初情期前母猪性腺外促性腺激素受体表达的研究
摘要
本研究系统评述了促卵泡素受体(FSHR)和促黄体素受体(LH/CGR)以及能量对繁殖性能影响的研究进展,探讨了初情期前母猪子宫、输卵管中FSHR和LH/CGR的分布与活性,研究了不同日粮能量水平对子宫、输卵管FSHR和LH/CGR表达的影响。研究发现,初情期前母猪子宫、输卵管中存在功能性的FSHR和LH/CGR,FSHR存在于子宫内膜上皮细胞和输卵管粘膜上皮细胞,LH/CGR存在于子宫内膜上皮细胞、子宫肌层血管平滑肌细胞和输卵管粘膜上皮细胞。促性腺激素可能直接调节子宫、输卵管上相应受体表达,也可能通过卵巢上分泌的性腺激素间接调节。不同日粮能量水平能够影响初情期前母猪外周血清FSH、LH、生长激素(GH)、瘦素、胰岛素(INS)及胰岛素样生长因子-I(IGF-I)浓度。高能量水平可提高血清FSH、LH、瘦素、GH、INS及IGF-Ⅰ的浓度;低能量水平则降低血清中这些激素的浓度;不同日粮能量水平影响初情期前母猪子宫、输卵管FSHR和LH/CGR的表达。高能量水平具有促进作用,低能量水平则起抑制作用。母猪发育阶段和发情周期不同,子宫、输卵管中FSHR和LH/CGR表达量存在差异。卵泡期子宫中FSHR和黄体期子宫中LH/CGR的表达量均高于初情期前子宫,输卵管中FSHR和LH/CGR的表达量均在卵泡期最高。能量可能通过血清INS、IGF-Ⅰ、瘦素、GH等代谢媒介引起垂体促性腺激素分泌的变化,影响FSHR和LH/CGR的表达。
Two months old crossbred (Landrace×Large White) gilts were used to investigate: 1) the expression and location of the functional FSHR and LH/CGR in both uterus and oviducts. 2) the effects of dietary energy on hormones release in prepubertal gilts 3) the effects of dietary energy on the expression of gonadotropins receptors in extragonadal prepubertal gilts. The present study was to clarify the preliminary molecular biology mechanism of the regulating role of extragonadal gonadotropins in the extragonadal tissues , and to establish theoretical basis for feeding of prepubertal gilts as well.
Firstly, we determined the localization of FHSR and LH/CGR in the uteri and oviducts of prepubertal gilts with immunohistochemical method. The uterus, oviducts and ovaries were collected from five prepubertal gilts. the ovaries were used as the positive control, and the negtive control was non-specific IgG instead of specific antibodies. The results showed that LH/CGR and FSHR were present in the uteri and oviducts of prepubertal gilts. In the uteri, FSHR was expressed in the endometrial epithelial cells, but not in the myometrium and perimetrium. LH/CGR was present in the endometrial epithelial cells and the myometrial vascular smooth muscle. However, there was no LH/CGR in the perimetrium. FSHR and LH/CGR were all expressed in the oviductal mucosal epithelium, but not in the myosalpinx and the serosa. The resluts confirmed that the ovaries was not the only target organ of gonadotropins receptors, and FSHR and LH/CGR were also located in the prepubertal extragonadal organs such as uterus and oviducts.
Secondly, we investigated the expression of functional FHSR and LH/CGR in the prepubertal uterus and oviducts with radioactive immunoassay, and Western blotting. Fifteen growth-matched crossbred prepubertal gilts were randomly allocated to three groups after one week reared in the same room. One group animals (n=5) were performed ovariectomy. Two days later, the fasting vein blood was collected from all three groups. Then we carried out treatments according to the experimental design. The groupⅠgilts were injected with physiological saline (5 ml per gilt); the groupⅡgilts were injected, i.m., with eCG (300IU per gilt); the groupⅢgilts (ovariectomizd) were injected with eCG (300IU per gilt). Twenty four hours after eCG treatment, vein blood was collected from all three groups for serum preparation, uteri and oviducts were also removed for detection. The results showed that the concentrations of serum FSH and LH were significantly increased after eCG injection(p<0.01)in groupⅡand groupⅢ. The ovary excision removed the major source of in vivo estrogen synthesis and significantly decreased the concentration of serum estradiol compared with that of intact gilts(p<0.01). After eCG injection, the concentration of serum estradiol in the ovariectomized gilts was significantly lower than that of saline injection pig (p<0.01). Regardless of ovaries removed or not, eCG injection could promote the expression of FSHR and LH/CGR in the uterus and oviducts. The FSHR expression level of uterus and oviducts in intact gilts was significantly higher than that of ovariectomized gilts. The similar consequence was found in oviducal LH/CGR(p<0.05). These results showed that the FSHR and LH/CGR in prepubertal uterus and oviducts were active and functional, and the expression of LH/CGR and FSHR were regulated by FSH, LH, and the estrogen.
Finally, we observed the effects of dietary energy on the hormones release and the expression of FSHR and LH/CGR on prepubertal gilts with radioactive immunoassay, fluorescence quantitative-PCR and Western blotting. Fifteen gilts were randomly allocated to three groups, the groupⅠfed with the moderate energy level (100%of NRC energy requirements), the groupⅡfed with the low energy level (90% of NRC energy requirements), the groupⅢfed with the diet of high energy level (100%of NRC energy requirements), for 12d. Then the fasting vein blood was collected from all three groups for serum preparation on d1, d5, d9, and d12.Then uterus and oviducts were collected at the slaughter on d12 The results showed that dietary energy could affect the concentrations of peripheral serum FSH, LH, GH, leptin, Ins, and IGF-Ⅰin prepuberty gilts. High energy level significantly enhanced the concentrations of serum FSH, LH, leptin, GH, insulin and IGF-Ⅰ, whereas the change of GH concentration was not significantly(p<0.01). Low energy level significantly reduced the concentrations of serum FSH, LH, GH, leptin, Ins and IGF-Ⅰ. Dietary energy level affected the expression of FSHR and LH/CGR in prepubertal gilts. High energy intake significantly enhanced the expresioon of uterine and oviducal FSHR and LH/CGR, whereas insufficient energy intake restricted FSHR and LH/CGR expression. In the uterus, the expression level of FSHR mRNA was the highest in follicular phase and LH/CGR mRNA expression level was the highest in luteal phase. In the oviducts, The expression level of FSHR mRNA and LH/CGR mRNA were all the highest in follicular phase .
Overall, our results indicated that there were functional FSHR and LH/CGR in uterus and oviducts of prepubertal gilts, and confirmed for the first time that FSHR was present in the endometrium epithelial cells. FSH, LH and estrogens regulated the expression of FSHR and LHR in the uterus and oviducts. Gonadotropins could regulate the uterine and oviductal function not only indirectly through gonadal hormone secreted by ovaries. but also directly through gonadotrophins receptors in the uterus and oviducts. These results imply that dietary energy level could influence the expression of FSHR and LH/CGR in uterus and oviducts which might be mediated by insulin, IGF-Ⅰ, leptin and GH, These results provided the hypothesis that gonadotropins might play direct roles on the extragonadal organs and consummate the mechanism of gonadotropins in animal reproduction. Furthermore, we explored the effect and mechanism of energy on reproductive development of prepubertal gilts from aspects of reproductive endocrinology, gene and protein expression, which might provide a theoretical basis for feeding of the prepubertal gilts.
Two months old crossbred (Landrace×Large White) gilts were used to investigate: 1) the expression and location of the functional FSHR and LH/CGR in both uterus and oviducts. 2) the effects of dietary energy on hormones release in prepubertal gilts 3) the effects of dietary energy on the expression of gonadotropins receptors in extragonadal prepubertal gilts. The present study was to clarify the preliminary molecular biology mechanism of the regulating role of extragonadal gonadotropins in the extragonadal tissues , and to establish theoretical basis for feeding of prepubertal gilts as well.
Firstly, we determined the localization of FHSR and LH/CGR in the uteri and oviducts of prepubertal gilts with immunohistochemical method. The uterus, oviducts and ovaries were collected from five prepubertal gilts. the ovaries were used as the positive control, and the negtive control was non-specific IgG instead of specific antibodies. The results showed that LH/CGR and FSHR were present in the uteri and oviducts of prepubertal gilts. In the uteri, FSHR was expressed in the endometrial epithelial cells, but not in the myometrium and perimetrium. LH/CGR was present in the endometrial epithelial cells and the myometrial vascular smooth muscle. However, there was no LH/CGR in the perimetrium. FSHR and LH/CGR were all expressed in the oviductal mucosal epithelium, but not in the myosalpinx and the serosa. The resluts confirmed that the ovaries was not the only target organ of gonadotropins receptors, and FSHR and LH/CGR were also located in the prepubertal extragonadal organs such as uterus and oviducts.
Secondly, we investigated the expression of functional FHSR and LH/CGR in the prepubertal uterus and oviducts with radioactive immunoassay, and Western blotting. Fifteen growth-matched crossbred prepubertal gilts were randomly allocated to three groups after one week reared in the same room. One group animals (n=5) were performed ovariectomy. Two days later, the fasting vein blood was collected from all three groups. Then we carried out treatments according to the experimental design. The groupⅠgilts were injected with physiological saline (5 ml per gilt); the groupⅡgilts were injected, i.m., with eCG (300IU per gilt); the groupⅢgilts (ovariectomizd) were injected with eCG (300IU per gilt). Twenty four hours after eCG treatment, vein blood was collected from all three groups for serum preparation, uteri and oviducts were also removed for detection. The results showed that the concentrations of serum FSH and LH were significantly increased after eCG injection(p<0.01)in groupⅡand groupⅢ. The ovary excision removed the major source of in vivo estrogen synthesis and significantly decreased the concentration of serum estradiol compared with that of intact gilts(p<0.01). After eCG injection, the concentration of serum estradiol in the ovariectomized gilts was significantly lower than that of saline injection pig (p<0.01). Regardless of ovaries removed or not, eCG injection could promote the expression of FSHR and LH/CGR in the uterus and oviducts. The FSHR expression level of uterus and oviducts in intact gilts was significantly higher than that of ovariectomized gilts. The similar consequence was found in oviducal LH/CGR(p<0.05). These results showed that the FSHR and LH/CGR in prepubertal uterus and oviducts were active and functional, and the expression of LH/CGR and FSHR were regulated by FSH, LH, and the estrogen.
Finally, we observed the effects of dietary energy on the hormones release and the expression of FSHR and LH/CGR on prepubertal gilts with radioactive immunoassay, fluorescence quantitative-PCR and Western blotting. Fifteen gilts were randomly allocated to three groups, the groupⅠfed with the moderate energy level (100%of NRC energy requirements), the groupⅡfed with the low energy level (90% of NRC energy requirements), the groupⅢfed with the diet of high energy level (100%of NRC energy requirements), for 12d. Then the fasting vein blood was collected from all three groups for serum preparation on d1, d5, d9, and d12.Then uterus and oviducts were collected at the slaughter on d12 The results showed that dietary energy could affect the concentrations of peripheral serum FSH, LH, GH, leptin, Ins, and IGF-Ⅰin prepuberty gilts. High energy level significantly enhanced the concentrations of serum FSH, LH, leptin, GH, insulin and IGF-Ⅰ, whereas the change of GH concentration was not significantly(p<0.01). Low energy level significantly reduced the concentrations of serum FSH, LH, GH, leptin, Ins and IGF-Ⅰ. Dietary energy level affected the expression of FSHR and LH/CGR in prepubertal gilts. High energy intake significantly enhanced the expresioon of uterine and oviducal FSHR and LH/CGR, whereas insufficient energy intake restricted FSHR and LH/CGR expression. In the uterus, the expression level of FSHR mRNA was the highest in follicular phase and LH/CGR mRNA expression level was the highest in luteal phase. In the oviducts, The expression level of FSHR mRNA and LH/CGR mRNA were all the highest in follicular phase .
Overall, our results indicated that there were functional FSHR and LH/CGR in uterus and oviducts of prepubertal gilts, and confirmed for the first time that FSHR was present in the endometrium epithelial cells. FSH, LH and estrogens regulated the expression of FSHR and LHR in the uterus and oviducts. Gonadotropins could regulate the uterine and oviductal function not only indirectly through gonadal hormone secreted by ovaries. but also directly through gonadotrophins receptors in the uterus and oviducts. These results imply that dietary energy level could influence the expression of FSHR and LH/CGR in uterus and oviducts which might be mediated by insulin, IGF-Ⅰ, leptin and GH, These results provided the hypothesis that gonadotropins might play direct roles on the extragonadal organs and consummate the mechanism of gonadotropins in animal reproduction. Furthermore, we explored the effect and mechanism of energy on reproductive development of prepubertal gilts from aspects of reproductive endocrinology, gene and protein expression, which might provide a theoretical basis for feeding of the prepubertal gilts.
引文
[1]楼航应.卵巢颗粒细胞FSHR的表达与IVF周期卵巢反应性及妊娠结局关系的研究[D].浙江:浙江大学医学院,2003.
[2]李俊杰,贾青,田树军.雌性哺乳动物LH及其受体的研究进展[J].黄牛杂志,2003,29(3):26-29.
[3]李俊杰,贾青,张正珊.雌性哺乳动物促卵泡素受体与促黄体素受体的结构与表达[J]..生命的化学,2003,23(4):289-291.
