绵羊常年发情的内分泌学研究及卵巢差异表达基因筛选
详细信息    本馆镜像全文|  推荐本文 |  |   获取CNKI官网全文
摘要
绵羊属于季节性发情动物,一般在短日照的秋冬季节发情配种。季节性发情不仅限制了母羊的生产力,还导致季节性产羔,致使产肉性能也呈现出一定的季节性,造成肥羔肉不能实现一年四季均衡供应。我国大部分绵羊品种表现为季节性发情,因此季节性发情成为我国养羊业高效和可持续发展以及实现国家养羊业发展模式战略转变的瓶颈。然而,要从分子水平上改变绵羊季节性发情的特性,使其能常年发情和配种,就必须首先了解常年发情的内分泌学机制和涉及的关键基因。
     本文旨在通过检测不同季节绵羊外周血液中褪黑激素(MLT)和雌二醇(E2)、孕酮(P4)、促卵泡激素(FSH)、促黄体激素(LH)等生殖激素的变化,同时检测绵羊卵巢组织不同繁殖状态下相关基因表达模式,期望揭示绵羊繁殖季节性的内分泌变化规律和卵巢组织中的关键基因,为进一步阐明绵羊常年发情的分子机理提供理论基础。
     采集小尾寒羊和滩羊春夏秋冬四个季节各一个完整发情周期的血液,检测褪黑激素(MLT)和上述4种生殖激素的浓度。春季采集小尾寒羊不同发情阶段(发情间期末、发情前期、发情期)和滩羊乏情期的卵巢组织,利用高通量测序技术RNA-Seq进行转录组水平检测。本实验获得结果如下:
     (1)外周血液中褪黑激素(MLT)分泌呈现一定的变化规律。MLT在夜间分泌极显著高于白昼(P<0.01),上升与下降的速度非常快,往往在日落或日出1小时左右即可完成,之后一直保持在高水平或低水平状态直至日出或日落时。小尾寒羊MLT分泌没有明显的季节性变化,而滩羊则具有显著的季节性变化特征:秋分和冬至显著高于春分和夏至(P <0.05)。进一步分析了MLT白天和夜间的季节性变化规律,白天MLT水平小尾寒羊四季变化平稳,滩羊在秋冬季节有上升趋势,但变化不显著;夜间MLT小尾寒羊四季变化平稳,滩羊秋冬季节显著上升(P<0.05)。结果提示夜间MLT水平对调控绵羊的季节性发情具有重要作用。
     (2)外周血液中生殖激素的分泌具有鲜明的季节性变化特征。小尾寒羊外周血液中四种生殖激素在春、夏、秋、冬四个季节变化基本稳定。滩羊生殖激素变化具有显著的季节性,E2变化基本稳定;P4分泌量秋冬季节显著高于春夏季节(P <0.05);FSH和LH分泌量秋冬季节显著高于春季和夏季(P <0.05),夏季显著高于春季(P <0.05)。通过对两品种间激素变化研究发现,小尾寒羊与滩羊四个季节血液中E2浓度差异不显著(P>0.05);春夏季节时小尾寒羊P4和FSH含量显著高于滩羊(P <0.05);小尾寒羊LH含量在四个季节均高于滩羊,但仅在冬季时差异显著(P <0.05)。上述结果表明,稳定的E2水平及一定浓度的P4可能与绵羊的常年发情有关,高水平的FSH和LH对开启或维持绵羊的发情具有重要作用,或与多胎具有一定的关系。
     (3)本研究利用RNA-seq技术获得了绵羊不同繁殖状态下的基因表达模式。卵巢的聚类结果表明,乏情期与发情间期,发情前期与发情期分别首先聚在一起。这表明处于不同繁殖状态的绵羊,乏情期与发情间期,发情前期与发情期具有更相似的基因表达模式。
     (4)不同繁殖状态下(乏情期、发情间期末、发情前期、发情期)卵巢共获得差异基因767个,其中已知基因396个、未知371个,表达上调的有702个、下调的有65个。对已知的差异表达基因进行GO功能富集和KEGG Pathway分析。由GO功能分类可知,卵巢组织不同繁殖状态差异表达基因功能主要涉及结合活性、细胞器官、细胞、催化活性、生物调控、生物过程调控、代谢过程等类型。KEGG pathway富集分析表明这些基因主要参与了TGF信号通路、类固醇的生物合成、Notch信号通路、黄酮及黄酮醇的生物合成、孕酮介导的卵母细胞成熟、胰岛素信号通路等调控途径。其中TGF-β信号通路中FSTL3、SMAD1、SMAD4和TGFBRII基因及胰岛素信号通路中的BAD基因可能对常年发情有一定的调控作用,需要进一步的分析验证。
Sheep are seasonal breeders. The breeding season begins at the end of summer or at the beginningof spring. This seasonal estrus limited the ewe productivity and leads to the seasonal lambing, so theprime lamb provided has the seasonal variations depend on the sheep seasonal estrus. Most of sheep areseasonal breeders in China, which limited the efficient and sustainable development of the sheepindustry. In order to change the seasonal estrus at the genetic level, the endocrine basis and the keygenes related to seasonal estrus should be understood firstly.
     The objective of this study is to analyze the seasonal changes of year-round and seasonal estrussheep.Melatonin, E2, P4, FSH and LH concentration in serum and the gene expression pattern of ovaryin different reproductive status were determined. Our results could provide theoretical basis forregulating the seasonal reproduction in sheep.
     Blood samples were collected from five Small Tail Han sheep and five Tan sheep in spring,summer, autumn and winter, respectively. The melatonin, E2, P4, FSH and LH in serum were measuredby radioimmunoassay. Ovaries were collected from Small Tail Han sheep in the different estrous periodincluding the end of dioestrus, proestrus and estrus, and Tan sheep during the anestrous in spring. TheRNA-Seq was used to screen the differently expressed genes in ovary. Our results were listed as follows:
     (1) Secretion pattern of Serum MLT. MLT level at night was significantly higher (P<0.05) than that inthe day. The increase and decrease of the speed was fastand normally completed in1hours.Then theMLT remained at either a high level or low level until sunrise or sunset. The MLT level of Small TailHan sheep showed no significant changes around the year. However, it was different in Tan sheep. MLTlevel in the autumnal equinox and winter solstice were significantly higher (P<0.05) than that in springequinox and summer solstice. The day and night MLT levels of Small Tail Han sheep showed nosignificant changes during four seasons. However, the night MLT level of Tan sheep showedsignificantly increased (P<0.05) in autum and winter. The results indicated that the night MLT levelplays an important role in seasonal reproduction regulation in sheep.
     (2) Secretion pattern of Serum reproductive hormone. The results showed that the concentrations ofthese four hormones in Small Tail Han sheep had no significant changes around the year. However, itwas different in Tan sheep. P4, FSH and LH concentration in estrous season were significantly higher(P<0.05) than those in anestrous season. According to the analysis between two breeds, E2level has nosignificant difference(P>0.05)around the year. The serum P4and FSH concentration in Small TailHan sheep were significantly higher (P<0.05) than in Tan sheep during spring and summer. The LHlevel in Small Tail Han sheep was higher than in Tan sheep, however, has no significantdifference(P>0.05) except in winter. These results indicated that a certain level of E2and P4wereneeded for sheep year-round estrous, and high level of FSH and LH were necessary for year-roundestrus or polyembryony in sheep.
     (3) RNA-Seq was used for determining the gene expression pattern in the different reproductive status.According to the hierachical clustering results of differentially expressed genes from the different reprodctive status. The results showed that the genes of anestrus and diestrus, proestrus and estrus wereclustered together firstly. These indicated that sheep in anestrus and diestrus, proestrus and estrus had asimilar gene expression pattern.
