黄体期和卵泡期小尾寒羊KiSS-1与RFRP-3组织表达研究
|
摘要
为分析KiSS-1和RFRP-3基因在小尾寒羊不同繁殖时期(黄体期和卵泡期)下丘脑、垂体、性腺轴各组织中的表达差异,阐明这2个基因在绵羊发情转换中的表达模式,以常年发情的小尾寒羊为研究对象,采用实时荧光定量PCR技术,对比分析了KiSS-1和RFRP-3基因在黄体期和卵泡期的小尾寒羊下丘脑—垂体—性腺轴各组织中的表达差异。结果表明:1)KiSS-1和RFRP-3基因在下丘脑、垂体、卵巢等10种组织中均有表达;2)黄体期和卵泡期下丘脑中KiSS-1基因表达量均显著高于其他组织(P<0.05),卵泡期下丘脑和松果体中KiSS-1基因表达量均显著高于黄体期(P<0.05);3)黄体期和卵泡期下丘脑和卵巢中RFRP-3基因的表达量均显著高于其他组织(P<0.05),且卵巢中RFRP-3基因在黄体期的表达量显著高于卵泡期(P<0.05)。综上,研究发现KiSS-1和RFRP-3基因在绵羊发情调控中均发挥重要作用,KiSS-1促进发情而RFRP-3抑制发情。
The aim of this study was to analyze the expression of KiSS-1 and RFRP-3 genes in the tissues of the hypothalamuspituitary-gonadal axis in Small Taili Han sheep during different reproductive periods(luteal and follicular periods),and to elucidate the expression pattern of these two genes in estrus conversion of sheep.Small Tail Han ewes at one year-round estrous breed were selected in the study,and Real-time PCR method was used to detect the expression of KiSS-1 and RFRP-3 genes in hypothalamus-pituitary-gonadal axis' tissues during luteal and follicular periods.The results showed that KiSS-1 and RFRP-3 genes were all expressed in ten tissues such as hypothalamus,pituitary and ovary.The expression of KiSS-1 in hypothalamus was significantly higher(P<0.05)than that in other tissues at both luteal and follicular periods.The expression of KiSS-1 in hypothalamus and pineal gland was significantly higher(P<0.05)in follicular period than that in luteal period.The expression of RFRP-3 in hypothalamus and ovary was significantly higher(P<0.05)than that in other tissues both luteal and follicular periods.The expression of RFRP-3 in ovary was significantly higher(P<0.05)in luteal period than that in follicular period.In conclusion,the KiSS-1 and RFRP-3 genes played important roles in sheep estrus regulation.KiSS-1 promoted estrus while RFRP-3 inhibited estrus.
The aim of this study was to analyze the expression of KiSS-1 and RFRP-3 genes in the tissues of the hypothalamuspituitary-gonadal axis in Small Taili Han sheep during different reproductive periods(luteal and follicular periods),and to elucidate the expression pattern of these two genes in estrus conversion of sheep.Small Tail Han ewes at one year-round estrous breed were selected in the study,and Real-time PCR method was used to detect the expression of KiSS-1 and RFRP-3 genes in hypothalamus-pituitary-gonadal axis' tissues during luteal and follicular periods.The results showed that KiSS-1 and RFRP-3 genes were all expressed in ten tissues such as hypothalamus,pituitary and ovary.The expression of KiSS-1 in hypothalamus was significantly higher(P<0.05)than that in other tissues at both luteal and follicular periods.The expression of KiSS-1 in hypothalamus and pineal gland was significantly higher(P<0.05)in follicular period than that in luteal period.The expression of RFRP-3 in hypothalamus and ovary was significantly higher(P<0.05)than that in other tissues both luteal and follicular periods.The expression of RFRP-3 in ovary was significantly higher(P<0.05)in luteal period than that in follicular period.In conclusion,the KiSS-1 and RFRP-3 genes played important roles in sheep estrus regulation.KiSS-1 promoted estrus while RFRP-3 inhibited estrus.