[4] SPRENGEL R, BRAUN T, NIKOLICS K, et al. The testicular receptor for follicle stimulating hormone: structure and functional expression of cloned Cdna [J]. Mol Endocrinol, 1990, 4(4):525-530.
[5] GUDERMANN T, NURNBERG B, SCHULTZ G. Receptors and G proteins as primary components of transmembrane signal transduction. Part 1. G-protein-coupled receptors: structure and function [J]. J Mol Med, 1995, 73(2):51-63.
[6] KENNELLY P J, KERBS E G.. Consensus sequences as substrste specificity determinants for protein kinases and protain phosphatases [J]. J Biol Chem, 1991, 266:15555-15558.
[7] QUINTANA J, HIPKIN RW, SANCHEZ-YAGUE J, et al. Follitropin (FSH) and a phorbol ester stimulate the phosphorylation of the FSH receptor in intact cells [J]. J Biol Chem, 1994, 269:8772-8779.
[8] MINEGISHI T, IGARASHI S, NAKAMURA K, et al. Functional expression of the recombinant human FSH receptor [J]. J Endocrinol, 1994, 141(2):369-375.
[9] VANNIER B, LOOSFELT H, MEDYRI G, et al. Anti-human FSH receptor monoclonal antibodies: immunochemical and immunocytochemical characterization of the receptor [J]. Biochemistry, 1996, 35(5):1358-1366.
[10] GROMOLL J, RIED T, HOLTGREVE-GREZ H, et al. Localization of the human FSH receptor to chromosome 2 p21 using a genomic probe comprising exon 10 [J]. J Mol Endocrinol, 1994, 12:265-271.
[11] ROUSSEAU-MERCK M F, MISRAHI M, LOOSFELT H, et al. Localization of the human luteinizing hormone receptor gene to chromosome 2 p21 [J]. Cytogenet Cell Genet, 1990, 54: 77-79.
[12] ROUEEEAU-MERCK M F, MISRAHI M, LOOSFELT H, et al. The chromosomal localization of the human follicle-stimulating hormone receptor gene on 2p21-p16 is similar to that of the luteinizing hormone receptor gene [J]. Genomics, 1993,15: 222-224.
[13] HECKERT L. L., GRISWOLD M. D. Expression of follicle-stimulating hormone receptor mRNA in rat testes and Sertolicells [J]. Molec Endocr, 1991, 5:670-677.
[14] HECKERT L L, GRISWOLD M D. Expression of the FSH receptor in the testis [J]. Rec.Prog. Horm. Res, 1992, 48: 61-78.
[15] LAPOLT P S, TILLY J L, AIHARA T, et al. Gonadotropin-induced up and down regulation of ovarian follicle stimulating hormone (FSH) receptor gene expression in immature rats: effects of pregnant mare's serum gonadotropin, human chorionic gonadotropin and recombinant FSH [J]. Endocrinology, 1992, 130: 1289-1295.
[16] CAMP T A, RAHAL J O. MAYO K E, Cellular localization and hormonal regulation of follicle stimulating hormone and luteinizing hormone receptor messenger RNAs in the rat ovary [J]. Molec Endocr, 1991, 5: 1405-1417.
[17] AITTOMAKI K, LUCENA J L, PAKARINEN P, et al. Mutation in the follicle stimulati- ng hormone receptor gene causes hereditary hypergonadotropic ovarian failure [J]. Cell, 1995, 82(6):959-968.
[18] MEANS A R, VAITUKAITIS J. Peptide hormone“receptor”: specific binding of 3 H-FSH to testis [J]. Endocrinology, 1972, 90(1):39-46.
[19] RANNIKKO A, PENTTILA T L, ZHANG F P, et al. Stage-specific expression of the FSH receptor gene in the prepubertal and adult rat seminiferous epithelium [J]. J Endocrinol. 1996, 151(1):29-35.
[20] ORTH J, CHRISTENSEN A K. Autoradiographic localization of specifically bound 125I-labeled follicle-stimulating hormone on spermatogonia of the rat testis [J]. Endocrinology. 1978, 103(5): 1944-1951.
[21] RANNIKKI A S, ZHANG F P, HUHTANIEMI I T. Ontogeny of follicle-stimulating hormone receptor gene expression in the rat testis and ovary [J]. Mol Cell Endocrinol, 1995, 107(2): 199-208.
[22] SASAKI M,YAMAMOTO M,ARISHIMA K,et al.Effects of follicle stimulating hormone on intermediate filaments and cell division of Sertoli cellsof fetal rat testis in culture [J].J Vet Med Sci, 1998, 60 (1):35-39.
[23]陈雪雁,陈克铨,陈实平. FSH受体的特性和表达的调控[J].解剖学报. 1999, 30(3):284-287.
[24]丁允闵,叶世钧,陈咏梅,等.大鼠支持细胞生后发育过程中卵泡刺激素受体mRNA水平的动态观察[J].中国医学科学院学报, 1996, 18(2):99-103.
[25] SOKKA T, HUHTANIEMI I. Ontogeny of gonadotrophin receptors and gonadotrophin stimulated cyclic AMP production in the neonatal rat ovary [J]. J Endocrinol, 1990, 127(2):297-303.
[26] O'SHAUGHNESSY P J, MARSH P, DUDLEY K. Follicle-stimulating hormone receptor mRNA in the mouse ovary during post-natal development in the normal mouse and in the adult hypogonadal (hpg) mouse: structure of alternate transcripts [J]. Mol Cell Endocrinol, 1994, 101(1-2):197-201.
[27] O'SHAUGHNESSY P J, DUDLEY K, RAJAPAKSHA W R. Expression of follicle stimulating hormone-receptor mRNA during gonadal development [J]. Mol Cell Endocrinol, 1996, 125(1-2):169-175.
[28] MARGRET W Z, MAGID M S, KRAMER E E, et al. Follicle-stimulating hormone receptor is expressed in human ovarian surface epithelium and fallopian tube. Am J Pathol, 1996, 148(1):47-53.
[29] DUNKEL L, TILLY J L, SHIKONE T, et al. Follicle-stimulating hormone receptor expression in the rat ovary: increases during prepubertal development and regulation by the opposing actions of transforming growth factors beta and alpha [J]. Biol Reprod, 1994, 50(4):940-948.
[30] MINEGISHI T, TANO M, IGARASHI M, et al. Expression of follicle-stimulating hormone receptor in human ovary [J]. Eur J Clin Invest, 1997, 27(6):469-474.
[31] ZHANG C q, SHIMADA K, SAITO N. Expression of Messenger Ribonucleic Acids of Luteinizing Hormone and Follicle-Stimulating Hormone Receptors in Granulosa and Theca Layers of Chicken Preovulatory Follicles [J]. 1997, 105: 402-409.
[32] MIZARACHI D, SHEMESH M. Follicle-Stimulating Hormone Receptor and Its Messenger Ribonucleic Acid Are Present in the Bovine Cervix and Can Regulate Cervical Prostanoid Synthesis [J]. Biology of reproduction, 1999, 61:776-784.
[33] SHEMESH M, MIZRACHI D, GUREVICH M, et al. Functional importance of bovine myometrial and vascular LH receptors and cervical FSH receptors [J]. Semin Reprod Med,2001, 19(1):87-96.
[34] DIAS J A, LINDAU-SHEPARD B, HAUER C, et al. Human follicle-stimulating hormone structure- activity relationships [J]. Biol Reprod 1998, 58:1331-1336.
[35] KORNYEI J L, LI X, LEI Z M, et al. Restoration of human chorionic gonadotropin response in human myometrial smooth muscle cells by treatment with follicle-stimulating hormone (FSH): evidence for the presence of FSH receptors in human myometrium [J]. European Journal of Endocrinology, 1996, 134 (2) :225-231.
[36] GALWAY A B, LAPOLT P S, TSAFRIRI A, et al. Recombinant follicle-stimulating hormone induces ovulation and tissue plasminogen activator expression in hypophysectomized rats [J]. Endocrinology, 1990, 127(6):3023-3028.
[37] LAPOLT P S, TILLY J L, AIHARA T, et al. Gonadotropin-induced up- and down-regulation of ovarian follicle-stimulating hormone (FSH) receptor gene expression in immature rats: effects of pregnant mare's serum gonadotropin, human chorionic gonadotropin, and recombinant FSH[J]. Endocrinology, 1992, 130(3):1289-1295.
[38] TANO M, MINEGISHI T, KISHI H, et al. The effect of follicle-stimulating hormone (FSH) on the expression of FSH receptor in cultured rat granulosa cells [J]. Life Sci, 1999, 64(12):1063-1069.
[39] MINEGISHI T, TANO M, KISJI H, et al. Follicle-stimulating hormone regulation on its receptor messenger ribonucleic acid levels in cultured rat granulosa cells [J]. Biochim Biophys Acta, 1997, 1359(2):165-173.
[40] MURPHY B D, Dobias M. Homologous and heterologous ligands downregulate follicle-stimulating hormone receptor mRNA in porcine granulosa cells [J]. Mol Reprod Dev, 1999, 53(2):198-207.
[41] MINEGISHI T, TANO M, NAKAMURA K, et al. Regulation of follicle-stimulating hormone receptor [J]. Horm Res., 1996;46(1):37-44.
[42] O'SHAUGHNESSY P J, GRAY S A. Gonadotropin-dependent and gonadotropin independent development of inhibin subunit messenger ribonucleic acid levels in the mouse ovary [J]. Endocrinology, 1995, 136(5):2060-2965.
[43] KUMAR T R, WANG Y. LU N, et al.. Follicle stimulating hormone is required for ovarian follicle maturation but not male fertility [J]. Nat.Genet. 1997, 15(2):201-204.
[44] RICHARDS J S. Maturation of ovarian follicles: actions and interactions of pituitary and ovarianhormones on follicular cell development [J]. Recent Prog Horm Res, 1980, 32: 477-527.
[45] REILLY C M, CANNADY W E, MAHESH V B, et al. Duration of estrogen exposure prior to follicle-stimulating hormone stimulation is critical to granulosa cell growth and differentiation in rats [J]. Biology of reproduction, 1996, 54:1336-1342.
[46] WEIL S, VENDOLA K, ZHOU J, et al. Androgen and follicle-stimulating hormone interactions in primate ovarian follicle development. J. Clin Endocrinol Metab, 1999, 84(8):2951-2956.
[47] CARDENAS H, HERRICK J R, POPE W F, Increased ovulation rate in gilts treated with dihydrotestosterone [J]. Reproduction, 2002, 123(4):527-533.
[48] MINEGISHI T, Hirakawa T, KISHI H. et al. A role of insulin-like growth factor I for follicle-stimulating hormone receptor expression in rat granulosa cells [J]. Biol Reprod, 2000, 62(2):325-333.
[49] OTSUKA F, YAMAMOTO S, ERICKSON G F, et al. Bone morphogenetic protein-15 inhibits follicle-stimulating homone (FSH) action by suppressing FSH receptor expression [J]. J Biol Chem, 2001, 276(14):11387-11392.
[50] TILLY J L, LAPOLT P S, HSUEH A J. Hormonal regulation of follicle-stimulating hormone receptor messenger ribonucleic acid levels in cultured rat granulosa cells [J]. Endocrinology, 1992, 130(3):1296-1302.
[51] FINDLAY, J K. An update on the roles of inhibin, activin and follistatin as local regulators of folliculogenesis [J]. Biol Reprod, 1993, 48:15-23.
[52] MENON K M J, MUNSHI U M, C louser C L, et al. Regulation of Luteinizing Hormone/ Human Chorionic Gonadotropin Receptor Expression: A Perspective1 [J]. Biol Reprod, 2004, 70: 861-866.
[53] MCFARLAND K C, SPRENGEL R, PHILLIPS H S, et al. Lutropin-choriogonadotropin receptor: an unusual member of the G protein-coupled receptpr family [J]. Science, 1989, 245(4917): 494-499.
[54] KISHI M, LIU X, HIRAKAWA T, et al. Identification of two distinct structural motifs that, when added to the C-terminal tail of the rat LH receptor, redirect the internalized hormone-receptor complex from a degradation to a recycling pathway [J]. Mol Endocrinol, 2001, 15: 1624-1635.
[55] TSAI-MORRIS C H, WEI W, Buczko E, et al.. Promoter and regulatory regions of the ratluteinizing hormone receptor gene [J]. J Biol Chem, 1993, 268:4447-4452.
[56] TSAI-MORRIS C H, GEN Y, XIE X Z, et al. Transcriptional protein binding domains governing basal expression of the rat luteinizing hormone receptor gene [J]. J Biol Chem, 1994, 269:15868-15875.
[57] TSAI-MORRIS C H, GEN Y, BUCZKO E, et al. Characterozation of diverse functionalelements in the upstream Sp1 domain of the rat luteinizing hormone receptor gene promoter [J]. J Biol Chem, 1995, 270:7487-7494.
[58] ZHANG F P, HAMALAINEN T, KAIPIA A, et al. Ontogeny of luteinizing hormone receptor gene expression in the rat testis [J]. Endocrinology, 1994, 134(5):2206-2213.
[59] TENA-SEMPERE M, ZHANG F P , HHTANIEMI I. Persistent expression of a truncated form of the lueinizing hormone receptor messenger ribonucleic acid in the rat testis after selective leydig cell destruction by ethylene dimethane sulfonate [J]. Endodrinol, 1994, 135:1018-1024.