     (4) RNA-Seq analysis of differentially expressed genes generated a list of767genes with396as knownand371as unknown, and with702as up-regulated and65as down-regulated. The most significantlyoverrepresented GO terms are related to binding, organell, cell, cell part, cell process and metabolicprocess, etc. Moreover, KEGG pathway analysis revealed that many pathways such as TGFβ signalingpathway, Steriod biosynthesis, insulin signaling pathway, Notch signaling pathway, Flavone andflavonol biosynthesis as well as Progesterone-mediated oocyte maturation, etc. had been regulatedremarkably at the different reproductive status of sheep ovary. And FSTL3, SMAD1, SMAD4andTGFBRII and BAD gene may the important candidate gene for year-round estrus regulation.
引文
1.程瑞禾,韦来红,张泉福.湖羊考力代羊发情周期外周血液中LH禾孕酮含量的变化.南京农业大学学报[J],1991,14(3):126-127.
    2.高云芳,高更更,白绪祥,张新利,陈超,李保国.雌性川金丝猴尿液中雌二醇与孕酮水平的季节性变化[J].西北大学学报(自然科学版),2005,35(5):592-596.
    3.葛晨霞,王龙涛,李向军,贾博寅,马磊,姜怀志. Bad基因在黄体期绵羊生殖器官中表达的比较研究[J].中国兽医学报,2011,31(3):428-432.
    4.黄冬维,储明星.动物季节性繁殖分子调控机理研究进展[J].遗传,2011,(07):35-46.
    5.侯衍猛,曹洪防,徐云华,钟为,王树迎.莱芜黑山羊发情周期中FSH、LH、E2和P的分泌规律[J].中国兽医学报,2006,26(3):340-343.
    6.刘兰平,何亚维.波尔山羊繁殖周期外周血浆中生殖激素的变化[J].畜禽业,2004,(11):42-42.
    7.罗庆苗.利用数字基因表达谱对小尾寒羊高繁性状相关基因的筛选研究[D].北京:中国农业科学院,2012.
    8.赖平,王凭青,张宝云,储明星,刘重旭,谭颖,樊奇.哺乳动物季节性繁殖的神经内分泌调节机制[J].遗传,2012,(03):281-288.
    9.马友记,李发弟,张利.褪黑素对动物生殖调控机理的研究进展[J].中国草食动物,2007,85-88.
    10.毛文智.绵羊生殖周期卵巢形态学特点的研究[D].吉林:吉林农业大学,2007.
    11.彭夏雨,杨梅,陈童,汪立芹,郭志勤.动态添加FSH对绵羊卵巢皮质组织体外培养的影响[J].畜牧兽医学报,2010,41(11):1387-1393.
    12.权凯,黄炎坤,张长兴.青海高原半细毛羊同期发情的研究[J].黑龙江畜牧兽医,2005,(7):30-31.
    13.祁云霞,刘永斌,荣威恒.转录组研究新技术:RNA-Seq及其应用[J].遗传,2011,(11):1191-1202.
    14.王金富,冷青文,邓湘泉,孔繁军,杨勇军.中国美利奴军垦型细毛羊正常发情周期内血清孕酮含量的变化[J].四川畜牧兽医,2000,27(11):22-22.
    15.王建民,秦孜娟,曲绪仙,王中华,李福昌.山东小尾寒羊种质特性及利用途径的研究进展(上)[J].草食家畜,1997,(3):6-10.
    16.王训立,周建华,范春梅.季节性变化对雌性恒河猴生殖功能的影响[J].中国实验动物学报,2002,10(1):21-26.
    17.王曦,汪小我,王立坤,冯智星,张学工.新一代高通量RNA测序数据的处理与分析[J].生物化学与生物物理进展,2010,(08):834-846.
    18.徐业芬. Smad信号转导通路相关基因表达与湖羊高繁殖力关系的研究[D].南京:南京农业大学,2010.
    19.张海容,张勇,张海涛,李发弟.甘肃高山细毛母羊血浆中雌二醇和孕酮的含量变化[J].甘肃农业大学学报,2005,40(2):138-141.
    20.国家畜禽遗传资源委员会组编.中国畜禽遗传资源志羊志[M].北京:中国农业出版社,2011.
    21.章亨璆.谈小尾寒羊繁殖性能的开发利用[J].中国养羊,1995,(2):27-29.
    22.朱娜,贾洪响,刘晓坤,赵晓娥,魏强,马保华.小鼠发情周期卵泡发育动态及其对超数排卵的影响[J].动物学研究,2012,33(3):276-282.
    23.赵伟,王公金,徐晓波.波尔山羊发情周期和妊娠期外周血浆中孕酮、雌二醇和睾酮的浓度变化[J].江苏农业科学,2004,(3):60-61.
    24.张小辉. MLT、LH和FSH对绵羊季节性发情的调控作用研究[D].西安:西北农林科技大学,2004.
    25.张英杰,刘月琴,储明星.小尾寒羊高繁殖力和常年发情内分泌机理研究[J].畜牧兽医学报,2001,32:510-516.
    26. Abe M, Herzog E D, Yamazaki S, Straume M, Tei H, Sakaki Y, Menaker M, Block G D. Circadianrhythms in isolated brain regions. J Neurosci,2002,22(1):350-356.
    27. Adachi S, Yamada S, Takatsu Y, Matsui H, Kinoshita M, Takase K, Sugiura H, Ohtaki T,Matsumoto H, Uenoyama Y, Tsukamura H, Inoue K, Maeda K. Involvement of anteroventralperiventricular metastin/kisspeptin neurons in estrogen positive feedback action on luteinizinghormone release in female rats. J Reprod Dev,2007,53(2):367-378.
    28. Anderson G M, Hardy S L, Valent M, Billings H J, Connors J M, Goodman R L. Evidence thatthyroid hormones act in the ventromedial preoptic area and the premammillary region of the brainto allow the termination of the breeding season in the ewe. Endocrinology,2003,144(7):2892-2901.
    29. Arck P, Hansen P J, Mulac Jericevic B, Piccinni M P, Szekeres-Bartho J. Progesterone duringpregnancy: endocrine-immune cross talk in mammalian species and the role of stress. Am J ReprodImmunol,2007,58(3):268-279.
    30. Arendt J. Melatonin and the pineal gland: influence on mammalian seasonal and circadianphysiology. Rev Reprod,1998,3(1):13-22.
    31. Arendt J, Symons A M, Laud C A, Pryde S J. Melatonin can induce early onset of the breedingseason in ewes. J Endocrinol,1983,97(3):395-400.
    32. Ashburner M, Ball C A, Blake J A, Botstein D, Butler H, Cherry J M, Davis A P, Dolinski K,Dwight S S, Eppig J T, Harris M A, Hill D P, Issel-Tarver L, Kasarskis A, Lewis S, Matese J C,Richardson J E, Ringwald M, Rubin G M, Sherlock G. Gene ontology: tool for the unification ofbiology. The Gene Ontology Consortium. Nat Genet,2000,25(1):25-29.
    33. Baby T E, Bartlewski P M. Circulating concentrations of ovarian steroids and follicle-stimulatinghormone (FSH) in ewes with3or4waves of antral follicle emergence per estrous cycle. ReprodBiol,2011,11(1):19-36.
    34. Balsalobre A. Clock genes in mammalian peripheral tissues. Cell Tissue Res,2002,309(1):193-199.
    35. Barrell G K, Moenter S M, Caraty A, Karsch F J. Seasonal changes of gonadotropin-releasinghormone secretion in the ewe. Biol Reprod,1992,46(6):1130-1135.