引文
[1] Greives T J,Mason A O,Scotti M A L,Levine J,Ketterson E D,Kriegsfeld L J,Demas G E.Environmental control of kisspeptin:Implications for seasonal reproduction[J].Endocrinology,2007,148(3):1158-1166
[2] Lee J H,Miele M E,Hicks D J,Phillips K K,Trent J M,Weissman B E,Welch D R.KiSS-1,a novel human malignant melanoma metastasis-suppressor gene[J].Journal of the National Cancer Institute,1996,88(23):1731-1737
[3] Ohtaki T,Shintani Y,Honda S,Matsumoto H,Hori A,Kanehashi K,Ohtaki T L,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 peptide ligand of a G-protein-coupled receptor[J].Nature,2001,411(6837):613-617
[4] Novaira H J, Ng Y, Wolfe A,Radovick S.Kisspeptin increases GnRH mRNA expression and secretion in GnRH secreting neuronal cell lines[J].Molecular and Cellular Endocrinology,2009,311(1-2):126-134
[5] Papaoiconomou E,Msaouel P,Makri A,Diamanti-Kandarakis E,Koutsilieris M.The role of kisspeptin/GPR54in the reproductive system[J].In Vivo,2011,25(3):343-354
[6]郭若婷,鲁慧文,欧科鹏,王颖,刘乙,顾真真,刘小军.绵羊Otx-2基因的分子克隆、组织表达及其与季节性繁殖关系的研究[J].畜牧兽医学报,2015,46(5):696-703Guo R T,Lu H W,Ou K P,Wang Y,Liu Y,Gu Z Z,Liu X J.Molecular cloning,tissue expression and its relationship with seasonal reproduction of Otx-2gene in sheep[J].Journal of Animal and Veterinary Sciences,2015,46(5):696-703(in Chinese)
[7] Thompson E L,Patterson M,Murphy K G,Smith K L,Dhillo W S,Todd J F,Ghatei M A,Bloom S R.Central and peripheral administration of kisspeptin-10 stimulates the hypothalamic-pituitary-gonadalaxis[J]. Journal of Neuroendocrinology,2004,16(10):850-858
[8] Messager S,Chatzidaki E E,Ma D,Hendrick A G,Zahn D,Dixon J,Thresher R R,Malinge I,Lomet D,Carlton M B L,Colledge W H,Caraty A C,Aparicio S A J R.Kisspeptin directly stimulates gonadotropin-releasing hormone release via G protein-coupled receptor 54[J]. Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1761-1766
[9] Caraty A,Smith J T,Lomet D,Ben Sa6d S,Morrissey A,Cognie J,Doughton G,Baril C B,Clarke I J.Kisspeptin synchronizes preovulatory surges in cyclical ewes and causes ovulation in seasonally acyclic ewes[J].Endocrinology,2007,148(11):5258-5267
[10] Tsutsui K,Saigoh E,Ukena K,Teranishi H,Fujisawa Y,Kikuchi M,Ishii S,Sharp P J.A novel avian hypothalamic peptide inhibiting gonadotropin release[J].Biochemical and Biophysical Research Communications,2000,275(2):661-667.
[11]向伟,赖平,张宝云,王凭青,储明星,樊奇,刘雪旭,潭颖.RFRP-3对哺乳动物生殖功能和能量平衡的影响[J].遗传,2012,34(8):969-976Xiang W,Lai P,Zhang B Y,Wang P Q,Chu M X,Fan Q,Liu C X,Tan Y.Effects of RFRP-3 on reproductive function and energy balance in mammals[J].Genetics,2012,34(8):969-976(in Chinese)
[12] Ducret E,Anderson G M,Herbison A E.RFamide-related peptide-3,a mammalian gonadotropin-inhibitory hormone ortholog,regulates gonadotropin-releasing hormone neuron firing in the mouse[J].Endocrinology,2009,150(6):2799-2804
[13] Wu M,Dumalska I,Morozova E,Van Den Pol A N,Alreja M.Gonadotropin inhibitory hormone inhibits basal forebrain vGluT2-gonadotropin-releasing hormone neurons via a direct postsynaptic mechanism[J].The Journal of Physiology,2009,587(7):1401-1411
[14] Smith J T.The role of kisspeptin and gonadotropin inhibitory hormone in the seasonal regulation of reproduction in sheep[J].Domestic Animal Endocrinology,2012,43(2):75-84
[15] Livak K J,Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the2-ΔΔCT method[J].Methods,2001,25(4):402-408