[60] SOKKA T A, HAMALAINEN T M, KAIPIA A, et al. Development of Luteinizing Hormone Action in the Perinatal Rat Ovary [J]. Bio Reprod, 1996, 55:663-670.
[61] O'SHAUGHNESSY P J, MCLELLAND D, MCBRIDE M W. Regulation of Luteiniz ing Horm one-Receptor and Follicle-Stimulating Hormone-Receptor Messenger Ribonucleic Acid Levels during Development in the Neonatal Mouse Ovary [J]. Bio Reprod, 1997, 57:602-608.
[62] BAO B, GARVERICK H A, SMITH G W, et al. Changes in messenger ribonucleic acid encoding luteinizing hormone receptor, cytochrome P450-side chain cleavage, and aromatase are associated with recruitment and selection of bovine ovarian follicles [J]. Biol Reprod, 1997, 56(5):1158-1168.
[63] ABDENNEBI L, MONGET P, PISSELET C, et al. Comparative expression of luteinizing hormone and follicle-stimulating hormone receptors in ovarian follicles from high and low prolific sheep breeds [J]. Biol Reprod, 1999, 60(4):845-854.
[64] DERECKA K, PIETILA E M, RAJANIEMI H J, et al. Cycle-dependent LH/hCG receptor gene expression in porcine nongonadal reproductive tissues [J]. J Physiol Pharmacol, 1995, 46(1):77-85.
[65] LIN J, LEI Z M, LOJUN S, et al. Increased expression of luteinizing hormone/ human chorionic gonadotropin receptor gene in human endometrial carcinomas [J]. J ClinEndocrinol Metab, 1994,79:1483-1491.
[66] HAN S W, LEI Z M, RAO CH V, Homologous down-regulation of luteinizing hormone/chorionic gonadotropin receptors by increasing the degradation of receptor transcripts in human uterine endometrial stromal cells. Biol. Reprod. 1997, 57:158-164.
[67] SHEMESH M, MIZRACHI D, GUREVICH M, et al. Expression of functional luteinizing hormone (LH) receptor and its messenger ribonucleic acid in bovine endometrium: LH augmentation of cAMP and inositol phosphate in vitro and human chorionic gonadotropin (hCG) augmentation of peripheral prostaglandin in vivo [J]. Reprod Biol, 2001, 1:13-32.
[68] PIETILA E M, AATSINKI J T, Rajaniemi H.J.. Expression of the LH/CGreceptor gene in various rat tissues [J]. J Physiol Pharmacol, 1996, 47(1):5-14.
[69] ZHENG M, SHI H, SEGALOFF D L, et al. Expression and localization of luteinizing hormone receptor in the female mouse Reproductive tract [J]. Biol Reprod, 2001, 64:179-187.
[70] SHEMESH M. Actions of gonadotrophins on the uterus [J]. Reproduction, 2001, 121: 835-842.
[71] ZIECIK A J, OSTROWSKA G, KISIELEWSKA J, et al. Distribution and cycle phase dependency of gonadot ropin receptors in musculature and blood vessels of the porcine broad ligament [J]. Exp Clin Endocrinol Diabetes, 1995,103:44-51.
[72] SHEMESH M, GUREVICH M, FIELDS M J, et al. Expression of functional luteinizing hormone (LH) receptorand it s messenger ribonucleic acid in bovine uterine veins: LH induction of cyclooxygenase and augmentation of prostaglandin production in bovine uterine veins [J]. Endocrinology, 1997, 138:4844-4851.
[73] RODWAY M R, RAO C V. A novel prospective on t he role of human chorionic gonadot ropin during pregnancy and gestational trophoblastic disease. Early Pregnancy [J]. Biology and Medicine, 1995, 3:176-187.
[74] LEI Z M, RESHEF E, RAO V. The expression of human chorionic gonadotropin/ luteinizing hormone receptors in human endometrial and myometrial blood vessels [J]. J Clin Endocrinol Metab, 1992, 75(2):651-659.
[75] SHEMESH M., GUREVICH M., SHOREL S, et al. Regulation of cyclooxygenase and products of the cyclooxygenase pathway in the bovine uterus. Reprod Dom Anim., 1996,31 :431-436.
[76] STEPIEN A, SHEMESH M, ZIECIK A J. Luteinising hormone receptor kinetic and LH induced prostaglandin production throughout the oestrous cycle in porcine endometrium [J]. Reprod Nutr Dev,1999, 39:663-674.
[77] ZIECIK A J, BODEK G, BILTEK A. Nongonadal LH receptors, their involvement in femail reproductive function and a new applicable approach [J]. Vet J, 2005,169 (1):75-84.
[78] ZIECIK A J, STEPIEN A, GAWRONSKA B. Importance of endometrial luteinizing hormone receptors in induction of luteolysis and maternal recognition of pregnancy in the pig [J]. Reproduction in Domestic Animals, 2000, 35 :190-192.
[79] LIN P C, LI X, LEI Z M, et al. Human cervix contains functional luteinizing hormone/ human chorionic gonadot ropin receptors [J]. The Journal of Clinical Endocrinology & Metabolism, 2003, 88:3409-3414.
[80] STEPIEN A, DERECKA K, GAWRONSKA B, et al. LH/ hCG receptors in the porcine uterus a new evidence of their presence in the cervix and myometrium [J]. Physiol Pharmacol, 2000, 51:917-931.
[81]申颖,侯志高,王树迎.促性腺激素受体在子宫和输卵管中分布和作用的研究进展[J].动物医学进展. 2007, 28(6):86-88.
[82] LEI Z M, TOTH P, RAO Ch V, PRIDHAM D. Novel coexpression of human chorionic gonadotropin (hCG)/human luteinizing hormone receptors and their ligand hCG in human fallopian tubes [J]. J Clin Endocrinol Metab, 1993,77:863-872.
[83] HAN S W, LEI Z M, RAO C V. Up-regulation of cyclooxygenase-2 gene expression by chorionic gonadotropin in mucosal cells from human fallopian tubes [J]. Endocrinology, 1996, 137:2929-2937.
[84] SUN T, LEI Z M, RAO CH V. A novel regulation of the oviductal glycoprotein gene expression by luteinizing hormone in bovine tubal epithelial cells [J]. Mol Cell Endo, 1997, 131:97-108.
[85] GAWRONSKA B, PAUKKU T, HUHTANIEMI I, et al. Estrogen-dependent expression of LH/hCG receptors in pig fallopian tube and their role in relaxation of the oviduct [J]. J, Reprod Fertil, 1999, 115:293-301.
[86] DERECKA K, PIETILA E M, RAJANIEMI H J, et al. Cycle-dependent LH/hCG recaptor gene expression in porcine nongonadal reproductive tissues [J]. J Physiol Pharmacol, 1995, 46:77-85.
[87] GAWRONSKA B, STEPIEN A, ZIECIK A J. Effect of estradiol and progesterone on oviductal LH-receptors and LH-dependent relaxation of the porcine oviduct [J]. Theriogenology, 2000, 53:659-672.
[88] GAWRONSKA B, STEPIEN A, ZIECIK A J. Role of luteinizing hormone in control of oviduct function [J]. Reprod Domest Anim, 2000, 35:129-133.
[89] MISHRA S, LEI Z M, RAO Ch V, A novel role of luteinizing hormone in the embryodevelopment in cocultures [J]. Biol Reprod, 2003, 68:1455-1462.
[90] YOU S, KIM H, HSU C C, et al. Three different turkey luteinizing hormone receptor (tLH-R) isoforms I: characterization of alternative spliced tLH-R isoforms and their regulated expression in diverse tissues [J]. Biol Reprod, 2000, 62:108-116.
[91] FIELDS M J, SHEMESH M. Extragonadal luteinizing hormone receptors in the reproductive tract of domestic animals [J]. Bio Reprod, 2004, 71:1412-1418.
[92] SHI H, SEGALOFF D L. A role for increased lutropin/choriogonadotropin receptor (LHR) gene transcription in the follitropin-stimulated induction of the LHR in granulosa cells [J]. Mol Endocrinol, 1995, 9(6):734-744.
[93] IKEDA S, NAKAMURA K, KOGURE K., et al. Effect of estrogen on the expression of luteinizing hormone-human chorionic gonadotropin receptor messenger ribonucleic acid in cultured rat granulosa cells [J]. Endocrinology, 2008, 149(4):1524-1533.
[94] HIRAKAWA T, MINEGISHI T, ABE K, et al. A role of insulin-like growth factor I in luteinizing hormone receptor expression in granulosa cells [J]. Endocrinology, 1999, 140(11):4965-4971.
[95] ROY S K. Epidermal growth factor and transforming growth factor-beta modulation of follicle-stimulating hormone-induced deoxyribonucleic acid synthesis in hamster preantral and early antral follicles [J]. Biol Reprod, 1993 48(3):552-557.
[96]唐杰.山羊促性腺激素及其受体mRNA丰度的研究[D].河北:河北农业大学,2007.
[97] JOHANN S, HYESOOK Y, MYOUNGKUN J, et al. Orientation of Follicle-stimulati ng Hormone (FSH) Subunits Complexed with the FSH Receptor p subunit toward the N terminus of exodomain and a subunit to exoloop[J]. J Biol Chem, 2003, 278(48) :47868-47876.
[98] ANDREWS W V, MAURER R A, CONN P M. Stimulation of rat luteinizing hormon-ebeta messenger RNA levels by gonadotropin releasing hormone. Apparent role for protein kinaseC [J]. J Biol Chern, 1988, 263:13755-13761.
[99] BEN-MENAHEM D, NAOR Z. Regulation of gonadotropin mRNA levels in cultured rat pituitary cells by gonadotropin-releasing honnone(GnRH): role for Ca2+ and protein kinase C [J]. Biochemistry, 1994, 33:3698-3704.
[100] SUNDARESAN S, COLIN I M, PESTELL R Q et al. Stimulation of mitogenactivated protein kinase by gonadotropin-releasing hormone: evidence for the involvement of protein kinase [J]. Endocrinology, 1996, 137:304-311.
[101] MEUER K A, COX N M, MATAMOROS I A, et al. Decreased follicular steroids and insulin-like growth factor-I and increased atresia in diabetic gilts during follicular growth stimulated with PMSG [J]. J Reprod Fertil, 1991, 91(1):187-196.
[102] SHORT R E, ADAMS D C. Nutrition and hormonal interrelationships in beef cattle reproduction [J]. Anim Sci, 1998, 68:29-34.
[103] KLINDT J,YEN J T, Christenson R K, et al. Effect of prepubertal feeding regimen on reproduce-tive performance of gilts [J]. Anim Sci, 1999, 4:620-624.
[104] REESE D E, MOSER B D, PEO ER J R, et al. Influence of energy intake during lactation on the interval from weaning to first estrus in sows [J].Anim Sci, 1982, 55: 590-598.
[105] SHORT R E, BELLOWS R A. Relationships among weight gains, age at puberty and reproductive performance in heifers [J]. J Anim Sci, 1971,32, 127-131.
[106] GONZALEZ-PADILLA E, WILTBANK J N, et al. Puberty in beef heifers. I. The interrelationship between pituitary, hypothalamic and ovarian hormones [J]. J Anim Sci, 1975, 40:1091-1104.
[107] MURPHY M G, ENRIGHT W J, CROWE M A, et al. Effects of dietary intake on pattern of growth of dominant follicles during the oestrous cycle in beef heifers [J]. J Reprod Fertil, 1991, 92: 333-338.
[108] BEAM S W, BUTTER W R. Energy balance effects on follicular development and first ovulation in post-partum cows [J]. J Reprod Fertil Suppl, 1999, 54:411-424.
[109] BOOTH P J, COSGROVE J R, FOXCROFT G R. Endodrine and Metabolic Responses to Realimentation in Feed-Restricted Prepubertal Gilts: Associations Among Gonadotropins, Metabolic Hormones, Glucose, and Uteroovarian Development [J]. Anim Sci, 1996, 74:840-848.
[110] BOOTH P J, COSGROVE J R, FOXCROFT G R. Nutritional Manipulation of Growth andMetabolic and Reproductive Status in Prepubertal Gilts [J]. J Anim Sci, 1994, 72:2415-2424.
[111] ARMSTRONG J D, BRITT J H. Nutritionally-induced anestrus in gilts: metabolic and endocrine changes associated with cessation and resumption of estrous cycles [J]. J Anim Sci, 1987, 65(2):508-523.
[112] ZAK L J, COSGROVE J R, AHERNE F X, et al. Pattern of feed intake and associated metabolic and endocrine changes differentially affect postweaning fertility in primiparous lactating sows [J]. J Anim Sci, 1997, 75(1):208-216.
[113] VANDEN Brand H, DIELEMAN S J, SOEDE N M, et al. Dietary energy source at two feeding levels during lactation of primiparous sows: I. Effects on glucose, insulin, and luteinizing hormone and on follicle development, weaning-to-estrus interval, and ovulation rate [J]. J Anim Sci, 2000, 78:396-404.
[114] ZAK L J, XU X, HARDIN R T, et al. Impact of different patterns of feed intake during lactation in the primiparous sow on follicular development and oocyte maturation [J]. J Reprod Fertil, 1997, 110(1):99-106.