    36. Barrett P, Ebling F J, Schuhler S, Wilson D, Ross A W, Warner A, Jethwa P, Boelen A, Visser T J,Ozanne D M, Archer Z A, Mercer J G, Morgan P J. Hypothalamic thyroid hormone catabolism actsas a gatekeeper for the seasonal control of body weight and reproduction. Endocrinology,2007,148(8):3608-3617.
    37. Beard A P, Lamming G E. Oestradiol concentration and the development of the uterine oxytocinreceptor and oxytocin-induced PGF2alpha release in ewes. J Reprod Fertil,1994,100(2):469-475.
    38. Bechtold D A, Loudon A S. Hypothalamic thyroid hormones: mediators of seasonal physiology.Endocrinology,2007,148(8):3605-3607.
    39. Berezikov E, Thuemmler F, van Laake L W, Kondova I, Bontrop R, Cuppen E, Plasterk R H.Diversity of microRNAs in human and chimpanzee brain. Nat Genet,2006,38(12):1375-1377.
    40. Billings H J, Viguie C, Karsch F J, Goodman R L, Connors J M, Anderson G M. Temporalrequirements of thyroid hormones for seasonal changes in LH secretion. Endocrinology,2002,143(7):2618-2625.
    41. Bittman E L, Karsch F J. Nightly duration of pineal melatonin secretion determines thereproductive response to inhibitory day length in the ewe. Biol Reprod,1984,30(3):585-593.
    42. Bittman E L, Karsch F J, Hopkins J W. Role of the pineal gland in ovine photoperiodism:regulation of seasonal breeding and negative feedback effects of estradiol upon luteinizinghormone secretion. Endocrinology,1983,113(1):329-336.
    43. Bittman E L, Kaynard A H, Olster D H, Robinson J E, Yellon S M, Karsch F J. Pineal melatoninmediates photoperiodic control of pulsatile luteinizing hormone secretion in the ewe.Neuroendocrinology,1985,40(5):409-418.
    44. Boguski M S, Tolstoshev C M, Bassett D E, Jr. Gene discovery in dbEST. Science,1994,265(5181):1993-2004.
    45. Bosch M A, Xue C, Ronnekleiv O K. Kisspeptin expression in guinea pig hypothalamus: effects of17beta-estradiol. J Comp Neurol,2012,520(10):2143-2162.
    46. Brenner S, Johnson M, Bridgham J, Golda G, Lloyd D H, Johnson D, Luo S, McCurdy S, Foy M,Ewan M, Roth R, George D, Eletr S, Albrecht G, Vermaas E, Williams S R, Moon K, Burcham T,Pallas M, DuBridge R B, Kirchner J, Fearon K, Mao J, Corcoran K. Gene expression analysis bymassively parallel signature sequencing (MPSS) on microbead arrays. Nat Biotechnol,2000,18(6):630-634.
    47. Burnside J, Bernberg E, Anderson A, Lu C, Meyers B C, Green P J, Jain N, Isaacs G, Morgan R W.Marek's disease virus encodes MicroRNAs that map to meq and the latency-associated transcript. JVirol,2006,80(17):8778-8786.
    48. Chada M, Prusa R, Bronsky J, Kotaska K, Sidlova K, Pechova M, Lisa L. Inhibin B, folliclestimulating hormone, luteinizing hormone and testosterone during childhood and puberty in males:changes in serum concentrations in relation to age and stage of puberty. Physiol Res,2003,52(1):45-51.
    49. Chemineau P, Gauthier D, Poirier J C, et al. Plasma levels of LH, FSH, prolactin, oestradiol-17βand progesterone during nature and induced oestrus in the dairy goat [J]. Theriogenology,1982,17:313-323.
    50. Clark T A, Sugnet C W, Ares M, Jr. Genomewide analysis of mRNA processing in yeast usingsplicing-specific microarrays. Science,2002,296(5569):907-910.
    51. Clarke I J. GnRH and ovarian hormone feedback. Oxf Rev Reprod Biol,1987,9:54-95.
    52. Clarke I J. Variable patterns of gonadotropin-releasing hormone secretion during the estrogen-induced luteinizing hormone surge in ovariectomized ewes. Endocrinology,1993,133(4):1624-
    1632.
    53. Clarkson J, d'Anglemont de Tassigny X, Colledge W H, Caraty A, Herbison A E. Distribution ofkisspeptin neurones in the adult female mouse brain. J Neuroendocrinol,2009,21(8):673-682.
    54. Cloonan N, Forrest A R, Kolle G, Gardiner B B, Faulkner G J, Brown M K, Taylor D F, Steptoe AL, Wani S, Bethel G, Robertson A J, Perkins A C, Bruce S J, Lee C C, Ranade S S, Peckham H E,Manning J M, McKernan K J, Grimmond S M. Stem cell transcriptome profiling via massive-scalemRNA sequencing. Nat Methods,2008,5(7):613-619.
    55. Costa V, Angelini C, De Feis I, Ciccodicola A. Uncovering the complexity of transcriptomes withRNA-Seq. J Biomed Biotechnol,2010,2010:853916.
    56. Cunningham N F, Symons A M, Saba N. Levels of progesterone, LH and FSH in the plasma ofsheep during the oestrous cycle. J Reprod Fertil,1975,45(1):177-180.
    57. Dahl G E, Evans N P, Thrun L A, Karsch F J. Thyroxine is permissive to seasonal transitions inreproductive neuroendocrine activity in the ewe. Biol Reprod,1995,52(3):690-696.
    58. Dardente H. Melatonin-dependent timing of seasonal reproduction by the pars tuberalis: pivotalroles for long daylengths and thyroid hormones. J Neuroendocrinol,2012,24(2):249-266.
    59. de Reviers M M, Ravault J P, Tillet Y, Pelletier J. Melatonin binding sites in the sheep parstuberalis. Neurosci Lett,1989,100(1-3):89-93.
    60. Decourt C, Tillet Y, Caraty A, Franceschini I, Briant C. Kisspeptin immunoreactive neurons in theequine hypothalamus Interactions with GnRH neuronal system. J Chem Neuroanat,2008,36(3-4):131-137.
    61. Denamur R, Martinet J, Short R V. Pituitary control of the ovine corpus luteum. J Reprod Fertil,1973,32(2):207-220.
    62. Dennis G, Jr., Sherman B T, Hosack D A, Yang J, Gao W, Lane H C, Lempicki R A. DAVID:Database for Annotation, Visualization, and Integrated Discovery. Genome Biol,2003,4(5):3-3.
    63. Deveson S L, Arendt J, Forsyth I A, The influence of the pineal gland and melatonin on thereproductive performance of domesticated female ungulates.1992,30:113-134.
    64. Dore M A. Structural aspects of luteal function and regression in the ovary of the domestic dog. JReprod Fertil Suppl,1989,39:41-53.
    65. Dubocovich M L, Rivera-Bermudez M A, Gerdin M J, Masana M I. Molecular pharmacology,regulation and function of mammalian melatonin receptors. Front Biosci,2003,8: d1093-1108.
    66. Ebling F J. Photoperiodic regulation of puberty in seasonal species. Mol Cell Endocrinol,2010,324(1-2):95-101.
    67. Estrada K M, Clay C M, Pompolo S, Smith J T, Clarke I J. Elevated KiSS-1expression in thearcuate nucleus prior to the cyclic preovulatory gonadotrophin-releasing hormone/lutenisinghormone surge in the ewe suggests a stimulatory role for kisspeptin in oestrogen-positive feedback.J Neuroendocrinol,2006,18(10):806-809.
    68. Follett B K, Sharp P J. Circadian rhythmicity in photoperiodically induced gonadotrophin releaseand gonadal growth in the quail. Nature,1969,223(5209):968-971.
    69. Franceschini I, Lomet D, Cateau M, Delsol G, Tillet Y, Caraty A. Kisspeptin immunoreactive cellsof the ovine preoptic area and arcuate nucleus co-express estrogen receptor alpha. Neurosci Lett,2006,401(3):225-230.