[16] Schmittgen T D,Livak K J.Analyzing real-time PCR data by the comparative CT method[J].Nature Protocols,2008,3(6):1101-1108
[17]张伟,刘晓娜,邓双义,王世银,石国庆,阿勒泰羊不同繁殖状态下丘脑Kiss1基因表达规律与其季节性繁殖的关联性分析[J].基因组学与应用生物学,2016,35(1):45-52Zhang W,Liu X N,Deng S Y,Wang S Y,Shi G Q.Correlation analysis of Kiss1gene expression and seasonal reproduction in the hypothalamus of Altay sheep[J].Genomics and Applied Biology,2016,35(1):45-52(in Chinese)
[18]黄冬维,储明星,狄冉,刘秋月,胡文萍,王翔宇,潘章源,郭晓飞.山羊繁殖季节性相关基因KiSS-1与RFRP的表达研究[J].畜牧兽医学报,2014,46(6):924-931Huang D W,Chu M X,Di R,Liu Q Y,Hu W P,Wang X Y,Pan Z Y,Guo X F.Study on expression of KiSS-1 and RFRPin seasonal goat breeding[J].Journal of Animal and Veterinary Sciences,2014,46(6):924-931(in Chinese)
[19]朱碧泉,李卓昭,吴德,方正锋,林燕,车炼强.梅山与长大母猪下丘脑—垂体—卵巢轴Kiss1和GPR54基因的表达差异[D].中国畜牧杂志,2011,47(13):21-25Zhu B C,Li ZZ,Wu D,Fang Z F,Lin Y,Che L Q.Differences in expression of Kiss1 and GPR54 genes in Meishan and growing sows′hypothalamus-pituitary-ovary axis[D].Chinese Journal of Animal Science,2011,47(13):21-25(in Chinese)
[20] Tomikawa J,Homma T,Tajima S,Shibata T,Inamoto Y,Takase K,Inoue N,Ohkura S,Uenoyama Y,Maeda K,Tsukamura H. Molecular characterization and estrogen regulation of hypothalamic KiSS1 gene in the pig[J].Biology of Reproduction,2010,82(2):313-319
[21] Xiong J,Karsch F J,Lehman M N.Evidence for seasonal plasticity in the gonadotropin-releasing hormone(GnRH)system of the ewe:Changes in synaptic inputs onto GnRH neurons[J].Endocrinology,1997,138(3):1240-1250
[22] Merkley C M,Coolen L M,Goodman R L,Lehman M N.Evidence for changes in numbers of synaptic inputs onto KNDy and GnRH neurones during the preovulatory LH surge in the ewe[J].Journal of Neuroendocrinology,2015,27(7):624-635
[23] El-Tarabany M S,Zaglool A W,El-Tarabany A A,Awad A.Association analysis of polymorphism in KiSS1gene with reproductive traits in goats[J]. Animal Reproduction Science,2017,180:92-99
[24] Messager S,Chatzidaki E E,Ma D,Hendrick A G,Zahn D,Dixon J,Thresher R R,Malinge I,Lomet D,Carlton M B L,Colledge W H,Caraty A,Colledge W H.Kisspeptin directly stimulates gonadotropin-releasing hormone release via G protein-coupled receptor 54[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1761-1766
[25]王军,孙蕾,吕文发,娄玉杰.KiSS-1/GPR54系统在不同发情周期母羊下丘脑中的表达规律[J].西北农林科技大学学报:自然科学版,2011,39(12):49-52Wang J,Sun L,Lu W F,Qi Y J.Expression patterns of KiSS-1/GPR54system in different hypothalamus of estrous cycle in ewe[J].Journal of Northwest A&F University:Natural Science Edition,2011,39(12):49-52(in Chinese)
[26] Robert L.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,Clarke I J.Kisspeptin neurons in the arcuate nucleus of the ewe express both dynorphin A and neurokinin B[J].Endocrinology,2007,148(12):5752-5760
[27] Wagner G C,Johnston J D,Clarke I J,Lincoln G A,Hazlerigg D G. Redefining the limits of day length responsiveness in a seasonal mammal[J].Endocrinology,2007,149(1):32-39
[28] Luridiana S,Mura M C,Daga C,Cosso G,Bodano S,Farci F,Zidda F,Carcangiu V.Influences of melatonin treatment,melatonin receptor 1A(MTNR1A)and kisspeptin(KiSS-1)gene polymorphisms on first conception in Sarda ewe lambs[J].Reproduction Fertility and Development,2016,28(6):750-756