[115] ALMIDAl F R, KIRKWOOD R N, AHERNE F X, et al. Consequences of different patterns of feed intake during the estrous cycle in gilts on subsequent fertility [J]. J Anim Sci, 2000, 78(6):1556-1563.
[116] QUESNEL H, PASQUIER A, MOUNIER A M, et al. Influence of Feed Restriction During Lactation on Gonadotropic Hormones and Ovarian Development in Primiparous Sows [J]. J Anmi Sci, 1998,76:856-863.
[117] BRITT J H, ARMSTRONG J D, COX N M, et al. Control of follicular development during and after lactation in sows [J]. J Reprod Fertil Suppl, 1985, 33:37-54.
[118] MAURASSE C., MARTON P, DUFOUR J J, et al. Ovarian follicular populations at two stages of anoestrus cycle in heifers given high energy diets [J]. J anim Sci, 1985, 61: 1194-1200.
[119] GASSER C L, BEHLKE E J, GRUM D E, et al. Effect of timing of feeding a high-concentrate diet on growth and attainment of puberty in early-weaned heifers [J]. J Anim Sci, 2006, 84(11):3118-3122.
[120] MACKEY D R, SREENAN J M, ROCHE J F, et al. Effect of acute nutritional restriction on incidence ofanovulation and periovulatory estradiol and gonadotropin concentrations in beef heifers [J]. Biol Reprod, 1999, 61:1601-1607.
[121] KENDRICK K W, BAILEY T L, GARST A S, et al. Effects of energy balance of hormones, ovarian activity, and recovered oocytes in lactating Holstein cows using transvaginal follicular aspiration [J]. Dairy Sci, 1999, 82:1731-1741.
[122] JORRITSMA R, GROOT M W, VOS P L, et al. Acute fasting in heifers as a model for assessing the relationship between plasma and follicular fluid NEFA concentrations [J]. Theriogenology, 2003, 60(1):151-161.
[123]李勇,吴德.能量对家畜卵泡发育的影响及调控[J].饲料工业, 2007, 28(21):15-19.
[124]唐晓龙.不同能量和蛋白水平日粮对母羊繁殖性能的影响[D].河北:河北农业大学, 2007.
[125] KILLEEN ID. Effects of fasting ewes before mating on their reproductive performance [J]. Theriogenology, 1982,17:433-435.
[126] KIYMA Z, ALEXANDERl B M, VAN KIRK E A. Effects of feed restriction on reproductive and metabolichormones in ewes [J]. Anim Sci, 2004, 82:2548-2557.
[127]刘立文.日粮能量对初情期前母猪FSH、LH、GH、胰岛素及IGF-I释放的影响[D].吉林:吉林大学, 2006.
[128]于淼瑛.不同日粮能量水平对初情期前母猪卵巢及子宫LH受体和FSH受体mRNA表达的影响[D].吉林:吉林大学, 2006.
[129] COX N M, STUART M J, ALTHEN T G, et al. Enhancement of ovulation rate in gilts by increasing dietary energy and administering insulin during follicular growth [J]. J Anim Sci, 1987, 64(2):507-516.
[130] BOOTH P J, COSGROVE J R, FOXCROFT G R. Endocrine and metabolic responses to realimentation in feed-restricted prepubertal gilts: associations among gonadotropins, metabolic hormones, glucose, and uteroovarian development [J]. J Anim Sci, 1996, 74(4): 840-848.
[131] MATAMOROS I A, COX N M, MOORE A B. Effects of exogenous insulin and body condition on metabolic hormones and gonadotropin-induced follicular development in prepuberal gilts [J]. J Anim Sci, 1991, 69(5):2081-2091.
[132]周平,吴德,王延忠,等.营养对母猪卵泡发育的影响[J].饲料博览. 2007, 15:25-27.
[133]于淼瑛,周虚.胰岛素在能量影响猪卵泡发育中的作用.黑龙江动物繁殖[J]. 2005, 13(2):12-15.
[134] HELEN Q, ANNE P, ARMELLE P, et al. Feed restriction in cyclic gilts: Gonadotrophin independent effects on follicular growth[J]. Reprod Nutr Dev, 2000, 40(4): 405-414.
[135] DRIANCOURT M A, LOCATELLI A, PRUNIER A. Effects of gonadotrophin deprivation on follicular growth in gilts [J]. Reprod Nutr Dev, 1995, 35(6):663-673.
[136] ALMEIDA F R, MAO J, NOVAK S, et al. Effects of different patterns offeed restriction and insulin treatmen tduring the luteal phase on reproductive, metabolic, and endocrine parameters in cyclic gilts [J]. J Anim Sci, 2001,79 (1) :200-212.
[137] MAY J V, SCHOMBERG D W. Granulosa cell differentiation in vitro: effect of insulin on growth and functional integrity [J]. Biol Reprod, 1981,25(2):421-431.
[138]李波.能量对初情期前母猪卵巢和子宫IGF-ⅠR和EGFR mRNA表达的影响[D].吉林:吉林大学, 2007.
[139] ADAM C L, GADD T S, FINDLAY P A, et al. IGF-1 stimulation of luteinizing hormone secretion, IGF-binding proteins (IGFBPs) and expression of mRNAs for IGFs, IGF receptors and IGFBPs in the ovine pituitary gland [J]. J Endocri, 2000, 166: 247-254.
[140] RIVERA G M, FORTUNE J E. Proteolysis of insulin-like growth factor binding proteins-4 and -5 in bovine follicular fluid :implications for ovarian follicular selection and dominance [J]. Endocrinology, 2003, 144 (7) :2977-2987.
[141] LEEUWENBERG B R, HURST P R, MCNATTY K P. Expression of IGF-I mRNA in the ovine ovary [J]. J Mol Endocrinol, 1995, 15(3):251-258.
[142] DEVOTO L, KOHEN P, CASTRO O, et al. Multihormonal regulation of progesterone synthesis in cultured human midluteal cells [J]. J Clin Endocrinol Metab, 1995, 80(5):1566-1570.
[143] FOSTER C M, HASHIMOTO N, RANDOLPH J F, et al. Comparison of growth hormone and insulin-like growth factor-I regulation of estradiol and progesterone production in human luteinized granulosa cells [J]. Pediatr Res, 1995, 38(5):763-767.
[144]刘丽,李琦华,贾俊静,等.瘦素对动物能量代谢及繁殖性能的影响[J].饲料研究, 2008, 6, 26-28.
[145] BARB C R, HAUSMAN G J, CZAJA K, et al. Leptin: a Metabolic Signal Affecting Central Regulation of Reproduction in the Pig [J]. Domest Anim Endocrinol, 2005, 29(1):186-192.
[146] RUIZ-CORTE′S Z T, MEN T, PALIN M F, et al. Porcine leptin receptor: molecular structure and expression in the ovary [J]. Mol Reprod Dev, 2000, 56:465-474.
[147] SPICER L J, CHAMBERLAIN C S, FRANCISCO C C, et al. Ovarian action of leptin: effects on insulin-like growth factor-I-stimulated function of granulosa and thecal cells [J].Endocrine, 2000, 12(1):53-59.
[148] Guo X, Chen S, Xing F, et al. Effects of leptin on estradiol and progesterone production by human luteinized granulosa cells in vitro [J]. Zhonghua Fu Chan Ke Za Zhi. 2001, 36(2): 95-97.
[149] GHIZZONI L, BARRECA A, MASTORAKOS G, et al. Leptin inhibits steroid biosynthesis by human granulosa-lutein cells [J]. Horm Metab Res, 2001, 33(6):323-328.
[150] TATIANA RUIZ-CORTES Z, MARTEL-KENNES Y, NICOLAS Y, et al. Biphasic Effects of Leptin in Porcine Granulosa Cells [J]. Biol Reprod, 2003, 68:789-796.
[151] GONG J G, BRAMLEY T, WEBB R. The effect of recombinant bovine somatotropinon ovarian function in heifers. follicular populations and peripheral hormones [J]. BiolReprod, 1991, 45: 941-949.
[152] HERRLER A, EINSPANIER R, SCHAMS D, et al. Effect of recombinant bovine somatotropin (rBST) on follicular IGF-1contents and the ovarian response following superovulatrory treatment in dairy cows . Apreliminary study [J]. Theriogenology, 1994, 41:601-611.
[153] GONG J G, BAXTER G, BRAMLEY T A, et al. Enhancement of ovarian follicle development in heifers by treatment with recombinant bovine somatotropin. adose-response study [J]. J Reprod Fertil, 1997, 110:91-97.
[154] ZIECIK A J, DERECKA-RESZKA K, RZUCIDLO S J. Extragonadal gonadotropin receptors, their distribution and function [J]. J Physiol Pharmacol, 1992, 43:33-49.
[155] FRIEDMAN S, GUREVICH M, SHEMESH M. Bovine cyclic endometrium contains high-affinity luteinizing hormone/human chorionic gonadotropin binding sites [J]. Biol Reprod, 1995, 92:1020-1026.
[156] ZIECIK A J, STANCHEV P D, TILTON J E. Evidence for the presence of luteinizing hormone/human chorionic gonadotropin-binding sites in the porcine uterus[J]. Endocrinology, 1986, 119:1159-1163.
[157] RZUCIDLO S J, WEIGL R M, TILTON J E. Myometrial LH/hCG receptors during the estrous cycle and pregnancy in pigs [J]. Anim Reprod Sci, 1998, 51:249-257.
[158] BONNAMY P J, BENHAIM A, LEYMARIE P. Human chorionic gonadotropin affects tissue levels of progesterone and cyclic adenosine 39,59-monophosphate in the metestrous rat uterus in vitro [J]. Biol Reprod, 1989, 40:511-516.
[159] SAWITZKE A L, ODELL W D. Uterine binding sites for LH/hCG can be modulated byhormonal status in rabbits and rats [J]. Acta Endocrinol, 1991, 124:322-330.
[160] JENSEN T D, ODELL W D. Identification of LH/hCG receptors in rabbit uterus [J]. Proc Soc Exp Biol Med, 1988, 189:28-30.
[161] RESHEF E, LEI Z M, RAO CH V, et al. The presence of gonadotropin receptors in nonpregnant human uterus, human placenta, fetal membranes, and decidua [J]. J Clin Endocrinol Metab, 1990, 70:421-430.
[162] SHEMESH M, Hansel W. Levels of prostaglandin F(PGF) in bovine endometrium , uterine venous , ovarian arterial and jugular plasma during the estrous cycle [J]. Proc Soc Exp Biol Med, 1975, 148 :123-126.
[163] TOTH P, LI X, RAO C V, et al. Expression of functional human chorionic gonadotropin/ humanluteinizing hormone receptor gene in human uterine arteries [J]. J Clin Endocrinol Metab, 1994, 79:307-315.
[164] LIN D X , LEI Z M. Dependence of Uterine Cyclooxygenase Expression on Luteinizing Hormone Signaling [J]. Biol Reprod, 2005, 73:256-260.
[165] KORNYEI J L, LI X, LEI Z M, et al. Restoration of human chorionic gonadotropin response in human myometrial smooth muscle cells by treatment with follicle-stimulating hormone (FSH): evidence for the presence of FSH receptors in human myometrium [J]. European Journal of Endocrinology, 1996, 134(2): 225-231.
[166] GAWRONSKA B, STEPIEN A , ZIECIK A J . Effect of estradiol and progesterone on oviduc-tal LH-receptors and LH-dependent relaxation of the porcine oviduct[J]. Theriogenology, 2000, 53:659-672.
[167] RZUCIDLO S J, WIGL R M, TILTON J E. Myometrial LH/hCG receptors during the estrous cycle and pregnancy in pigs [J]. Anim Reprod Sci, 1998, 51:249-257.
[168] ZIECIK A J, KOTWICA G. Involvement of gonadotropins in induction of luteolysis in pigs [J]. Reprod Biol, 2001, 1:33-50.
[169] WEEMS Y S, KIM L, HUMPHREYS V, et al. Effect of luteinizing hormone (LH), pregnancy specific protein B (PSPB), or arachidonic acid (AA) on ovine endometrium of the estrous cycle or placental secretion of prostaglandins E2 (PGE2) and F2α(PGF2α) and progesterone in vitro [J]. Prostaglandins, 2003, 71:55-73.
[170] ZHOU X L, LEI Z M, RAO CH V. Treatment of human endometrial gland epithelial cells with chorionic gonadotropin/luteinizing hormone increases the expression of thecyclooxygenase-2 gene [J]. J Clin Endocrinol Metab, 1999, 84:3364-3377.
[171] COSGROVE J R, TILTON J E, HUNTER M G, et al. Gonadotrogen-independent mechanisms in feed–restricted prepubertal gilts [J]. Bio Reprod, 1992, 47: 736-745.
[172] COSGOVE J R, TITON T G, ALTHEN W A, et al. Maturational changes in gonadonatrophin secretion the LH response to realimentation and a nocturnal in crement in LH secrection of feed-restricted prepubertal gilts [J]. Reprod. Fertil, 1993, 98: 293-305.
[173] BLANK J L, DESJARDINS C. Differential effects of food restriction on pituitary- testicular function in mice [J]. Am J Physiol, 1985, 248: 181-189.
[174] FOSTER D L, OLSTER D H. Effect of restricted nutrition on puberty in the lamb: patterns of tonic luteinizing hormone (LH) secretion and competency of the LH surge system [J]. Endocrinology, 1985, 116:375-381.