    70. Gerhard D S, Wagner L, Feingold E A, Shenmen C M, Grouse L H, Schuler G, Klein S L, Old S,Rasooly R, Good P, Guyer M, Peck A M, Derge J G, Lipman D, Collins F S, Jang W, Sherry S,Feolo M, Misquitta L, Lee E, Rotmistrovsky K, Greenhut S F, Schaefer C F, Buetow K, Bonner T I,Haussler D, Kent J, Kiekhaus M, Furey T, Brent M, Prange C, Schreiber K, Shapiro N, Bhat N K,Hopkins R F, Hsie F, Driscoll T, Soares M B, Casavant T L, Scheetz T E, Brown-stein M J, UsdinT B, Toshiyuki S, Carninci P, Piao Y, Dudekula D B, Ko M S, Kawakami K, Suzuki Y, Sugano S,Gruber C E, Smith M R, Simmons B, Moore T, Waterman R, Johnson S L, Ruan Y, Wei C L,Mathavan S, Gunaratne P H, Wu J, Garcia A M, Hulyk S W, Fuh E, Yuan Y, Sneed A, Kowis C,Hodgson A, Muzny D M, McPherson J, Gibbs R A, Fahey J, Helton E, Ketteman M, Madan A,Rodrigues S, Sanchez A, Whiting M, Madari A, Young A C, Wetherby K D, Granite S J, Kwong PN, Brinkley C P, Pearson R L, Bouffard G G, Blakesly R W, Green E D, Dickson M C, RodriguezA C, Grimwood J, Schmutz J, Myers R M, Butterfield Y S, Griffith M, Griffith O L, Krzywinski MI, Liao N, Morin R, Palmquist D, Petrescu A S, Skalska U, Smailus D E, Stott J M, Schnerch A,Schein J E, Jones S J, Holt R A, Baross A, Marra M A, Clifton S, Makowski K A, Bosak S, MalekJ. The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian GeneCollection (MGC). Genome Res,2004,14(10B):2121-2127.
    71. Ginther O J. Influence of exogenous progesterone and the uterus on ovarian activity in sheep.Endocrinology,1968,83(3):613-615.
    72. Goding J R, Catt K J, Brown J M, Kaltenbach C C, Cumming I A, Mole B J. Radioimmunoassayfor ovine luteinizing hormone. Secretion of luteinizing hormone during estrus and followingestrogen administration in sheep. Endocrinology,1969,85(1):133-142.
    73. Goldman B D. Mammalian photoperiodic system: formal properties and neuroendocrinemechanisms of photoperiodic time measurement. J Biol Rhythms,2001,16(4):283-301.
    74. Goodman R L, Bittman E L, Foster D L, Karsch F J. Alterations in the control of luteinizinghormone pulse frequency underlie the seasonal variation in estradiol negative feedback in the ewe.Biol Reprod,1982,27(3):580-589.
    75. Goodman R L, Lehman M N, Smith J T, Coolen L M, de Oliveira C V, Jafarzadehshirazi M R,Pereira A, Iqbal J, Caraty A, Ciofi P, Clarke I J. Kisspeptin neurons in the arcuate nucleus of theewe express both dynorphin A and neurokinin B. Endocrinology,2007,148(12):5752-7560.
    76. Govindan S, Ahmad S N, Vedicherla B, Kodati V, Jahan P, Rao K P, Ahuja Y R, Hasan Q.Association of progesterone receptor gene polymorphism (PROGINS) with endometriosis, uterinefibroids and breast cancer. Cancer Biomark,2007,3(2):73-78.
    77. Greives T J, Humber S A, Goldstein A N, Scotti M A, Demas G E, Kriegsfeld L J. Photoperiod andtestosterone interact to drive seasonal changes in kisspeptin expression in Siberian hamsters(Phodopus sungorus). J Neuroendocrinol,2008,20(12):1339-1347.
    78. Greives T J, Mason A O, Scotti M A, Levine J, Ketterson E D, Kriegsfeld L J, Demas G E.Environmental control of kisspeptin: implications for seasonal reproduction. Endocrinology,2007,148(3):1158-1166.
    79. Hanon E A, Lincoln G A, Fustin J M, Dardente H, Masson-Pevet M, Morgan P J, Hazlerigg D G.Ancestral TSH mechanism signals summer in a photoperiodic mammal. Curr Biol,2008,18(15):1147-1152.
    80. Harbers M, Carninci P. Tag-based approaches for transcriptome research and genome annotation.Nat Methods,2005,2(7):495-502.
    81. Harris T D, Buzby P R, Babcock H, Beer E, Bowers J, Braslavsky I, Causey M, Colonell J, DimeoJ, Efcavitch J W, Giladi E, Gill J, Healy J, Jarosz M, Lapen D, Moulton K, Quake S R, SteinmannK, Thayer E, Tyurina A, Ward R, Weiss H, Xie Z. Single-molecule DNA sequencing of a viralgenome. Science,2008,320(5872):106-109.
    82. Homer N, Merriman B, Nelson S F. BFAST: an alignment tool for large scale genomeresequencing. PLoS One,2009,4(11): e7767.
    83. Hrabovszky E, Ciofi P, Vida B, Horvath M C, Keller E, Caraty A, Bloom S R, Ghatei M A, DhilloW S, Liposits Z, Kallo I. The kisspeptin system of the human hypothalamus: sexual dimorphismand relationship with gonadotropin-releasing hormone and neurokinin B neurons. Eur J Neurosci,2010,31(11):1984-1998.
    84. Hu Z, Hung J H, Wang Y, Chang Y C, Huang C L, Huyck M, DeLisi C. VisANT3.5: multi-scalenetwork visualization, analysis and inference based on the gene ontology. Nucleic Acids Res,2009,37(Web Server issue): W115-121.
    85. Hutchinson J S, Robertson H A. The growth of the follicle and corpus luteum in the ovary of thesheep. Res Vet Sci,1966,7(1):17-24.
    86. I'Anson H, Legan S J. Changes in LH pulse frequency and serum progesterone concentrationsduring the transition to breeding season in ewes. J Reprod Fertil,1988,82(1):341-351.
    87. Johnston J D. Measuring seasonal time within the circadian system: regulation of thesuprachiasmatic nuclei by photoperiod. J Neuroendocrinol,2005,17(7):459-465.
    88. Johnston J D, Ebling F J, Hazlerigg D G. Photoperiod regulates multiple gene expression in thesuprachiasmatic nuclei and pars tuberalis of the Siberian hamster (Phodopus sungorus). Eur JNeurosci,2005,21(11):2967-2974.
    89. Karsch F J, Bittman E L, Foster D L, Goodman R L, Legan S J, Robinson J E. Neuroendocrinebasis of seasonal reproduction. Recent Prog Horm Res,1984,40:185-232.
    90. Karsch F J, Dahl G E, Evans N P, Manning J M, Mayfield K P, Moenter S M, Foster D L. Seasonalchanges in gonadotropin-releasing hormone secretion in the ewe: alteration in response to thenegative feedback action of estradiol. Biol Reprod,1993,49(6):1377-1383.
    91. Karsch F J, Goodman R L, Legan S J. Feedback basis of seasonal breeding: test of an hypothesis. JReprod Fertil,1980,58(2):521-535.
    92. Kinoshita M, Tsukamura H, Adachi S, Matsui H, Uenoyama Y, Iwata K, Yamada S, Inoue K,Ohtaki T, Matsumoto H, Maeda K. Involvement of central metastin in the regulation ofpreovulatory luteinizing hormone surge and estrous cyclicity in female rats. Endocrinology,2005,146(10):4431-4436.