[29] Smith J T,Clarke I J.Seasonal breeding as a neuroendocrine model for puberty in sheep[J].Molecular and Cellular Endocrinology,2010,324(1-2):102-109
[30] Jafarzadeh Shirazi M R,Zamiri M J,Salehi M S,Moradi S,Tamadon A,Namavar M R,Akhlaghi A,Tsutsui K,Caraty A.Differential expression of RFamide-related peptide,a mammalian gonadotrophin-inhibitory hormone orthologue,and kisspeptin in the hypothalamus of Abadeh ecotype does during breeding and anoestrous seasons[J].Journal of Neuroendocrinology,2014,26(3):186-194
[31] Beskonakli E,Palaoglu S,Aksaray S,Alanoglu G,Turhan T,Taskin Y.Effect of pinealectomy on immune parameters in ratswith Staphylococcus aureus infection[J].Neurosurgical Review,2001,24(1):26-30
[32] Tsutsui K,Bentley G E,Kriegsfeld L J,Osugi T,Seong J Y, Vaudry H. Discovery and evolutionary history of gonadotrophin-inhibitory hormone and kisspeptin:New key neuropeptides controlling reproduction[J]. Journal of Neuroendocrinology,2010,22(7):716-727
[33]王梁,刘国世,史文清,史建民,高超,田秀芝,张璐,赵广华,商业,云鹏,朱世恩.外源褪黑激素对梅花鹿超数排卵及体内促性腺激素的影响[J].中国鹿业进展,2011Wang L,Liu G S,Shi W Q,Shi J M,Gao C,Tian X Z,Zhang L,Zhao G H,Shang Y,Yun P,Zhu S E.Effects of exogenous melatonin on superovulation and gonadotropin in Sika deer[J].China Deer Industry Progress,2011(in Chinese)
[34] Li X,Su J,Lei Z,Zhao Y,Jin M,Fang R,Zheng L C,Jiao Y.Gonadotropin-inhibitory hormone(GnIH)and its receptor in the female pig:cDNA cloning,expression in tissues and expression pattern in the reproductive axis during the estrous cycle[J].Peptides,2012,36(2):176-185
[35] Li H,Song H,Huang M,Nie H,Wang Z,Wang F.Impact of food restriction on ovarian development,RFamide-related peptide-3 and the hypothalamic-pituitary-ovarian axis in prepubertal ewes[J].Reproduction in Domestic Animals,2014,49(5):831-838.
[36]李卉.RFRP-3在幼龄绵羊组织中的表达及其参与繁殖营养调控的研究[D].江苏:南京农业大学,2014Li H.Expression of RFRP-3 in young sheep tissues and its participation in reproductive regulation[D].Jiangsu:Nanjing Agricultural University,2014(in Chinese)
[37]甘超,胡深强,苟华,袁鑫,王继文,李亮.鹅GnIH受体基因的克隆及组织表达特性研究[J].中国家禽,2014,36(1):4Gan C,Hu S Q,Yan H,Yuan X,Wang J W,Li L.Cloning and characterization of GnIH receptor genes in geese[J].Chinese Poultry,2014,36(1):4(in Chinese)
[38]侯玉晶,雷治海,苏娟,姚远,李月勤,于晓磊,刘倩.兔神经肽S及其受体基因的克隆,组织表达及热应激对其表达的影响[J].中国农业科学,2011,44(12):2589-2600Hou Y J,Lei Z H,Su J,Yao Y,Li Y Q,Yu X L,Liu Q.Cloning,tissue expression and effect of heat stress on expression of rabbit neuropeptide S and its receptors[J].Chinese Agricultural Science,2011,44(12):2589-2600(in Chinese)
[39] Clarke I J,Sari I P,Qi Y,Smith J T,Parkington H C,Ubuka T,Iqbal J,Li Q,Tilbrook A,Morgan K,Pawson A J,Tsutsui K,Millar R P,Bentley G E.Potent action of RFamide-relatedpeptide-3 onpituitarygonadotropes indicative of a hypophysiotropic role in the negative regulation of gonadotropin secretion[J].Endocrinology,2008,149(11):5811-5821
[40] Ancel C,Bentsen A H,Sébert M E,Tena-Sempere M,Mikkelsen J D,Simonneaux V.Stimulatory effect of RFRP-3on the gonadotrophic axis in the male Syrian hamster:The exception proves the rule[J].Endocrinology,2012,153(3):1352-1363
[41] Dardente H,Birnie M,Lincoln G A, Hazlerigg D G.RFamide-related peptide and its cognate receptor in the sheep:cDNAcloning, mRNAdistributioninthe hypothalamus and the effect of photoperiod[J].Journal of Neuroendocrinology,2008,20(11):1252-1259