[175] THOMAS G B, MERCER J E, KARALIS T, et al. Effect of restricted feeding on the concentrations of growth hormone (GH), gonadotropins, and prolactin (PRL) in plasma, and on the amounts of messenger ribonucleic acid for GH, gonadotropin subunits,and PRL in the pituitary glands of adult ovariectomized ewes [J]. Endocrinology, 1990, 126:1361-1367.
[176] CAMERON J L, NOSBISCH C. Suppression of pulsatile luteinizing hormone and testosterone secretion during short term food restriction in the adult male rhesus monkey (Macaca mulatta) [J]. Endocrinology, 1991, 128:1532-1540.
[177] ROJDMARK S. Influence of short-term fasting on the pituitary-testicular axis in normal men [J]. Horm Res, 1987, 25:140-146.
[178] RHODES M T, MINION J E, STEVENSON J S, et al. Endocrine changes associated with flushing and altrenogest treatment in gilts [J]. Anim. Sci, 1987, 65(1):154-161.
[179] FLOWERSl B, MARTIN M J, CANTLEY T C. et al. Endocrine Changes Associated with a Dietary-Induced Increase in Ovulation Rate (Flushing) in Gilts [J]. J Anim Sci, 1989, 67:771-778.
[180] COSGROVE J R, TILTON J E, HUNTER M G, et al. Gonadotrogen-independent mechanisms in feed–restricted prepubertal gilts [J]. Bio Reprod, 1992, 47: 736-745.
[181] SNYDER J L, CLAPPERl J A, ROBERTS A J, et al. Insulin-like growth factor-I, insulin-like growth factor-binding proteins, and gonadotropins in the hypothalamicpituitary axis and serum of nutrient restricted ewes [J]. Biol. Reprod,1999, 61: 219-224.
[182] KILE J P, ALEXANDEl B M, MOSS G E, et al. Gonadotropin-releasing hormone overridesthe negative effect of reduced dietary energy on gonadotropin synthesis and secretion in ewes [J]. Endocrinology,1991, 128:843-849.
[183] RHIND S M, LESLIE L D, GUNN R G, et al. Plasma FSH, LH, prolactin and progesterone profiles of Cheviot ewes with different levels of intake before and after mating, and associated effects on reproductive performance [J]. Anim Reprod Sci, 1985, 8:301-313.
[184] GOMBE S, HANSEL W. Plasma luteinizing hormone (LH) and progesterone levels in heifers on restricted energy intakes [J].Anim Sci, 1973, 37:728-733.
[185] MCCANN J P, HANSEL W. Relationships between insulin and glucose metabolism and pituitary-ovarian functions in fasted heifers [J]. Biol. Reprod, 1986, 34:630-641.
[186] SCHRICK F N, SPITZER J C, JENKINS T C, et al. Effect of dietary energy restriction on metabolic and endocrine responses during the estrous cycle of the suckled beef cow [J]. Anim Sci, 1990, 68:3313-3321.
[187] COOPER K, BROOKSR P H, COLE D J A, et al. The effect of feed level during the estrous cycle on ovulation, embryo survival and anterior pituitary LH potency in the gilt [J]. J Reprod Fertil, 1973, 32:71-78.
[188] KIRKPATRICK, R L, HOWLAND B E, FIRST N L, et al. Ovarian and pituitary gland changes in gilts on two nutrient energy levels [J]. J Anita Sci, 1967, 26:358-364.
[189] KEMP B, SOEDE N M, HELMOND F A, et al. Effects of energy source in the diet on reproductive hormones and insulin during lactation and subsequent estrus in multiparous sows [J]. Anim Sci, 1995, 73:3022-3029.
[190] KOKETSU Y, DIAL G D, PELLIGREW J E, et al. Influence of imposed feed intake patterns during lactation on reproductive performance and on circulating levels of glucose, insulin, and luteinizing hormone in primiparous sows [J]. Anim Sci, 1996, 74:1036-1046.
[191] BAIDOO S K, LYTHGOEy E S, KIRKWOOD R N, et al. Effect of lactation feed intake on endocrine status and metabolite levels in sows [J]. Anim Sci, 1992, 72:799-807.
[192] DOMINGUEZ M M. Effects of body condition, reproductive status and breed on follicular population and oocyte quality in cows [J].Theriogenology, 1995, 43:1405-1418.
[193] GHARIB S D, WIERMAN M E, SHUPNIK M A, et al. Molecular biology of the pituitary Gonadotropins [J]. Endocr Rev, 1990, 11:177-199.
[194] MARSHALL JC M, CHRISTINE A E, CHRISTOPER R M. Hypothalamic dysfunction [J]. Mol Cell Endocrinol, 2001, 183: 29-32.
[195] NETT T M, TURZILLO A M, BARATTA M, et al. Pituitary effects of steroid hormones on secretion of follicle-stimulating hormone and luteinizing hormone [J]. Dourest Anim Endocrinol, 2002, 23: 33-42.
[196] WADE G N, SCHNEIDER J E. Metabolic fuels and reproduction in female animal [J]. Neurosci, Biobehav, 1992, 16: 235-272.
[197] MONGET P, MARTIN G B. Involvement of insulin-like growth factor in the interactions between nutrition and reproduction in female mammals [J]. Human Reprod, (Oxf.), 1997, 12: 33-52.
[198] MOUNIZH K, LU R , CHEHAB F F. Leptin treatmeant rescues the terility of genetically ob/ob males [J]. Endocrinology, 1997, 138: 1190-1193.
[199] VIGUIE C, BATTAGLIA D F, KRASA H A, et al. Thyroid hormones act primarily within the brain to promote the seasonal inhibition of luteinizing hormone secretion in the ewe [J]. Endocrinology, 1999, 140: 1111-1117.
[200] LANDAU, B R, TAKAOKA Y, ABRAMS M A, et al. Binding of insulin by monkey and pig hypothalamus [J]. Diabetes, 1983, 32:284-291.
[201] HAVRANKOVA J, SCHMECHEL D, ROTH J, et al. Identification of insulin in rat brain [J]. Proc Natl Acad Sci, USA 1978, 75:5737-5742.
[202] ADASHI E Y HSUEHA J W, YENS S C. Insulin enhancement of luteinizing hormone and follicle-stimulating hormone release by cultured pituitary cell [J]. Endocrinology, 1981, 108:1441-1449.
[203] ARMSTRONG D G, MCEVOY T G, BAXTER G J. et al. Effect of dietary energy and protein on bovine follicular dynamics and embryo production in vitro: associations with the ovarian insulin-like growth factor system [J]. Biol Reprod, 2001, 64:1624-1632.
[204] BOSSIS I R. WETTEMANN P, WELTY S D J. et al. Nutritionally induced anovulation in beef heifers: ovarian and endocrine function during realimentation and resumptions of ovulation [J]. Biol Reprod, 2000, 62:1436-1444.
[205] GRIGSBY M E, TRENKLE A. Plasma growth hormone, insulin, glucocorticoids and thyroid hormones in large, medium and small breeds of steers with and without an estradiol implant [J]. Domest Anim Endocrinol, 1986, 3:261-267.
[206] RICHARDS M W, WELLEMAN R P, SPICER L J, et al. Nutritional anestrus in beef cows: Effect of body condition and ovariectomy on serum luteinizing hormone and insulin-likegrowth factor-I [J]. Biol Reprod, 1991, 44:961-966.
[207] SIMPSON R B, CHASE C C, SPICER L J, et al. Effect of exogenous estradiol on plasma concentrations of somatotropin, insulin-like growth factor-I, insulin- like growth factor binding protein activity, and metabolites in ovariectomized Angus and Brahman cows [J]. Domest Anim Endocrinol, 1997, 14:367-380.
[208] ARMSTRONG, D G, GONG J G, WEBB R. Interactions between nutrition and ovarian activity in cattle: Physiological, cellular and molecular mechanisms [J]. Repro duction Suppl, 2003, 61:403-414.
[209] ZAMORANO P L, DE SEVILLA L, MAHESH V B, et al. Production and refolding of recombinant leptin [J]. Biotechniques, 1997, 23(5):800-804.
[210] LONG J, BARB C R, KRAELELING R R, et al. Growth hormone releaseing factor decrease long form Leptin receptor expression in porcine anterior pituitary cells [J]. Domestic Anim Endocrinology, 2003, 141:333-339.
[211] SANJAY K, AGARW A L, KLARA L, et al. Leptin Antagonizes the Insulin-Like Growth Factor-I Augmentation of Steroidogenesis in Granulosa and Theca Cells of the Human Ovary [J].Clin Endocrinol Metab, 1999, 84: 1072-1076.
[212] WISHANNT C S, HARELL R J. Leptin and luteinnizing homone concentration in pigs. College of agriculture and life sciences department of animal science annual swine report, 2001. 27:367-378.
[213] AMSTALDEN M, GARCIA M R, WILLIAMS S W, et al. Leptin gene expression, circulating leptin, and luteinizing hormone pulsatility are acutely responsive to short-term fasting in prepubertal heifers: Relationships to circulating insulin and insulin-like growth factor-I [J]. Biol Reprod, 2000, 63:127-133.
[214] BRYAN K A. Effect of doses of porcine somatotropin (pST) on growth, carcass traits and granulosa cell function of gilts [J]. Anim Sci, 1988, 66 (1): 379-385.
[215] BRYAN K A, HAMMOND J M, CANNING S, et al. Reproductive and growth responses of gilts to exogenous porcine pituitary omatotropin [J]. Anim Sci, 1989, 67: 196-205.
[216] ADASHI, E Y, HSUEH AJ W, Yen S S C. Insulin enhancement of luteinizing hormone and follicle-stimulating hormone release by cultured pituitary cell [J]. Endocrinology, 1981, 108:1441-1449.
[217] WANDJI S A, WOOD T L, CRAWFORD J, et al . Expression of mouse ovarian insulin-likegrowth factor system components during follicular development and atresia [J]. Endocrinol, 1998, 139 (12): 5205-5214.
[218]葛立军,芮荣.促黄体素受体在母畜生殖道的分布与表达[J].动物医学进展, 2007, 28 (11) :92-95.
[219] MINEGISHI T, HIRAKAWA T, ABE K. A Role of Insulin-Like Growth Factor I in Luteinizing Hormone Receptor Expression in Granulosa Cells [J]. Endocrinology, 1999, 11: 4965-4971.
[220] ZHOU J, KUMAR T R, MATZUK M M, et al. Insulin-like growth factor I regulates gonadotropin responsiveness in the murine ovary [J]. Mol Endocrinol, 1997, 11:1924-1933.
[221] BAKER J, HARDY M P, ZHOU J, et al. Effects of an Igf1 gene null mutation on mouse reproduction [J]. Mol Endocrinol, 1996, 10:903-918.
[222] TILLY J L, LAPOLT P S, HSUECH J W. Hormonal regulation of follicle stimulating hormone receptor messenger ribonucleic acid level in cultured rat granulose cells [J]. Endocrinology, 1992, 130(3) : 1296-1302.
[2]李俊杰,贾青,田树军.雌性哺乳动物LH及其受体的研究进展[J].黄牛杂志,2003,29(3):26-29.
[3]李俊杰,贾青,张正珊.雌性哺乳动物促卵泡素受体与促黄体素受体的结构与表达[J]..生命的化学,2003,23(4):289-291.
[4] SPRENGEL R, BRAUN T, NIKOLICS K, et al. The testicular receptor for follicle stimulating hormone: structure and functional expression of cloned Cdna [J]. Mol Endocrinol, 1990, 4(4):525-530.
[5] GUDERMANN T, NURNBERG B, SCHULTZ G. Receptors and G proteins as primary components of transmembrane signal transduction. Part 1. G-protein-coupled receptors: structure and function [J]. J Mol Med, 1995, 73(2):51-63.
[6] KENNELLY P J, KERBS E G.. Consensus sequences as substrste specificity determinants for protein kinases and protain phosphatases [J]. J Biol Chem, 1991, 266:15555-15558.
[7] QUINTANA J, HIPKIN RW, SANCHEZ-YAGUE J, et al. Follitropin (FSH) and a phorbol ester stimulate the phosphorylation of the FSH receptor in intact cells [J]. J Biol Chem, 1994, 269:8772-8779.
[8] MINEGISHI T, IGARASHI S, NAKAMURA K, et al. Functional expression of the recombinant human FSH receptor [J]. J Endocrinol, 1994, 141(2):369-375.
[9] VANNIER B, LOOSFELT H, MEDYRI G, et al. Anti-human FSH receptor monoclonal antibodies: immunochemical and immunocytochemical characterization of the receptor [J]. Biochemistry, 1996, 35(5):1358-1366.
[10] GROMOLL J, RIED T, HOLTGREVE-GREZ H, et al. Localization of the human FSH receptor to chromosome 2 p21 using a genomic probe comprising exon 10 [J]. J Mol Endocrinol, 1994, 12:265-271.
[11] ROUSSEAU-MERCK M F, MISRAHI M, LOOSFELT H, et al. Localization of the human luteinizing hormone receptor gene to chromosome 2 p21 [J]. Cytogenet Cell Genet, 1990, 54: 77-79.
[12] ROUEEEAU-MERCK M F, MISRAHI M, LOOSFELT H, et al. The chromosomal localization of the human follicle-stimulating hormone receptor gene on 2p21-p16 is similar to that of the luteinizing hormone receptor gene [J]. Genomics, 1993,15: 222-224.