    93. Kodzius R, Kojima M, Nishiyori H, Nakamura M, Fukuda S, Tagami M, Sasaki D, Imamura K,Kai C, Harbers M, Hayashizaki Y, Carninci P. CAGE: cap analysis of gene expression. NatMethods,2006,3(3):211-222.
    94. Kuenzel W J. The search for deep encephalic photoreceptors within the avian brain, using gonadaldevelopment as a primary indicator. Poult Sci,1993,72(5):959-967.
    95. Land R B, Pelletier J, Thimonier J, Mauleon P. A quantitative study of genetic differences in theincidence of oestrus, ovulation and plasma luteinizing hormone concentration in the sheep. JEndocrinol,1973,58(2):305-317.
    96. Lee J H, Miele M E, Hicks D J, Phillips K K, Trent J M, Weissman B E, Welch D R. KiSS-1, anovel human malignant melanoma metastasis-suppressor gene. J Natl Cancer Inst,1996,88(23):1731-1737.
    97. Legan S J, Karsch F J. Neuroendocrine regulation of the estrous cycle and seasonal breeding in theewe. Biol Reprod,1979,20(1):74-85.
    98. Legan S J, Karsch F J. Photoperiodic control of seasonal breeding in ewes: modulation of thenegative feedback action of estradiol. Biol Reprod,1980,23(5):1061-1068.
    99. Legan S J, Karsch F J, Foster D L. The endocrin control of seasonal reproductive function in theewe: a marked change in response to the negative feedback action of estradiol on luteinizinghormone secretion. Endocrinology,1977,101(3):818-824.
    100. Lincoln G A. Secretion of LH in rams exposed to two different photoperiods. J Reprod Fertil,1976,47(2):351-353.
    101. Lincoln G A. Effects of placing micro-implants of melatonin in the pars tuberalis, pars distalis andthe lateral septum of the forebrain on the secretion of FSH and prolactin, and testicular size in rams.J Endocrinol,1994,142(2):267-276.
    102. Lincoln G A, Lincoln C E, McNeilly A S. Seasonal cycles in the blood plasma concentration ofFSH, inhibin and testosterone, and testicular size in rams of wild, feral and domesticated breeds ofsheep. J Reprod Fertil,1990,88(2):623-633.
    103. Lincoln G A, Maeda K I. Reproductive effects of placing micro-implants of melatonin in themediobasal hypothalamus and preoptic area in rams. J Endocrinol,1992,132(2):201-215.
    104. Lincoln G A, Peet M J. Photoperiodic control of gonadotrophin secretion in the ram: a detailedstudy of the temporal changes in plasma levels of follicle-stimulating hormone, luteinizinghormone and testosterone following an abrupt switch from long to short days. J Endocrinol,1977,74(3):355-367.
    105. Lockhart D J, Winzeler E A. Genomics, gene expression and DNA arrays. Nature,2000,405(6788):827-836.
    106. Maeda K, Ohkura S, Uenoyama Y, Wakabayashi Y, Oka Y, Tsukamura H, Okamura H.Neurobiological mechanisms underlying GnRH pulse generation by the hypothalamus. Brain Res,2010,1364:103-115.
    107. Malpaux B, Daveau A, Maurice-Mandon F, Duarte G, Chemineau P. Evidence that melatonin actsin the premammillary hypothalamic area to control reproduction in the ewe: presence of bindingsites and stimulation of luteinizing hormone secretion by in situ microimplant delivery.Endocrinology,1998,139(4):1508-1516.
    108. Malpaux B, Daveau A, Maurice F, Gayrard V, Thiery J C. Short-day effects of melatonin onluteinizing hormone secretion in the ewe: evidence for central sites of action in the mediobasalhypothalamus. Biol Reprod,1993,48(4):752-760.
    109. Malpaux B, Daveau A, Maurice F, Locatelli A, Thiery J C. Evidence that melatonin binding sites inthe pars tuberalis do not mediate the photoperiodic actions of melatonin on LH and prolactinsecretion in ewes. J Reprod Fertil,1994,101(3):625-632.
    110. Malpaux B, Skinner D C, Maurice F. The ovine pars tuberalis does not appear to be targeted bymelatonin to modulate luteinizing hormone secretion, but may be important for prolactin release. JNeuroendocrinol,1995,7(3):199-206.
    111. Malpaux B, Tricoire H, Mailliet F, Daveau A, Migaud M, Skinner D C, Pelletier J, Chemineau P.Melatonin and seasonal reproduction: understanding the neuroendocrine mechanisms using thesheep as a model. Reprod Suppl,2002,59:167-179.
    112. Malpaux B, ViguiéC, Skinner D C, Thiéry J C, Pelletier J, Chemineau P, Seasonal breeding insheep: Mechanism of action of melatonin. Vol.42.1996.109-117.
    113. Mardis E R. The impact of next-generation sequencing technology on genetics. Trends Genet,2008,24(3):133-141.
    114. Margulies M, Egholm M, Altman W E, Attiya S, Bader J S, Bemben L A, Berka J, Braverman M S,Chen Y J, Chen Z, Dewell S B, Du L, Fierro J M, Gomes X V, Godwin B C, He W, Helgesen S, HoC H, Irzyk G P, Jando S C, Alenquer M L, Jarvie T P, Jirage K B, Kim J B, Knight J R, Lanza J R,Leamon J H, Lefkowitz S M, Lei M, Li J, Lohman K L, Lu H, Makhijani V B, McDade K E,McKenna M P, Myers E W, Nickerson E, Nobile J R, Plant R, Puc B P, Ronan M T, Roth G T,Sarkis G J, Simons J F, Simpson J W, Srinivasan M, Tartaro K R, Tomasz A, Vogt K A, Volkmer GA, Wang S H, Wang Y, Weiner M P, Yu P, Begley R F, Rothberg J M. Genome sequencing inmicrofabricated high-density picolitre reactors. Nature,2005,437(7057):376-380.
    115. Marioni J C, Mason C E, Mane S M, Stephens M, Gilad Y. RNA-seq: an assessment of technicalreproducibility and comparison with gene expression arrays. Genome Res,2008,18(9):1509-1517.
    116. Martin G B. Factors affecting the secretion of luteinizing hormone in the ewe. Biol Rev CambPhilos Soc,1984,59(1):1-87.
    117. Matos M H, Lima-Verde I B, Luque M C, Maia J E, Jr., Silva J R, Celestino J J, Martins F S, Bao SN, Lucci C M, Figueiredo J R. Essential role of follicle stimulating hormone in the maintenance ofcaprine preantral follicle viability in vitro. Zygote,2007,15(2):173-182.
    118. McCabe M, Waters S, Morris D, Kenny D, Lynn D, Creevey C. RNA-seq analysis of differentialgene expression in liver from lactating dairy cows divergent in negative energy balance. BMCGenomics,2012,13:193.
    119. Messager S. Kisspeptin and its receptor: new gatekeepers of puberty. J Neuroendocrinol,2005,17(10):687-688.
    120. Messager S, Chatzidaki E E, Ma D, Hendrick A G, Zahn D, Dixon J, Thresher R R, Malinge I,Lomet D, Carlton M B, Colledge W H, Caraty A, Aparicio S A. Kisspeptin directly stimulatesgonadotropin-releasing hormone release via G protein-coupled receptor54. Proc Natl Acad Sci U SA,2005,102(5):1761-1766.
    121. Morin R, Bainbridge M, Fejes A, Hirst M, Krzywinski M, Pugh T, McDonald H, Varhol R, Jones S,Marra M. Profiling the HeLa S3transcriptome using randomly primed cDNA and massivelyparallel short-read sequencing. Biotechniques,2008,45(1):81-94.