[42] 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 in kisspeptin and RFamide-related peptide(RFRP)expression and terminal connections to gonadotropin-releasing hormone neurons in the brain:A novel medium for seasonal breeding in the sheep[J].Endocrinology,2008,149(11):5770-5782
[43] Oishi H,Klausen C,Bentley G E,Osugi T,Tsutsui K,Gilks C B,Yano T,Leung P C K.The human gonadotropininhibitory hormone ortholog RFamide-related peptide-3suppresses gonadotropin-induced progesterone production in human granulosa cells[J].Endocrinology,2012,153(7):3435-3445
[44] Beymer M,Henningsen J,Bahougne T,Simonneaux V.The role of kisspeptin and RFRP in the circadian control of female reproduction[J].Molecular and Cellular Endocrinology,2016,438(15):89-99
[45] Singh P,Krishna A,Sridaran R,Tsutsui K.Immunohistochemical localization of GnRH and RFamide-related peptide-3in the ovaries of mice during the estrous cycle[J].Journal of Molecular Histology,2011,42(5):371
[46]龚金秋,阳美霞,唐娇美,曾杰,王水莲.RFRP-3对哺乳动物生殖激素的调节作用[J].中兽医医药杂志,2017,36(4):31-33Gong J Q,Yang M X,Tang J M,Zeng J,Wang S L.Regulatory effects of RFRP-3 on reproductive hormones in mammals[J].Journal of Veterinary Medicine,2017,36(4):31-33(in Chinese)
[2] Lee J H,Miele M E,Hicks D J,Phillips K K,Trent J M,Weissman B E,Welch D R.KiSS-1,a novel human malignant melanoma metastasis-suppressor gene[J].Journal of the National Cancer Institute,1996,88(23):1731-1737
[3] Ohtaki T,Shintani Y,Honda S,Matsumoto H,Hori A,Kanehashi K,Ohtaki T L,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 peptide ligand of a G-protein-coupled receptor[J].Nature,2001,411(6837):613-617
[4] Novaira H J, Ng Y, Wolfe A,Radovick S.Kisspeptin increases GnRH mRNA expression and secretion in GnRH secreting neuronal cell lines[J].Molecular and Cellular Endocrinology,2009,311(1-2):126-134
[5] Papaoiconomou E,Msaouel P,Makri A,Diamanti-Kandarakis E,Koutsilieris M.The role of kisspeptin/GPR54in the reproductive system[J].In Vivo,2011,25(3):343-354
[6]郭若婷,鲁慧文,欧科鹏,王颖,刘乙,顾真真,刘小军.绵羊Otx-2基因的分子克隆、组织表达及其与季节性繁殖关系的研究[J].畜牧兽医学报,2015,46(5):696-703Guo R T,Lu H W,Ou K P,Wang Y,Liu Y,Gu Z Z,Liu X J.Molecular cloning,tissue expression and its relationship with seasonal reproduction of Otx-2gene in sheep[J].Journal of Animal and Veterinary Sciences,2015,46(5):696-703(in Chinese)
[7] Thompson E L,Patterson M,Murphy K G,Smith K L,Dhillo W S,Todd J F,Ghatei M A,Bloom S R.Central and peripheral administration of kisspeptin-10 stimulates the hypothalamic-pituitary-gonadalaxis[J]. Journal of Neuroendocrinology,2004,16(10):850-858
[8] Messager S,Chatzidaki E E,Ma D,Hendrick A G,Zahn D,Dixon J,Thresher R R,Malinge I,Lomet D,Carlton M B L,Colledge W H,Caraty A C,Aparicio S A J R.Kisspeptin directly stimulates gonadotropin-releasing hormone release via G protein-coupled receptor 54[J]. Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1761-1766
[9] Caraty A,Smith J T,Lomet D,Ben Sa6d S,Morrissey A,Cognie J,Doughton G,Baril C B,Clarke I J.Kisspeptin synchronizes preovulatory surges in cyclical ewes and causes ovulation in seasonally acyclic ewes[J].Endocrinology,2007,148(11):5258-5267
[10] Tsutsui K,Saigoh E,Ukena K,Teranishi H,Fujisawa Y,Kikuchi M,Ishii S,Sharp P J.A novel avian hypothalamic peptide inhibiting gonadotropin release[J].Biochemical and Biophysical Research Communications,2000,275(2):661-667.