[13] HECKERT L. L., GRISWOLD M. D. Expression of follicle-stimulating hormone receptor mRNA in rat testes and Sertolicells [J]. Molec Endocr, 1991, 5:670-677.
[14] HECKERT L L, GRISWOLD M D. Expression of the FSH receptor in the testis [J]. Rec.Prog. Horm. Res, 1992, 48: 61-78.
[15] LAPOLT P S, TILLY J L, AIHARA T, et al. Gonadotropin-induced up and down regulation of ovarian follicle stimulating hormone (FSH) receptor gene expression in immature rats: effects of pregnant mare's serum gonadotropin, human chorionic gonadotropin and recombinant FSH [J]. Endocrinology, 1992, 130: 1289-1295.
[16] CAMP T A, RAHAL J O. MAYO K E, Cellular localization and hormonal regulation of follicle stimulating hormone and luteinizing hormone receptor messenger RNAs in the rat ovary [J]. Molec Endocr, 1991, 5: 1405-1417.
[17] AITTOMAKI K, LUCENA J L, PAKARINEN P, et al. Mutation in the follicle stimulati- ng hormone receptor gene causes hereditary hypergonadotropic ovarian failure [J]. Cell, 1995, 82(6):959-968.
[18] MEANS A R, VAITUKAITIS J. Peptide hormone“receptor”: specific binding of 3 H-FSH to testis [J]. Endocrinology, 1972, 90(1):39-46.
[19] RANNIKKO A, PENTTILA T L, ZHANG F P, et al. Stage-specific expression of the FSH receptor gene in the prepubertal and adult rat seminiferous epithelium [J]. J Endocrinol. 1996, 151(1):29-35.
[20] ORTH J, CHRISTENSEN A K. Autoradiographic localization of specifically bound 125I-labeled follicle-stimulating hormone on spermatogonia of the rat testis [J]. Endocrinology. 1978, 103(5): 1944-1951.
[21] RANNIKKI A S, ZHANG F P, HUHTANIEMI I T. Ontogeny of follicle-stimulating hormone receptor gene expression in the rat testis and ovary [J]. Mol Cell Endocrinol, 1995, 107(2): 199-208.
[22] SASAKI M,YAMAMOTO M,ARISHIMA K,et al.Effects of follicle stimulating hormone on intermediate filaments and cell division of Sertoli cellsof fetal rat testis in culture [J].J Vet Med Sci, 1998, 60 (1):35-39.
[23]陈雪雁,陈克铨,陈实平. FSH受体的特性和表达的调控[J].解剖学报. 1999, 30(3):284-287.
[24]丁允闵,叶世钧,陈咏梅,等.大鼠支持细胞生后发育过程中卵泡刺激素受体mRNA水平的动态观察[J].中国医学科学院学报, 1996, 18(2):99-103.
[25] SOKKA T, HUHTANIEMI I. Ontogeny of gonadotrophin receptors and gonadotrophin stimulated cyclic AMP production in the neonatal rat ovary [J]. J Endocrinol, 1990, 127(2):297-303.
[26] O'SHAUGHNESSY P J, MARSH P, DUDLEY K. Follicle-stimulating hormone receptor mRNA in the mouse ovary during post-natal development in the normal mouse and in the adult hypogonadal (hpg) mouse: structure of alternate transcripts [J]. Mol Cell Endocrinol, 1994, 101(1-2):197-201.
[27] O'SHAUGHNESSY P J, DUDLEY K, RAJAPAKSHA W R. Expression of follicle stimulating hormone-receptor mRNA during gonadal development [J]. Mol Cell Endocrinol, 1996, 125(1-2):169-175.
[28] MARGRET W Z, MAGID M S, KRAMER E E, et al. Follicle-stimulating hormone receptor is expressed in human ovarian surface epithelium and fallopian tube. Am J Pathol, 1996, 148(1):47-53.
[29] DUNKEL L, TILLY J L, SHIKONE T, et al. Follicle-stimulating hormone receptor expression in the rat ovary: increases during prepubertal development and regulation by the opposing actions of transforming growth factors beta and alpha [J]. Biol Reprod, 1994, 50(4):940-948.
[30] MINEGISHI T, TANO M, IGARASHI M, et al. Expression of follicle-stimulating hormone receptor in human ovary [J]. Eur J Clin Invest, 1997, 27(6):469-474.
[31] ZHANG C q, SHIMADA K, SAITO N. Expression of Messenger Ribonucleic Acids of Luteinizing Hormone and Follicle-Stimulating Hormone Receptors in Granulosa and Theca Layers of Chicken Preovulatory Follicles [J]. 1997, 105: 402-409.
[32] MIZARACHI D, SHEMESH M. Follicle-Stimulating Hormone Receptor and Its Messenger Ribonucleic Acid Are Present in the Bovine Cervix and Can Regulate Cervical Prostanoid Synthesis [J]. Biology of reproduction, 1999, 61:776-784.
[33] SHEMESH M, MIZRACHI D, GUREVICH M, et al. Functional importance of bovine myometrial and vascular LH receptors and cervical FSH receptors [J]. Semin Reprod Med,2001, 19(1):87-96.
[34] DIAS J A, LINDAU-SHEPARD B, HAUER C, et al. Human follicle-stimulating hormone structure- activity relationships [J]. Biol Reprod 1998, 58:1331-1336.
[35] KORNYEI J L, LI X, LEI Z M, et al. Restoration of human chorionic gonadotropin response in human myometrial smooth muscle cells by treatment with follicle-stimulating hormone (FSH): evidence for the presence of FSH receptors in human myometrium [J]. European Journal of Endocrinology, 1996, 134 (2) :225-231.
[36] GALWAY A B, LAPOLT P S, TSAFRIRI A, et al. Recombinant follicle-stimulating hormone induces ovulation and tissue plasminogen activator expression in hypophysectomized rats [J]. Endocrinology, 1990, 127(6):3023-3028.
[37] LAPOLT P S, TILLY J L, AIHARA T, et al. Gonadotropin-induced up- and down-regulation of ovarian follicle-stimulating hormone (FSH) receptor gene expression in immature rats: effects of pregnant mare's serum gonadotropin, human chorionic gonadotropin, and recombinant FSH[J]. Endocrinology, 1992, 130(3):1289-1295.
[38] TANO M, MINEGISHI T, KISHI H, et al. The effect of follicle-stimulating hormone (FSH) on the expression of FSH receptor in cultured rat granulosa cells [J]. Life Sci, 1999, 64(12):1063-1069.
[39] MINEGISHI T, TANO M, KISJI H, et al. Follicle-stimulating hormone regulation on its receptor messenger ribonucleic acid levels in cultured rat granulosa cells [J]. Biochim Biophys Acta, 1997, 1359(2):165-173.
[40] MURPHY B D, Dobias M. Homologous and heterologous ligands downregulate follicle-stimulating hormone receptor mRNA in porcine granulosa cells [J]. Mol Reprod Dev, 1999, 53(2):198-207.
[41] MINEGISHI T, TANO M, NAKAMURA K, et al. Regulation of follicle-stimulating hormone receptor [J]. Horm Res., 1996;46(1):37-44.
[42] O'SHAUGHNESSY P J, GRAY S A. Gonadotropin-dependent and gonadotropin independent development of inhibin subunit messenger ribonucleic acid levels in the mouse ovary [J]. Endocrinology, 1995, 136(5):2060-2965.
[43] KUMAR T R, WANG Y. LU N, et al.. Follicle stimulating hormone is required for ovarian follicle maturation but not male fertility [J]. Nat.Genet. 1997, 15(2):201-204.
[44] RICHARDS J S. Maturation of ovarian follicles: actions and interactions of pituitary and ovarianhormones on follicular cell development [J]. Recent Prog Horm Res, 1980, 32: 477-527.
[45] REILLY C M, CANNADY W E, MAHESH V B, et al. Duration of estrogen exposure prior to follicle-stimulating hormone stimulation is critical to granulosa cell growth and differentiation in rats [J]. Biology of reproduction, 1996, 54:1336-1342.
[46] WEIL S, VENDOLA K, ZHOU J, et al. Androgen and follicle-stimulating hormone interactions in primate ovarian follicle development. J. Clin Endocrinol Metab, 1999, 84(8):2951-2956.
[47] CARDENAS H, HERRICK J R, POPE W F, Increased ovulation rate in gilts treated with dihydrotestosterone [J]. Reproduction, 2002, 123(4):527-533.
[48] MINEGISHI T, Hirakawa T, KISHI H. et al. A role of insulin-like growth factor I for follicle-stimulating hormone receptor expression in rat granulosa cells [J]. Biol Reprod, 2000, 62(2):325-333.
[49] OTSUKA F, YAMAMOTO S, ERICKSON G F, et al. Bone morphogenetic protein-15 inhibits follicle-stimulating homone (FSH) action by suppressing FSH receptor expression [J]. J Biol Chem, 2001, 276(14):11387-11392.
[50] TILLY J L, LAPOLT P S, HSUEH A J. Hormonal regulation of follicle-stimulating hormone receptor messenger ribonucleic acid levels in cultured rat granulosa cells [J]. Endocrinology, 1992, 130(3):1296-1302.
[51] FINDLAY, J K. An update on the roles of inhibin, activin and follistatin as local regulators of folliculogenesis [J]. Biol Reprod, 1993, 48:15-23.
[52] MENON K M J, MUNSHI U M, C louser C L, et al. Regulation of Luteinizing Hormone/ Human Chorionic Gonadotropin Receptor Expression: A Perspective1 [J]. Biol Reprod, 2004, 70: 861-866.
[53] MCFARLAND K C, SPRENGEL R, PHILLIPS H S, et al. Lutropin-choriogonadotropin receptor: an unusual member of the G protein-coupled receptpr family [J]. Science, 1989, 245(4917): 494-499.
[54] KISHI M, LIU X, HIRAKAWA T, et al. Identification of two distinct structural motifs that, when added to the C-terminal tail of the rat LH receptor, redirect the internalized hormone-receptor complex from a degradation to a recycling pathway [J]. Mol Endocrinol, 2001, 15: 1624-1635.
[55] TSAI-MORRIS C H, WEI W, Buczko E, et al.. Promoter and regulatory regions of the ratluteinizing hormone receptor gene [J]. J Biol Chem, 1993, 268:4447-4452.
[56] TSAI-MORRIS C H, GEN Y, XIE X Z, et al. Transcriptional protein binding domains governing basal expression of the rat luteinizing hormone receptor gene [J]. J Biol Chem, 1994, 269:15868-15875.
[57] TSAI-MORRIS C H, GEN Y, BUCZKO E, et al. Characterozation of diverse functionalelements in the upstream Sp1 domain of the rat luteinizing hormone receptor gene promoter [J]. J Biol Chem, 1995, 270:7487-7494.
[58] ZHANG F P, HAMALAINEN T, KAIPIA A, et al. Ontogeny of luteinizing hormone receptor gene expression in the rat testis [J]. Endocrinology, 1994, 134(5):2206-2213.
[59] TENA-SEMPERE M, ZHANG F P , HHTANIEMI I. Persistent expression of a truncated form of the lueinizing hormone receptor messenger ribonucleic acid in the rat testis after selective leydig cell destruction by ethylene dimethane sulfonate [J]. Endodrinol, 1994, 135:1018-1024.
[60] SOKKA T A, HAMALAINEN T M, KAIPIA A, et al. Development of Luteinizing Hormone Action in the Perinatal Rat Ovary [J]. Bio Reprod, 1996, 55:663-670.
[61] O'SHAUGHNESSY P J, MCLELLAND D, MCBRIDE M W. Regulation of Luteiniz ing Horm one-Receptor and Follicle-Stimulating Hormone-Receptor Messenger Ribonucleic Acid Levels during Development in the Neonatal Mouse Ovary [J]. Bio Reprod, 1997, 57:602-608.
[62] BAO B, GARVERICK H A, SMITH G W, et al. Changes in messenger ribonucleic acid encoding luteinizing hormone receptor, cytochrome P450-side chain cleavage, and aromatase are associated with recruitment and selection of bovine ovarian follicles [J]. Biol Reprod, 1997, 56(5):1158-1168.
[63] ABDENNEBI L, MONGET P, PISSELET C, et al. Comparative expression of luteinizing hormone and follicle-stimulating hormone receptors in ovarian follicles from high and low prolific sheep breeds [J]. Biol Reprod, 1999, 60(4):845-854.
[64] DERECKA K, PIETILA E M, RAJANIEMI H J, et al. Cycle-dependent LH/hCG receptor gene expression in porcine nongonadal reproductive tissues [J]. J Physiol Pharmacol, 1995, 46(1):77-85.
[65] LIN J, LEI Z M, LOJUN S, et al. Increased expression of luteinizing hormone/ human chorionic gonadotropin receptor gene in human endometrial carcinomas [J]. J ClinEndocrinol Metab, 1994,79:1483-1491.
[66] HAN S W, LEI Z M, RAO CH V, Homologous down-regulation of luteinizing hormone/chorionic gonadotropin receptors by increasing the degradation of receptor transcripts in human uterine endometrial stromal cells. Biol. Reprod. 1997, 57:158-164.
