ZMORA N, ZEEVI D, DORI-BACHASH M, PEVSNER-FISCHER M, KARTVELISHVILY E, BRANDIS A, HARMELIN A, SHIBOLET O, HALPERN Z, HONDA K, AMIT I, SEGAL E, ELINAV E. Microbiota diurnal rhythmicity programs host transcriptome oscillations. Cell, 2016, 167(6): 1495-1510.

[93] MONTAGNER A, KORECKA A, POLIZZI A, LIPPI Y, BLUM Y, CANLET C, TREMBLAY-FRANCO M, GAUTIER-STEIN A, BURCELIN R, YEN Y C, JE H S, AL-ASMAKH M, MITHIEUX G, ARULAMPALAM V, LAGARRIGUE S, GUILLOU H, PETTERSSON S, WAHLI W. Hepatic circadian clock oscillators and nuclear receptors integrate microbiome-derived signals. Scientific Reports, 2016. 6: 20127.

[94] LEONE V, GIBBONS S M, MARTINEZ K, HUTCHISON A L, HUANG E Y, CHAM C M, PIERRE J F, HENEGHAN A F, NADIMPALLI A, HUBERT N, ZALE E, WANG Y, HUANG Y, THERIAULT B, DINNER A R, MUSCH M W, KUDSK K A, PRENDERGAST B J, GILBERT J A, CHANG E B. Effects of diurnal variation of gut microbes and high-fat feeding on host circadian clock function and metabolism. Cell Host & Microbe, 2015, 17(5): 681-689.

[95] LADIZESKY M G，BOGGIO V，ALBORNOZ L E, CASTRILLON P O, MAUTALEN C, CARDINALI D P. Melatonin increases oestradiol-induced bone formation in ovariectomized rats. Journal of Pineal Research, 2003, 34(2): 143-151.

[96] CONTI A, CONCONI S, HERTENS E, SKWARLO-SONTA K, MARKOWSKA M, MAESTRONI J M. Evidence for melatonin synthesis in mouse and human bone marrow cell. Journal of Pineal Research, 2000, 28(4): 193-202.

[97] CUTANDO A，ANEIROS-FERNANDEZ J, LOPEZ-VALVERDE A, ARIAS-SANTIAGO S, ANEIROS-CACHAZA J, REITER R J. A new perspective in Oral health: potential importance and actions of melatonin receptors MT1，MT2，MT3，and RZR/ROR in the oral cavity. Archives of Oral Biology, 2011, 56(10): 944-950.

[98] TAYLOR A C, HORVAT-GORDON M, MOORE A, BARTELL P A. The effects of melatonin on the physical properties of bones and egg shells in the laying hen. Plos one, 2013, 8(2): e55663.

[99] ARYA A K, SACHDEVA N. Parathyroid Hormone (PTH) Assays and Applications to Bone Disease: Overview on Methodology. Part of the Biomarkers in Disease: Methods, Discoveries and Applications book series (BDMDA), 2017: 127-154.

[100] FRASER W D, AHMAD A M, VORA J P. The physiology of the circadian rhythm of parathyroid hormone and its potential as a treatment for osteoporosis. Current Opinion in Nephrology and Hypertension, 2004, 13(4): 437-444.

[101]刘俊美. 蛋种鸡夜间补光的效果. 中国禽业导刊, 2006, 23(22): 30.

LIU J M. The effect of the night replenishing on the egg-laying breeder. Guide to Chinese Poultry, 2006, 23(22): 30. (in Chinese)