[11]向伟,赖平,张宝云,王凭青,储明星,樊奇,刘雪旭,潭颖.RFRP-3对哺乳动物生殖功能和能量平衡的影响[J].遗传,2012,34(8):969-976Xiang W,Lai P,Zhang B Y,Wang P Q,Chu M X,Fan Q,Liu C X,Tan Y.Effects of RFRP-3 on reproductive function and energy balance in mammals[J].Genetics,2012,34(8):969-976(in Chinese)
[12] Ducret E,Anderson G M,Herbison A E.RFamide-related peptide-3,a mammalian gonadotropin-inhibitory hormone ortholog,regulates gonadotropin-releasing hormone neuron firing in the mouse[J].Endocrinology,2009,150(6):2799-2804
[13] Wu M,Dumalska I,Morozova E,Van Den Pol A N,Alreja M.Gonadotropin inhibitory hormone inhibits basal forebrain vGluT2-gonadotropin-releasing hormone neurons via a direct postsynaptic mechanism[J].The Journal of Physiology,2009,587(7):1401-1411
[14] Smith J T.The role of kisspeptin and gonadotropin inhibitory hormone in the seasonal regulation of reproduction in sheep[J].Domestic Animal Endocrinology,2012,43(2):75-84
[15] Livak K J,Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the2-ΔΔCT method[J].Methods,2001,25(4):402-408
[16] Schmittgen T D,Livak K J.Analyzing real-time PCR data by the comparative CT method[J].Nature Protocols,2008,3(6):1101-1108
[17]张伟,刘晓娜,邓双义,王世银,石国庆,阿勒泰羊不同繁殖状态下丘脑Kiss1基因表达规律与其季节性繁殖的关联性分析[J].基因组学与应用生物学,2016,35(1):45-52Zhang W,Liu X N,Deng S Y,Wang S Y,Shi G Q.Correlation analysis of Kiss1gene expression and seasonal reproduction in the hypothalamus of Altay sheep[J].Genomics and Applied Biology,2016,35(1):45-52(in Chinese)
[18]黄冬维,储明星,狄冉,刘秋月,胡文萍,王翔宇,潘章源,郭晓飞.山羊繁殖季节性相关基因KiSS-1与RFRP的表达研究[J].畜牧兽医学报,2014,46(6):924-931Huang D W,Chu M X,Di R,Liu Q Y,Hu W P,Wang X Y,Pan Z Y,Guo X F.Study on expression of KiSS-1 and RFRPin seasonal goat breeding[J].Journal of Animal and Veterinary Sciences,2014,46(6):924-931(in Chinese)
[19]朱碧泉,李卓昭,吴德,方正锋,林燕,车炼强.梅山与长大母猪下丘脑—垂体—卵巢轴Kiss1和GPR54基因的表达差异[D].中国畜牧杂志,2011,47(13):21-25Zhu B C,Li ZZ,Wu D,Fang Z F,Lin Y,Che L Q.Differences in expression of Kiss1 and GPR54 genes in Meishan and growing sows′hypothalamus-pituitary-ovary axis[D].Chinese Journal of Animal Science,2011,47(13):21-25(in Chinese)
[20] Tomikawa J,Homma T,Tajima S,Shibata T,Inamoto Y,Takase K,Inoue N,Ohkura S,Uenoyama Y,Maeda K,Tsukamura H. Molecular characterization and estrogen regulation of hypothalamic KiSS1 gene in the pig[J].Biology of Reproduction,2010,82(2):313-319
[21] Xiong J,Karsch F J,Lehman M N.Evidence for seasonal plasticity in the gonadotropin-releasing hormone(GnRH)system of the ewe:Changes in synaptic inputs onto GnRH neurons[J].Endocrinology,1997,138(3):1240-1250
[22] Merkley C M,Coolen L M,Goodman R L,Lehman M N.Evidence for changes in numbers of synaptic inputs onto KNDy and GnRH neurones during the preovulatory LH surge in the ewe[J].Journal of Neuroendocrinology,2015,27(7):624-635
[23] El-Tarabany M S,Zaglool A W,El-Tarabany A A,Awad A.Association analysis of polymorphism in KiSS1gene with reproductive traits in goats[J]. Animal Reproduction Science,2017,180:92-99