[67] SHEMESH M, MIZRACHI D, GUREVICH M, et al. Expression of functional luteinizing hormone (LH) receptor and its messenger ribonucleic acid in bovine endometrium: LH augmentation of cAMP and inositol phosphate in vitro and human chorionic gonadotropin (hCG) augmentation of peripheral prostaglandin in vivo [J]. Reprod Biol, 2001, 1:13-32.
[68] PIETILA E M, AATSINKI J T, Rajaniemi H.J.. Expression of the LH/CGreceptor gene in various rat tissues [J]. J Physiol Pharmacol, 1996, 47(1):5-14.
[69] ZHENG M, SHI H, SEGALOFF D L, et al. Expression and localization of luteinizing hormone receptor in the female mouse Reproductive tract [J]. Biol Reprod, 2001, 64:179-187.
[70] SHEMESH M. Actions of gonadotrophins on the uterus [J]. Reproduction, 2001, 121: 835-842.
[71] ZIECIK A J, OSTROWSKA G, KISIELEWSKA J, et al. Distribution and cycle phase dependency of gonadot ropin receptors in musculature and blood vessels of the porcine broad ligament [J]. Exp Clin Endocrinol Diabetes, 1995,103:44-51.
[72] SHEMESH M, GUREVICH M, FIELDS M J, et al. Expression of functional luteinizing hormone (LH) receptorand it s messenger ribonucleic acid in bovine uterine veins: LH induction of cyclooxygenase and augmentation of prostaglandin production in bovine uterine veins [J]. Endocrinology, 1997, 138:4844-4851.
[73] RODWAY M R, RAO C V. A novel prospective on t he role of human chorionic gonadot ropin during pregnancy and gestational trophoblastic disease. Early Pregnancy [J]. Biology and Medicine, 1995, 3:176-187.
[74] LEI Z M, RESHEF E, RAO V. The expression of human chorionic gonadotropin/ luteinizing hormone receptors in human endometrial and myometrial blood vessels [J]. J Clin Endocrinol Metab, 1992, 75(2):651-659.
[75] SHEMESH M., GUREVICH M., SHOREL S, et al. Regulation of cyclooxygenase and products of the cyclooxygenase pathway in the bovine uterus. Reprod Dom Anim., 1996,31 :431-436.
[76] STEPIEN A, SHEMESH M, ZIECIK A J. Luteinising hormone receptor kinetic and LH induced prostaglandin production throughout the oestrous cycle in porcine endometrium [J]. Reprod Nutr Dev,1999, 39:663-674.
[77] ZIECIK A J, BODEK G, BILTEK A. Nongonadal LH receptors, their involvement in femail reproductive function and a new applicable approach [J]. Vet J, 2005,169 (1):75-84.
[78] ZIECIK A J, STEPIEN A, GAWRONSKA B. Importance of endometrial luteinizing hormone receptors in induction of luteolysis and maternal recognition of pregnancy in the pig [J]. Reproduction in Domestic Animals, 2000, 35 :190-192.
[79] LIN P C, LI X, LEI Z M, et al. Human cervix contains functional luteinizing hormone/ human chorionic gonadot ropin receptors [J]. The Journal of Clinical Endocrinology & Metabolism, 2003, 88:3409-3414.
[80] STEPIEN A, DERECKA K, GAWRONSKA B, et al. LH/ hCG receptors in the porcine uterus a new evidence of their presence in the cervix and myometrium [J]. Physiol Pharmacol, 2000, 51:917-931.
[81]申颖,侯志高,王树迎.促性腺激素受体在子宫和输卵管中分布和作用的研究进展[J].动物医学进展. 2007, 28(6):86-88.
[82] LEI Z M, TOTH P, RAO Ch V, PRIDHAM D. Novel coexpression of human chorionic gonadotropin (hCG)/human luteinizing hormone receptors and their ligand hCG in human fallopian tubes [J]. J Clin Endocrinol Metab, 1993,77:863-872.
[83] HAN S W, LEI Z M, RAO C V. Up-regulation of cyclooxygenase-2 gene expression by chorionic gonadotropin in mucosal cells from human fallopian tubes [J]. Endocrinology, 1996, 137:2929-2937.
[84] SUN T, LEI Z M, RAO CH V. A novel regulation of the oviductal glycoprotein gene expression by luteinizing hormone in bovine tubal epithelial cells [J]. Mol Cell Endo, 1997, 131:97-108.
[85] GAWRONSKA B, PAUKKU T, HUHTANIEMI I, et al. Estrogen-dependent expression of LH/hCG receptors in pig fallopian tube and their role in relaxation of the oviduct [J]. J, Reprod Fertil, 1999, 115:293-301.
[86] DERECKA K, PIETILA E M, RAJANIEMI H J, et al. Cycle-dependent LH/hCG recaptor gene expression in porcine nongonadal reproductive tissues [J]. J Physiol Pharmacol, 1995, 46:77-85.
[87] GAWRONSKA B, STEPIEN A, ZIECIK A J. Effect of estradiol and progesterone on oviductal LH-receptors and LH-dependent relaxation of the porcine oviduct [J]. Theriogenology, 2000, 53:659-672.
[88] GAWRONSKA B, STEPIEN A, ZIECIK A J. Role of luteinizing hormone in control of oviduct function [J]. Reprod Domest Anim, 2000, 35:129-133.
[89] MISHRA S, LEI Z M, RAO Ch V, A novel role of luteinizing hormone in the embryodevelopment in cocultures [J]. Biol Reprod, 2003, 68:1455-1462.
[90] YOU S, KIM H, HSU C C, et al. Three different turkey luteinizing hormone receptor (tLH-R) isoforms I: characterization of alternative spliced tLH-R isoforms and their regulated expression in diverse tissues [J]. Biol Reprod, 2000, 62:108-116.
[91] FIELDS M J, SHEMESH M. Extragonadal luteinizing hormone receptors in the reproductive tract of domestic animals [J]. Bio Reprod, 2004, 71:1412-1418.
[92] SHI H, SEGALOFF D L. A role for increased lutropin/choriogonadotropin receptor (LHR) gene transcription in the follitropin-stimulated induction of the LHR in granulosa cells [J]. Mol Endocrinol, 1995, 9(6):734-744.
[93] IKEDA S, NAKAMURA K, KOGURE K., et al. Effect of estrogen on the expression of luteinizing hormone-human chorionic gonadotropin receptor messenger ribonucleic acid in cultured rat granulosa cells [J]. Endocrinology, 2008, 149(4):1524-1533.
[94] HIRAKAWA T, MINEGISHI T, ABE K, et al. A role of insulin-like growth factor I in luteinizing hormone receptor expression in granulosa cells [J]. Endocrinology, 1999, 140(11):4965-4971.
[95] ROY S K. Epidermal growth factor and transforming growth factor-beta modulation of follicle-stimulating hormone-induced deoxyribonucleic acid synthesis in hamster preantral and early antral follicles [J]. Biol Reprod, 1993 48(3):552-557.
[96]唐杰.山羊促性腺激素及其受体mRNA丰度的研究[D].河北:河北农业大学,2007.
[97] JOHANN S, HYESOOK Y, MYOUNGKUN J, et al. Orientation of Follicle-stimulati ng Hormone (FSH) Subunits Complexed with the FSH Receptor p subunit toward the N terminus of exodomain and a subunit to exoloop[J]. J Biol Chem, 2003, 278(48) :47868-47876.
[98] ANDREWS W V, MAURER R A, CONN P M. Stimulation of rat luteinizing hormon-ebeta messenger RNA levels by gonadotropin releasing hormone. Apparent role for protein kinaseC [J]. J Biol Chern, 1988, 263:13755-13761.
[99] BEN-MENAHEM D, NAOR Z. Regulation of gonadotropin mRNA levels in cultured rat pituitary cells by gonadotropin-releasing honnone(GnRH): role for Ca2+ and protein kinase C [J]. Biochemistry, 1994, 33:3698-3704.
[100] SUNDARESAN S, COLIN I M, PESTELL R Q et al. Stimulation of mitogenactivated protein kinase by gonadotropin-releasing hormone: evidence for the involvement of protein kinase [J]. Endocrinology, 1996, 137:304-311.
[101] MEUER K A, COX N M, MATAMOROS I A, et al. Decreased follicular steroids and insulin-like growth factor-I and increased atresia in diabetic gilts during follicular growth stimulated with PMSG [J]. J Reprod Fertil, 1991, 91(1):187-196.
[102] SHORT R E, ADAMS D C. Nutrition and hormonal interrelationships in beef cattle reproduction [J]. Anim Sci, 1998, 68:29-34.
[103] KLINDT J,YEN J T, Christenson R K, et al. Effect of prepubertal feeding regimen on reproduce-tive performance of gilts [J]. Anim Sci, 1999, 4:620-624.
[104] REESE D E, MOSER B D, PEO ER J R, et al. Influence of energy intake during lactation on the interval from weaning to first estrus in sows [J].Anim Sci, 1982, 55: 590-598.
[105] SHORT R E, BELLOWS R A. Relationships among weight gains, age at puberty and reproductive performance in heifers [J]. J Anim Sci, 1971,32, 127-131.
[106] GONZALEZ-PADILLA E, WILTBANK J N, et al. Puberty in beef heifers. I. The interrelationship between pituitary, hypothalamic and ovarian hormones [J]. J Anim Sci, 1975, 40:1091-1104.
[107] MURPHY M G, ENRIGHT W J, CROWE M A, et al. Effects of dietary intake on pattern of growth of dominant follicles during the oestrous cycle in beef heifers [J]. J Reprod Fertil, 1991, 92: 333-338.
[108] BEAM S W, BUTTER W R. Energy balance effects on follicular development and first ovulation in post-partum cows [J]. J Reprod Fertil Suppl, 1999, 54:411-424.
[109] BOOTH P J, COSGROVE J R, FOXCROFT G R. Endodrine and Metabolic Responses to Realimentation in Feed-Restricted Prepubertal Gilts: Associations Among Gonadotropins, Metabolic Hormones, Glucose, and Uteroovarian Development [J]. Anim Sci, 1996, 74:840-848.
[110] BOOTH P J, COSGROVE J R, FOXCROFT G R. Nutritional Manipulation of Growth andMetabolic and Reproductive Status in Prepubertal Gilts [J]. J Anim Sci, 1994, 72:2415-2424.
[111] ARMSTRONG J D, BRITT J H. Nutritionally-induced anestrus in gilts: metabolic and endocrine changes associated with cessation and resumption of estrous cycles [J]. J Anim Sci, 1987, 65(2):508-523.
[112] ZAK L J, COSGROVE J R, AHERNE F X, et al. Pattern of feed intake and associated metabolic and endocrine changes differentially affect postweaning fertility in primiparous lactating sows [J]. J Anim Sci, 1997, 75(1):208-216.
[113] VANDEN Brand H, DIELEMAN S J, SOEDE N M, et al. Dietary energy source at two feeding levels during lactation of primiparous sows: I. Effects on glucose, insulin, and luteinizing hormone and on follicle development, weaning-to-estrus interval, and ovulation rate [J]. J Anim Sci, 2000, 78:396-404.
[114] ZAK L J, XU X, HARDIN R T, et al. Impact of different patterns of feed intake during lactation in the primiparous sow on follicular development and oocyte maturation [J]. J Reprod Fertil, 1997, 110(1):99-106.
[115] ALMIDAl F R, KIRKWOOD R N, AHERNE F X, et al. Consequences of different patterns of feed intake during the estrous cycle in gilts on subsequent fertility [J]. J Anim Sci, 2000, 78(6):1556-1563.
[116] QUESNEL H, PASQUIER A, MOUNIER A M, et al. Influence of Feed Restriction During Lactation on Gonadotropic Hormones and Ovarian Development in Primiparous Sows [J]. J Anmi Sci, 1998,76:856-863.
[117] BRITT J H, ARMSTRONG J D, COX N M, et al. Control of follicular development during and after lactation in sows [J]. J Reprod Fertil Suppl, 1985, 33:37-54.
[118] MAURASSE C., MARTON P, DUFOUR J J, et al. Ovarian follicular populations at two stages of anoestrus cycle in heifers given high energy diets [J]. J anim Sci, 1985, 61: 1194-1200.
[119] GASSER C L, BEHLKE E J, GRUM D E, et al. Effect of timing of feeding a high-concentrate diet on growth and attainment of puberty in early-weaned heifers [J]. J Anim Sci, 2006, 84(11):3118-3122.
[120] MACKEY D R, SREENAN J M, ROCHE J F, et al. Effect of acute nutritional restriction on incidence ofanovulation and periovulatory estradiol and gonadotropin concentrations in beef heifers [J]. Biol Reprod, 1999, 61:1601-1607.
[121] KENDRICK K W, BAILEY T L, GARST A S, et al. Effects of energy balance of hormones, ovarian activity, and recovered oocytes in lactating Holstein cows using transvaginal follicular aspiration [J]. Dairy Sci, 1999, 82:1731-1741.
[122] JORRITSMA R, GROOT M W, VOS P L, et al. Acute fasting in heifers as a model for assessing the relationship between plasma and follicular fluid NEFA concentrations [J]. Theriogenology, 2003, 60(1):151-161.
[123]李勇,吴德.能量对家畜卵泡发育的影响及调控[J].饲料工业, 2007, 28(21):15-19.
[124]唐晓龙.不同能量和蛋白水平日粮对母羊繁殖性能的影响[D].河北:河北农业大学, 2007.