    122. Morin R D, O'Connor M D, Griffith M, Kuchenbauer F, Delaney A, Prabhu A L, Zhao Y,McDonald H, Zeng T, Hirst M, Eaves C J, Marra M A. Application of massively parallelsequencing to microRNA profiling and discovery in human embryonic stem cells. Genome Res,2008,18(4):610-621.
    123. Mortazavi A, Williams B A, McCue K, Schaeffer L, Wold B. Mapping and quantifying mammaliantranscriptomes by RNA-Seq. Nat Methods,2008,5(7):621-628.
    124. Nagalakshmi U, Wang Z, Waern K, Shou C, Raha D, Gerstein M, Snyder M. The transcriptionallandscape of the yeast genome defined by RNA sequencing. Science,2008,320(5881):1344-1349.
    125. Nagoshi E, Saini C, Bauer C, Laroche T, Naef F, Schibler U. Circadian gene expression inindividual fibroblasts: cell-autonomous and self-sustained oscillators pass time to daughter cells.Cell,2004,119(5):693-705.
    126. Nakamura M, Carninci P.[Cap analysis gene expression: CAGE]. Tanpakushitsu Kakusan Koso,2004,49(17Suppl):2688-2693.
    127. Navarro V M, Castellano J M, Garcia-Galiano D, Tena-Sempere M. Neuroendocrine factors in theinitiation of puberty: the emergent role of kisspeptin. Rev Endocr Metab Disord,2007,8(1):11-20.
    128. Nicholls T J, Follett B K, Goldsmith A R, Pearson H. Possible homologies betweenphotorefractoriness in sheep and birds: the effect of thyroidectomy on the length of the ewe'sbreeding season. Reprod Nutr Dev,1988,28(2B):375-385.
    129. Nicholls T J, Goldsmith A R, Dawson A. Photorefractoriness in birds and comparison withmammals. Physiol Rev,1988,68(1):133-176.
    130. Nix D A, Courdy S J, Boucher K M. Empirical methods for controlling false positives andestimating confidence in ChIP-Seq peaks. BMC Bioinformatics,2008,9:523.
    131. O'Callaghan D, Wendling A, Karsch F J, Roche J F. Effect of exogenous thyroxine on timing ofseasonal reproductive transitions in ewes. Biol Reprod,1993,49(2):311-315.
    132. Oakley A E, Clifton D K, Steiner R A. Kisspeptin signaling in the brain. Endocr Rev,2009,30(6):713-743.
    133. Ohtaki T, Shintani Y, Honda S, Matsumoto H, Hori A, Kanehashi K, Terao Y, Kumano S, Takatsu Y,Masuda Y, Ishibashi Y, Watanabe T, Asada M, Yamada T, Suenaga M, Kitada C, Usuki S,Kurokawa T, Onda H, Nishimura O, Fujino M. Metastasis suppressor gene KiSS-1encodes peptideligand of a G-protein-coupled receptor. Nature,2001,411(6837):613-617.
    134. Okoniewski M J, Miller C J. Hybridization interactions between probesets in short oligomicroarrays lead to spurious correlations. BMC Bioinformatics,2006,7:276.
    135. Oliver J, Bayle J D. Brain photoreceptors for the photo-induced testicular response in birds.Experientia,1982,38(9):1021-1029.
    136. Ortavant R, Bocquier F, Pelletier J, Ravault J P, Thimonier J, Volland-Nail P. Seasonality ofreproduction in sheep and its control by photoperiod. Aust J Biol Sci,1988,41(1):69-85.
    137. Ortavant R, Pelletier J, Ravault J P, Thimonier J, Volland-Nail P. Photoperiod: main proximal anddistal factor of the circannual cycle of reproduction in farm mammals. Oxf Rev Reprod Biol,1985,7:305-345.
    138. Parkinson T J, Follett B K. Effect of thyroidectomy upon seasonality in rams. J Reprod Fertil,1994,101(1):51-58.
    139. Parkinson T J, Follett B K. Thyroidectomy abolishes seasonal testicular cycles of Soay rams. ProcBiol Sci,1995,259(1354):1-6.
    140. Peiffer J A, Kaushik S, Sakai H, Arteaga-Vazquez M, Sanchez-Leon N, Ghazal H, Vielle-Calzada JP, Meyers B C. A spatial dissection of the Arabidopsis floral transcriptome by MPSS. BMC PlantBiol,2008,8:43.
    141. Pelletier J, Almeida G. Short light cycles induce persistent reproductive activity in Ile-de-Francerams. J Reprod Fertil Suppl,1987,34:215-226.
    142. Pelletier J, Ortavant R. Photoperiodic control of LH release in the ram. I. Influence of increasingand decreasing light photoperiods. Acta Endocrinol (Copenh),1975,78(3):435-441.
    143. Pinilla L, Aguilar E, Dieguez C, Millar R P, Tena-Sempere M. Kisspeptins and reproduction:physiological roles and regulatory mechanisms. Physiol Rev,2012,92(3):1235-1316.
    144. Platt T E, Foster G S, Tarnavsky G K, Reeves J J. Effects of photoperiod and estradiol on tonicgonadotropins in ovariectomized ewes. J Anim Sci,1983,56(5):1180-1185.
    145. Prendergast B J, Pyter L M, Kampf-Lassin A, Patel P N, Stevenson T J. Rapid induction ofhypothalamic iodothyronine deiodinase expression by photoperiod and melatonin in juvenileSiberian hamsters (Phodopus sungorus). Endocrinology,2013,154(2):831-841.
    146. Reinartz J, Bruyns E, Lin J Z, Burcham T, Brenner S, Bowen B, Kramer M, Woychik R. Massivelyparallel signature sequencing (MPSS) as a tool for in-depth quantitative gene expression profilingin all organisms. Brief Funct Genomic Proteomic,2002,1(1):95-104.
    147. Reiter R J. The pineal and its hormones in the control of reproduction in mammals. Endocr Rev,1980,1(2):109-131.
    148. Revel F G, Ansel L, Klosen P, Saboureau M, Pevet P, Mikkelsen J D, Simonneaux V. Kisspeptin: akey link to seasonal breeding. Rev Endocr Metab Disord,2007,8(1):57-65.
    149. Revel F G, Masson-Pevet M, Pevet P, Mikkelsen J D, Simonneaux V. Melatonin controls seasonalbreeding by a network of hypothalamic targets. Neuroendocrinology,2009,90(1):1-14.
    150. Revel F G, Saboureau M, Masson-Pevet M, Pevet P, Mikkelsen J D, Simonneaux V. Kisspeptinmediates the photoperiodic control of reproduction in hamsters. Curr Biol,2006,16(17):1730-1735.
    151. Roa J, Aguilar E, Dieguez C, Pinilla L, Tena-Sempere M. New frontiers in kisspeptin/GPR54physiology as fundamental gatekeepers of reproductive function. Front Neuroendocrinol,2008,29(1):48-69.
    152. Robinson J E, Radford H M, Karsch F J. Seasonal changes in pulsatile luteinizing hormone (LH)secretion in the ewe: relationship of frequency of LH pulses to day length and response to estradiolnegative feedback. Biol Reprod,1985,33(2):324-334.
    153. Roche J F, Foster D L, Karsch F J, Cook B, Dziuk P J. Levels of luteinizing hormone in sera andpituitaries of ewes during the estrous cycle and anestrus. Endocrinology,1970,86(3):568-572.
    154. Ronaghi M, Uhlen M, Nyren P. A sequencing method based on real-time pyrophosphate. Science,1998,281(5375):363-365.
    155. Royce T E, Rozowsky J S, Gerstein M B. Toward a universal microarray: prediction of geneexpression through nearest-neighbor probe sequence identification. Nucleic Acids Res,2007,35(15): e99.
    156. Saldanha C J, Leak R K, Silver R. Detection and transduction of daylength in birds.Psychoneuroendocrinology,1994,19(5-7):641-656.