[24] Messager S,Chatzidaki E E,Ma D,Hendrick A G,Zahn D,Dixon J,Thresher R R,Malinge I,Lomet D,Carlton M B L,Colledge W H,Caraty A,Colledge W H.Kisspeptin directly stimulates gonadotropin-releasing hormone release via G protein-coupled receptor 54[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1761-1766
[25]王军,孙蕾,吕文发,娄玉杰.KiSS-1/GPR54系统在不同发情周期母羊下丘脑中的表达规律[J].西北农林科技大学学报:自然科学版,2011,39(12):49-52Wang J,Sun L,Lu W F,Qi Y J.Expression patterns of KiSS-1/GPR54system in different hypothalamus of estrous cycle in ewe[J].Journal of Northwest A&F University:Natural Science Edition,2011,39(12):49-52(in Chinese)
[26] Robert L.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,Clarke I J.Kisspeptin neurons in the arcuate nucleus of the ewe express both dynorphin A and neurokinin B[J].Endocrinology,2007,148(12):5752-5760
[27] Wagner G C,Johnston J D,Clarke I J,Lincoln G A,Hazlerigg D G. Redefining the limits of day length responsiveness in a seasonal mammal[J].Endocrinology,2007,149(1):32-39
[28] Luridiana S,Mura M C,Daga C,Cosso G,Bodano S,Farci F,Zidda F,Carcangiu V.Influences of melatonin treatment,melatonin receptor 1A(MTNR1A)and kisspeptin(KiSS-1)gene polymorphisms on first conception in Sarda ewe lambs[J].Reproduction Fertility and Development,2016,28(6):750-756
[29] Smith J T,Clarke I J.Seasonal breeding as a neuroendocrine model for puberty in sheep[J].Molecular and Cellular Endocrinology,2010,324(1-2):102-109
[30] Jafarzadeh Shirazi M R,Zamiri M J,Salehi M S,Moradi S,Tamadon A,Namavar M R,Akhlaghi A,Tsutsui K,Caraty A.Differential expression of RFamide-related peptide,a mammalian gonadotrophin-inhibitory hormone orthologue,and kisspeptin in the hypothalamus of Abadeh ecotype does during breeding and anoestrous seasons[J].Journal of Neuroendocrinology,2014,26(3):186-194
[31] Beskonakli E,Palaoglu S,Aksaray S,Alanoglu G,Turhan T,Taskin Y.Effect of pinealectomy on immune parameters in ratswith Staphylococcus aureus infection[J].Neurosurgical Review,2001,24(1):26-30
[32] Tsutsui K,Bentley G E,Kriegsfeld L J,Osugi T,Seong J Y, Vaudry H. Discovery and evolutionary history of gonadotrophin-inhibitory hormone and kisspeptin:New key neuropeptides controlling reproduction[J]. Journal of Neuroendocrinology,2010,22(7):716-727
[33]王梁,刘国世,史文清,史建民,高超,田秀芝,张璐,赵广华,商业,云鹏,朱世恩.外源褪黑激素对梅花鹿超数排卵及体内促性腺激素的影响[J].中国鹿业进展,2011Wang L,Liu G S,Shi W Q,Shi J M,Gao C,Tian X Z,Zhang L,Zhao G H,Shang Y,Yun P,Zhu S E.Effects of exogenous melatonin on superovulation and gonadotropin in Sika deer[J].China Deer Industry Progress,2011(in Chinese)
[34] Li X,Su J,Lei Z,Zhao Y,Jin M,Fang R,Zheng L C,Jiao Y.Gonadotropin-inhibitory hormone(GnIH)and its receptor in the female pig:cDNA cloning,expression in tissues and expression pattern in the reproductive axis during the estrous cycle[J].Peptides,2012,36(2):176-185
[35] Li H,Song H,Huang M,Nie H,Wang Z,Wang F.Impact of food restriction on ovarian development,RFamide-related peptide-3 and the hypothalamic-pituitary-ovarian axis in prepubertal ewes[J].Reproduction in Domestic Animals,2014,49(5):831-838.