    157. Scaramuzzi R J, Baird D T. Pulsatile release of luteinizing hormone and the secretion of ovariansteroids in sheep during anestrus. Endocrinology,1977,101(6):1801-1806.
    158. Scherbarth F, Steinlechner S. Endocrine mechanisms of seasonal adaptation in small mammals:from early results to present understanding. J Comp Physiol B,2010,180(7):935-952.
    159. Sheikheldin M A, Howland B E, Palmer W M. Seasonal profiles of melatonin in adult rams. JPineal Res,1992,12(2):58-63.
    160. Shiraki T, Kondo S, Katayama S, Waki K, Kasukawa T, Kawaji H, Kodzius R, Watahiki A,Nakamura M, Arakawa T, Fukuda S, Sasaki D, Podhajska A, Harbers M, Kawai J, Carninci P,Hayashizaki Y. Cap analysis gene expression for high-throughput analysis of transcriptionalstarting point and identification of promoter usage. Proc Natl Acad Sci U S A,2003,100(26):15776-15781.
    161. Silver R, Witkovsky P, Horvath P, Alones V, Barnstable C J, Lehman M N. Coexpression of opsin-and VIP-like-immunoreactivity in CSF-contacting neurons of the avian brain. Cell Tissue Res,1988,253(1):189-198.
    162. Simonneaux V, Ribelayga C. Generation of the melatonin endocrine message in mammals: areview of the complex regulation of melatonin synthesis by norepinephrine, peptides, and otherpineal transmitters. Pharmacol Rev,2003,55(2):325-395.
    163. Smith D R, Quinlan A R, Peckham H E, Makowsky K, Tao W, Woolf B, Shen L, Donahue W F,Tusneem N, Stromberg M P, Stewart D A, Zhang L, Ranade S S, Warner J B, Lee C C, Coleman BE, Zhang Z, McLaughlin S F, Malek J A, Sorenson J M, Blanchard A P, Chapman J, Hillman D,Chen F, Rokhsar D S, McKernan K J, Jeffries T W, Marth G T, Richardson P M. Rapid whole-genome mutational profiling using next-generation sequencing technologies. Genome Res,2008,18(10):1638-1642.
    164. Smith J T. Sex steroid control of hypothalamic Kiss1expression in sheep and rodents: comparativeaspects. Peptides,2009,30(1):94-102.
    165. Smith J T, Clarke I J. Kisspeptin expression in the brain: catalyst for the initiation of puberty. RevEndocr Metab Disord,2007,8(1):1-9.
    166. Smith J T, Clarke I J. Seasonal breeding as a neuroendocrine model for puberty in sheep. Mol CellEndocrinol,2010,324(1-2):102-109.
    167. Smith J T, Clay C M, Caraty A, Clarke I J. KiSS-1messenger ribonucleic acid expression in thehypothalamus of the ewe is regulated by sex steroids and season. Endocrinology,2007,148(3):1150-1157.
    168. Smith J T, Coolen L M, Kriegsfeld L J, Sari I P, Jaafarzadehshirazi M R, Maltby M, Bateman K,Goodman R L, Tilbrook A J, Ubuka T, Bentley G E, Clarke I J, Lehman M N. Variation inkisspeptin and RFamide-related peptide (RFRP) expression and terminal connections togonadotropin-releasing hormone neurons in the brain: a novel medium for seasonal breeding in thesheep. Endocrinology,2008,149(11):5770-5782.
    169. Smith J T, Cunningham M J, Rissman E F, Clifton D K, Steiner R A. Regulation of Kiss1geneexpression in the brain of the female mouse. Endocrinology,2005,146(9):3686-3692.
    170. Smith J T, Dungan H M, Stoll E A, Gottsch M L, Braun R E, Eacker S M, Clifton D K, Steiner R A.Differential regulation of KiSS-1mRNA expression by sex steroids in the brain of the male mouse.Endocrinology,2005,146(7):2976-2984.
    171. Souza C J, Campbell B K, Baird D T. Follicular dynamics and ovarian steroid secretion in sheepduring the follicular and early luteal phases of the estrous cycle. Biol Reprod,1997,56(2):483-488.
    172. Sogorescu E, Zamfirescu S, Anghel A H, et al. Seasonal variations of progesterone level andcharacteristics of breeding season and anoestrus period on Carpathian goats [J]. Journal of Animaland Veterinary Advances,2012,11(9):1472-1477.
    173. Suttie J M, Gluckman P D, Butler J H, Fennessy P F, Corson I D, Laas F J. Insulin-like growthfactor1(IGF-1) antler-stimulating hormone? Endocrinology,1985,116(2):846-848.
    174. Tena-Sempere M. GPR54and kisspeptin in reproduction. Hum Reprod Update,2006,12(5):631-639.
    175. Thiery J C, Martin G B. Neurophysiological control of the secretion of gonadotrophin-releasinghormone and luteinizing hormone in the sheep--a review. Reprod Fertil Dev,1991,3(2):137-173.
    176. Thimonier J. Control of seasonal reproduction in sheep and goats by light and hormones. J ReprodFertil Suppl,1981,30:33-45.
    177. Thorburn G D, Bassett J M, Smith I D. Progesterone concentration in the peripheral plasma ofsheep during the oestrous cycle. J Endocrinol,1969,45(3):459-469.
    178. Tournier B B, Menet J S, Dardente H, Poirel V J, Malan A, Masson-Pevet M, Pevet P, Vuillez P.Photoperiod differentially regulates clock genes' expression in the suprachiasmatic nucleus ofSyrian hamster. Neuroscience,2003,118(2):317-322.
    179. Trapnell C, Pachter L, Salzberg S L. TopHat: discovering splice junctions with RNA-Seq.Bioinformatics,2009,25(9):1105-1111.
    180. Trapnell C, Roberts A, Goff L, Pertea G, Kim D, Kelley D R, Pimentel H, Salzberg S L, Rinn J L,Pachter L. Differential gene and transcript expression analysis of RNA-seq experiments withTopHat and Cufflinks. Nat Protoc,2012,7(3):562-578.
    181. Trapnell C, Salzberg S L. How to map billions of short reads onto genomes. Nat Biotechnol,2009,27(5):455-457.
    182. Trapnell C, Williams B A, Pertea G, Mortazavi A, Kwan G, van Baren M J, Salzberg S L, Wold B J,Pachter L. Transcript assembly and quantification by RNA-Seq reveals unannotated transcripts andisoform switching during cell differentiation. Nat Biotechnol,2010,28(5):511-515.
    183. Tu H M, Kim S W, Salvatore D, Bartha T, Legradi G, Larsen P R, Lechan R M. Regionaldistribution of type2thyroxine deiodinase messenger ribonucleic acid in rat hypothalamus andpituitary and its regulation by thyroid hormone. Endocrinology,1997,138(8):3359-3368.
    184. Vallet J L, Lamming G E, Batten M. Control of endometrial oxytocin receptor and uterine responseto oxytocin by progesterone and oestradiol in the ewe. J Reprod Fertil,1990,90(2):625-634.
    185. Velculescu V E, Zhang L, Vogelstein B, Kinzler K W. Serial analysis of gene expression. Science,1995,270(5235):484-487.
    186. Vera J C, Wheat C W, Fescemyer H W, Frilander M J, Crawford D L, Hanski I, Marden J H. Rapidtranscriptome characterization for a nonmodel organism using454pyrosequencing. Mol Ecol,2008,17(7):1636-1647.
    187. Viguie C, Caraty A, Locatelli A, Malpaux B. Regulation of luteinizing hormone-releasing hormone(LHRH) secretion by melatonin in the ewe. I. Simultaneous delayed increase in LHRH andluteinizing hormone pulsatile secretion. Biol Reprod,1995,52(5):1114-1120.
    188. von Gall C, Stehle J H, Weaver D R. Mammalian melatonin receptors: molecular biology andsignal transduction. Cell Tissue Res,2002,309(1):151-162.
    189. von Gall C, Weaver D R, Moek J, Jilg A, Stehle J H, Korf H W. Melatonin plays a crucial role inthe regulation of rhythmic clock gene expression in the mouse pars tuberalis. Ann N Y Acad Sci,2005,1040:508-511.
    190. Walton J S, McNeilly J R, McNeilly A S, Cunningham F J. Changes in concentrations of follicle-stimulating hormone, luteinizing hormone, prolactin and progesterone in the plasma of ewes duringthe transition from anoestrus to breeding activity. J Endocrinol,1977,75(1):127-136.
    191. Wang L, Feng Z, Wang X, Zhang X. DEGseq: an R package for identifying differentially expressedgenes from RNA-seq data. Bioinformatics,2010,26(1):136-138.
    192. Wang L, Wang S, Li W. RSeQC: quality control of RNA-seq experiments. Bioinformatics,2012,28(16):2184-2185.
    193. Wang X W, Luan J B, Li J M, Bao Y Y, Zhang C X, Liu S S. De novo characterization of a whiteflytranscriptome and analysis of its gene expression during development. BMC Genomics,2010,11:400.
    194. Wang Z, Gerstein M, Snyder M. RNA-Seq: a revolutionary tool for transcriptomics. Nat Rev Genet,2009,10(1):57-63.
    195. Watanabe T, Yamamura T, Watanabe M, Yasuo S, Nakao N, Dawson A, Ebihara S, Yoshimura T.Hypothalamic expression of thyroid hormone-activating and-inactivating enzyme genes in relationto photorefractoriness in birds and mammals. Am J Physiol Regul Integr Comp Physiol,2007,292(1): R568-572.
    196. Wayne N L, Malpaux B, Karsch F J. Social cues can play a role in timing onset of the breedingseason of the ewe. J Reprod Fertil,1989,87(2):707-713.
    197. Webster G M, Haresign W. Seasonal changes in LH and prolactin concentrations in ewes of twobreeds. J Reprod Fertil,1983,67(2):465-471.
    198. Webster J R, Moenter S M, Barrell G K, Lehman M N, Karsch F J. Role of the thyroid gland inseasonal reproduction. III. Thyroidectomy blocks seasonal suppression of gonadotropin-releasinghormone secretion in sheep. Endocrinology,1991,129(3):1635-1643.
    199. Webster J R, Moenter S M, Woodfill C J, Karsch F J. Role of the thyroid gland in seasonalreproduction. II. Thyroxine allows a season-specific suppression of gonadotropin secretion insheep. Endocrinology,1991,129(1):176-183.
    200. Wilhelm B T, Marguerat S, Watt S, Schubert F, Wood V, Goodhead I, Penkett C J, Rogers J, BahlerJ. Dynamic repertoire of a eukaryotic transcriptome surveyed at single-nucleotide resolution.Nature,2008,453(7199):1239-1243.
    201. Woodfill C J, Robinson J E, Malpaux B, Karsch F J. Synchronization of the circannualreproductive rhythm of the ewe by discrete photoperiodic signals. Biol Reprod,1991,45(1):110-121.
    202. Xia Y, Sidis Y, Schneyer A. Overexpression of follistatin-like3in gonads causes defects in gonadaldevelopment and function in transgenic mice. Mol Endocrinol,2004,18(4):979-994.
    203. Yao Y, Guo G, Ni Z, Sunkar R, Du J, Zhu J K, Sun Q. Cloning and characterization of microRNAsfrom wheat (Triticum aestivum L.). Genome Biol,2007,8(6): R96.
    204. Yasuo S, Nakao N, Ohkura S, Iigo M, Hagiwara S, Goto A, Ando H, Yamamura T, Watanabe M,Watanabe T, Oda S, Maeda K, Lincoln G A, Okamura H, Ebihara S, Yoshimura T. Long-daysuppressed expression of type2deiodinase gene in the mediobasal hypothalamus of the Saanengoat, a short-day breeder: implication for seasonal window of thyroid hormone action onreproductive neuroendocrine axis. Endocrinology,2006,147(1):432-440.
    205. Yasuo S, Watanabe M, Okabayashi N, Ebihara S, Yoshimura T. Circadian clock genes andphotoperiodism: Comprehensive analysis of clock gene expression in the mediobasalhypothalamus, the suprachiasmatic nucleus, and the pineal gland of Japanese Quail under variouslight schedules. Endocrinology,2003,144(9):3742-3748.
    206. Yasuo S, Watanabe M, Tsukada A, Takagi T, Iigo M, Shimada K, Ebihara S, Yoshimura T.Photoinducible phase-specific light induction of Cry1gene in the pars tuberalis of Japanese quail.Endocrinology,2004,145(4):1612-1616.
    207. Yellon S M, Bittman E L, Lehman M N, Olster D H, Robinson J E, Karsch F J. Importance ofduration of nocturnal melatonin secretion in determining the reproductive response to inductivephotoperiod in the ewe. Biol Reprod,1985,32(3):523-529.
    208. Yoshimura T. Neuroendocrine mechanism of seasonal reproduction in birds and mammals. AnimSci J,2010,81(4):403-410.
    209. Yoshimura T, Yasuo S, Watanabe M, Iigo M, Yamamura T, Hirunagi K, Ebihara S. Light-inducedhormone conversion of T4to T3regulates photoperiodic response of gonads in birds. Nature,2003,426(6963):178-181.
    210. Yuthasastrakosol P, Palmer W M, Howland B E. Release of LH in anoestrous and cyclic ewes. JReprod Fertil,1977,50(2):319-321.
    211. Zarazaga L A, Malpaux B, Chemineau P. Amplitude of the plasma melatonin nycthemeral rhythmsis not associated with the dates of onset and offset of the seasonal ovulatory activity in the Ile-de-France ewe. Reprod Nutr Dev,2003,43(2):167-177.
    212. Zarazaga L A, Gatica M C, Celi I, et al. Exogenous melatonin after a treatment of artificial longdays is able to increase LH secretion during seasonal anoestrus in Payoya goats [J]. Journal ofApplied Animal Research,2013,41(1):34-38.
    213. Zeeberg B R, Feng W, Wang G, Wang M D, Fojo A T, Sunshine M, Narasimhan S, Kane D W,Reinhold W C, Lababidi S, Bussey K J, Riss J, Barrett J C, Weinstein J N. GoMiner: a resource forbiological interpretation of genomic and proteomic data. Genome Biol,2003,4(4): R28.
    214. Zhang G, Guo G, Hu X, Zhang Y, Li Q, Li R, Zhuang R, Lu Z, He Z, Fang X, Chen L, Tian W, TaoY, Kristiansen K, Zhang X, Li S, Yang H, Wang J. Deep RNA sequencing at single base-pairresolution reveals high complexity of the rice transcriptome. Genome Res,2010,20(5):646-654.
    215. Zhang J, Wu K, Zeng S, Teixeira da Silva J A, Zhao X, Tian C E, Xia H, Duan J. Transcriptomeanalysis of Cymbidium sinense and its application to the identification of genes associated withfloral development. BMC Genomics,2013,14:279.
    216. Zhao T, Li G, Mi S, Li S, Hannon G J, Wang X J, Qi Y. A complex system of small RNAs in theunicellular green alga Chlamydomonas reinhardtii. Genes Dev,2007,21(10):1190-1203.

© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号

地址:北京市海淀区学院路29号 邮编:100083

电话:办公室:(+86 10)66554848;文献借阅、咨询服务、科技查新:66554700