[36]李卉.RFRP-3在幼龄绵羊组织中的表达及其参与繁殖营养调控的研究[D].江苏:南京农业大学,2014Li H.Expression of RFRP-3 in young sheep tissues and its participation in reproductive regulation[D].Jiangsu:Nanjing Agricultural University,2014(in Chinese)
[37]甘超,胡深强,苟华,袁鑫,王继文,李亮.鹅GnIH受体基因的克隆及组织表达特性研究[J].中国家禽,2014,36(1):4Gan C,Hu S Q,Yan H,Yuan X,Wang J W,Li L.Cloning and characterization of GnIH receptor genes in geese[J].Chinese Poultry,2014,36(1):4(in Chinese)
[38]侯玉晶,雷治海,苏娟,姚远,李月勤,于晓磊,刘倩.兔神经肽S及其受体基因的克隆,组织表达及热应激对其表达的影响[J].中国农业科学,2011,44(12):2589-2600Hou Y J,Lei Z H,Su J,Yao Y,Li Y Q,Yu X L,Liu Q.Cloning,tissue expression and effect of heat stress on expression of rabbit neuropeptide S and its receptors[J].Chinese Agricultural Science,2011,44(12):2589-2600(in Chinese)
[39] Clarke I J,Sari I P,Qi Y,Smith J T,Parkington H C,Ubuka T,Iqbal J,Li Q,Tilbrook A,Morgan K,Pawson A J,Tsutsui K,Millar R P,Bentley G E.Potent action of RFamide-relatedpeptide-3 onpituitarygonadotropes indicative of a hypophysiotropic role in the negative regulation of gonadotropin secretion[J].Endocrinology,2008,149(11):5811-5821
[40] Ancel C,Bentsen A H,Sébert M E,Tena-Sempere M,Mikkelsen J D,Simonneaux V.Stimulatory effect of RFRP-3on the gonadotrophic axis in the male Syrian hamster:The exception proves the rule[J].Endocrinology,2012,153(3):1352-1363
[41] Dardente H,Birnie M,Lincoln G A, Hazlerigg D G.RFamide-related peptide and its cognate receptor in the sheep:cDNAcloning, mRNAdistributioninthe hypothalamus and the effect of photoperiod[J].Journal of Neuroendocrinology,2008,20(11):1252-1259
[42] 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 in kisspeptin and RFamide-related peptide(RFRP)expression and terminal connections to gonadotropin-releasing hormone neurons in the brain:A novel medium for seasonal breeding in the sheep[J].Endocrinology,2008,149(11):5770-5782
[43] Oishi H,Klausen C,Bentley G E,Osugi T,Tsutsui K,Gilks C B,Yano T,Leung P C K.The human gonadotropininhibitory hormone ortholog RFamide-related peptide-3suppresses gonadotropin-induced progesterone production in human granulosa cells[J].Endocrinology,2012,153(7):3435-3445
[44] Beymer M,Henningsen J,Bahougne T,Simonneaux V.The role of kisspeptin and RFRP in the circadian control of female reproduction[J].Molecular and Cellular Endocrinology,2016,438(15):89-99
[45] Singh P,Krishna A,Sridaran R,Tsutsui K.Immunohistochemical localization of GnRH and RFamide-related peptide-3in the ovaries of mice during the estrous cycle[J].Journal of Molecular Histology,2011,42(5):371
[46]龚金秋,阳美霞,唐娇美,曾杰,王水莲.RFRP-3对哺乳动物生殖激素的调节作用[J].中兽医医药杂志,2017,36(4):31-33Gong J Q,Yang M X,Tang J M,Zeng J,Wang S L.Regulatory effects of RFRP-3 on reproductive hormones in mammals[J].Journal of Veterinary Medicine,2017,36(4):31-33(in Chinese)