[125] KILLEEN ID. Effects of fasting ewes before mating on their reproductive performance [J]. Theriogenology, 1982,17:433-435.
[126] KIYMA Z, ALEXANDERl B M, VAN KIRK E A. Effects of feed restriction on reproductive and metabolichormones in ewes [J]. Anim Sci, 2004, 82:2548-2557.
[127]刘立文.日粮能量对初情期前母猪FSH、LH、GH、胰岛素及IGF-I释放的影响[D].吉林:吉林大学, 2006.
[128]于淼瑛.不同日粮能量水平对初情期前母猪卵巢及子宫LH受体和FSH受体mRNA表达的影响[D].吉林:吉林大学, 2006.
[129] COX N M, STUART M J, ALTHEN T G, et al. Enhancement of ovulation rate in gilts by increasing dietary energy and administering insulin during follicular growth [J]. J Anim Sci, 1987, 64(2):507-516.
[130] BOOTH P J, COSGROVE J R, FOXCROFT G R. Endocrine and metabolic responses to realimentation in feed-restricted prepubertal gilts: associations among gonadotropins, metabolic hormones, glucose, and uteroovarian development [J]. J Anim Sci, 1996, 74(4): 840-848.
[131] MATAMOROS I A, COX N M, MOORE A B. Effects of exogenous insulin and body condition on metabolic hormones and gonadotropin-induced follicular development in prepuberal gilts [J]. J Anim Sci, 1991, 69(5):2081-2091.
[132]周平,吴德,王延忠,等.营养对母猪卵泡发育的影响[J].饲料博览. 2007, 15:25-27.
[133]于淼瑛,周虚.胰岛素在能量影响猪卵泡发育中的作用.黑龙江动物繁殖[J]. 2005, 13(2):12-15.
[134] HELEN Q, ANNE P, ARMELLE P, et al. Feed restriction in cyclic gilts: Gonadotrophin independent effects on follicular growth[J]. Reprod Nutr Dev, 2000, 40(4): 405-414.
[135] DRIANCOURT M A, LOCATELLI A, PRUNIER A. Effects of gonadotrophin deprivation on follicular growth in gilts [J]. Reprod Nutr Dev, 1995, 35(6):663-673.
[136] ALMEIDA F R, MAO J, NOVAK S, et al. Effects of different patterns offeed restriction and insulin treatmen tduring the luteal phase on reproductive, metabolic, and endocrine parameters in cyclic gilts [J]. J Anim Sci, 2001,79 (1) :200-212.
[137] MAY J V, SCHOMBERG D W. Granulosa cell differentiation in vitro: effect of insulin on growth and functional integrity [J]. Biol Reprod, 1981,25(2):421-431.
[138]李波.能量对初情期前母猪卵巢和子宫IGF-ⅠR和EGFR mRNA表达的影响[D].吉林:吉林大学, 2007.
[139] ADAM C L, GADD T S, FINDLAY P A, et al. IGF-1 stimulation of luteinizing hormone secretion, IGF-binding proteins (IGFBPs) and expression of mRNAs for IGFs, IGF receptors and IGFBPs in the ovine pituitary gland [J]. J Endocri, 2000, 166: 247-254.
[140] RIVERA G M, FORTUNE J E. Proteolysis of insulin-like growth factor binding proteins-4 and -5 in bovine follicular fluid :implications for ovarian follicular selection and dominance [J]. Endocrinology, 2003, 144 (7) :2977-2987.
[141] LEEUWENBERG B R, HURST P R, MCNATTY K P. Expression of IGF-I mRNA in the ovine ovary [J]. J Mol Endocrinol, 1995, 15(3):251-258.
[142] DEVOTO L, KOHEN P, CASTRO O, et al. Multihormonal regulation of progesterone synthesis in cultured human midluteal cells [J]. J Clin Endocrinol Metab, 1995, 80(5):1566-1570.
[143] FOSTER C M, HASHIMOTO N, RANDOLPH J F, et al. Comparison of growth hormone and insulin-like growth factor-I regulation of estradiol and progesterone production in human luteinized granulosa cells [J]. Pediatr Res, 1995, 38(5):763-767.
[144]刘丽,李琦华,贾俊静,等.瘦素对动物能量代谢及繁殖性能的影响[J].饲料研究, 2008, 6, 26-28.
[145] BARB C R, HAUSMAN G J, CZAJA K, et al. Leptin: a Metabolic Signal Affecting Central Regulation of Reproduction in the Pig [J]. Domest Anim Endocrinol, 2005, 29(1):186-192.
[146] RUIZ-CORTE′S Z T, MEN T, PALIN M F, et al. Porcine leptin receptor: molecular structure and expression in the ovary [J]. Mol Reprod Dev, 2000, 56:465-474.
[147] SPICER L J, CHAMBERLAIN C S, FRANCISCO C C, et al. Ovarian action of leptin: effects on insulin-like growth factor-I-stimulated function of granulosa and thecal cells [J].Endocrine, 2000, 12(1):53-59.
[148] Guo X, Chen S, Xing F, et al. Effects of leptin on estradiol and progesterone production by human luteinized granulosa cells in vitro [J]. Zhonghua Fu Chan Ke Za Zhi. 2001, 36(2): 95-97.
[149] GHIZZONI L, BARRECA A, MASTORAKOS G, et al. Leptin inhibits steroid biosynthesis by human granulosa-lutein cells [J]. Horm Metab Res, 2001, 33(6):323-328.
[150] TATIANA RUIZ-CORTES Z, MARTEL-KENNES Y, NICOLAS Y, et al. Biphasic Effects of Leptin in Porcine Granulosa Cells [J]. Biol Reprod, 2003, 68:789-796.
[151] GONG J G, BRAMLEY T, WEBB R. The effect of recombinant bovine somatotropinon ovarian function in heifers. follicular populations and peripheral hormones [J]. BiolReprod, 1991, 45: 941-949.
[152] HERRLER A, EINSPANIER R, SCHAMS D, et al. Effect of recombinant bovine somatotropin (rBST) on follicular IGF-1contents and the ovarian response following superovulatrory treatment in dairy cows . Apreliminary study [J]. Theriogenology, 1994, 41:601-611.
[153] GONG J G, BAXTER G, BRAMLEY T A, et al. Enhancement of ovarian follicle development in heifers by treatment with recombinant bovine somatotropin. adose-response study [J]. J Reprod Fertil, 1997, 110:91-97.
[154] ZIECIK A J, DERECKA-RESZKA K, RZUCIDLO S J. Extragonadal gonadotropin receptors, their distribution and function [J]. J Physiol Pharmacol, 1992, 43:33-49.
[155] FRIEDMAN S, GUREVICH M, SHEMESH M. Bovine cyclic endometrium contains high-affinity luteinizing hormone/human chorionic gonadotropin binding sites [J]. Biol Reprod, 1995, 92:1020-1026.
[156] ZIECIK A J, STANCHEV P D, TILTON J E. Evidence for the presence of luteinizing hormone/human chorionic gonadotropin-binding sites in the porcine uterus[J]. Endocrinology, 1986, 119:1159-1163.
[157] RZUCIDLO S J, WEIGL R M, TILTON J E. Myometrial LH/hCG receptors during the estrous cycle and pregnancy in pigs [J]. Anim Reprod Sci, 1998, 51:249-257.
[158] BONNAMY P J, BENHAIM A, LEYMARIE P. Human chorionic gonadotropin affects tissue levels of progesterone and cyclic adenosine 39,59-monophosphate in the metestrous rat uterus in vitro [J]. Biol Reprod, 1989, 40:511-516.
[159] SAWITZKE A L, ODELL W D. Uterine binding sites for LH/hCG can be modulated byhormonal status in rabbits and rats [J]. Acta Endocrinol, 1991, 124:322-330.
[160] JENSEN T D, ODELL W D. Identification of LH/hCG receptors in rabbit uterus [J]. Proc Soc Exp Biol Med, 1988, 189:28-30.
[161] RESHEF E, LEI Z M, RAO CH V, et al. The presence of gonadotropin receptors in nonpregnant human uterus, human placenta, fetal membranes, and decidua [J]. J Clin Endocrinol Metab, 1990, 70:421-430.
[162] SHEMESH M, Hansel W. Levels of prostaglandin F(PGF) in bovine endometrium , uterine venous , ovarian arterial and jugular plasma during the estrous cycle [J]. Proc Soc Exp Biol Med, 1975, 148 :123-126.
[163] TOTH P, LI X, RAO C V, et al. Expression of functional human chorionic gonadotropin/ humanluteinizing hormone receptor gene in human uterine arteries [J]. J Clin Endocrinol Metab, 1994, 79:307-315.
[164] LIN D X , LEI Z M. Dependence of Uterine Cyclooxygenase Expression on Luteinizing Hormone Signaling [J]. Biol Reprod, 2005, 73:256-260.
[165] KORNYEI J L, LI X, LEI Z M, et al. Restoration of human chorionic gonadotropin response in human myometrial smooth muscle cells by treatment with follicle-stimulating hormone (FSH): evidence for the presence of FSH receptors in human myometrium [J]. European Journal of Endocrinology, 1996, 134(2): 225-231.
[166] GAWRONSKA B, STEPIEN A , ZIECIK A J . Effect of estradiol and progesterone on oviduc-tal LH-receptors and LH-dependent relaxation of the porcine oviduct[J]. Theriogenology, 2000, 53:659-672.
[167] RZUCIDLO S J, WIGL R M, TILTON J E. Myometrial LH/hCG receptors during the estrous cycle and pregnancy in pigs [J]. Anim Reprod Sci, 1998, 51:249-257.
[168] ZIECIK A J, KOTWICA G. Involvement of gonadotropins in induction of luteolysis in pigs [J]. Reprod Biol, 2001, 1:33-50.
[169] WEEMS Y S, KIM L, HUMPHREYS V, et al. Effect of luteinizing hormone (LH), pregnancy specific protein B (PSPB), or arachidonic acid (AA) on ovine endometrium of the estrous cycle or placental secretion of prostaglandins E2 (PGE2) and F2α(PGF2α) and progesterone in vitro [J]. Prostaglandins, 2003, 71:55-73.
[170] ZHOU X L, LEI Z M, RAO CH V. Treatment of human endometrial gland epithelial cells with chorionic gonadotropin/luteinizing hormone increases the expression of thecyclooxygenase-2 gene [J]. J Clin Endocrinol Metab, 1999, 84:3364-3377.
[171] COSGROVE J R, TILTON J E, HUNTER M G, et al. Gonadotrogen-independent mechanisms in feed–restricted prepubertal gilts [J]. Bio Reprod, 1992, 47: 736-745.
[172] COSGOVE J R, TITON T G, ALTHEN W A, et al. Maturational changes in gonadonatrophin secretion the LH response to realimentation and a nocturnal in crement in LH secrection of feed-restricted prepubertal gilts [J]. Reprod. Fertil, 1993, 98: 293-305.
[173] BLANK J L, DESJARDINS C. Differential effects of food restriction on pituitary- testicular function in mice [J]. Am J Physiol, 1985, 248: 181-189.
[174] FOSTER D L, OLSTER D H. Effect of restricted nutrition on puberty in the lamb: patterns of tonic luteinizing hormone (LH) secretion and competency of the LH surge system [J]. Endocrinology, 1985, 116:375-381.
[175] THOMAS G B, MERCER J E, KARALIS T, et al. Effect of restricted feeding on the concentrations of growth hormone (GH), gonadotropins, and prolactin (PRL) in plasma, and on the amounts of messenger ribonucleic acid for GH, gonadotropin subunits,and PRL in the pituitary glands of adult ovariectomized ewes [J]. Endocrinology, 1990, 126:1361-1367.
[176] CAMERON J L, NOSBISCH C. Suppression of pulsatile luteinizing hormone and testosterone secretion during short term food restriction in the adult male rhesus monkey (Macaca mulatta) [J]. Endocrinology, 1991, 128:1532-1540.
[177] ROJDMARK S. Influence of short-term fasting on the pituitary-testicular axis in normal men [J]. Horm Res, 1987, 25:140-146.
[178] RHODES M T, MINION J E, STEVENSON J S, et al. Endocrine changes associated with flushing and altrenogest treatment in gilts [J]. Anim. Sci, 1987, 65(1):154-161.
[179] FLOWERSl B, MARTIN M J, CANTLEY T C. et al. Endocrine Changes Associated with a Dietary-Induced Increase in Ovulation Rate (Flushing) in Gilts [J]. J Anim Sci, 1989, 67:771-778.
[180] COSGROVE J R, TILTON J E, HUNTER M G, et al. Gonadotrogen-independent mechanisms in feed–restricted prepubertal gilts [J]. Bio Reprod, 1992, 47: 736-745.
[181] SNYDER J L, CLAPPERl J A, ROBERTS A J, et al. Insulin-like growth factor-I, insulin-like growth factor-binding proteins, and gonadotropins in the hypothalamicpituitary axis and serum of nutrient restricted ewes [J]. Biol. Reprod,1999, 61: 219-224.
[182] KILE J P, ALEXANDEl B M, MOSS G E, et al. Gonadotropin-releasing hormone overridesthe negative effect of reduced dietary energy on gonadotropin synthesis and secretion in ewes [J]. Endocrinology,1991, 128:843-849.
[183] RHIND S M, LESLIE L D, GUNN R G, et al. Plasma FSH, LH, prolactin and progesterone profiles of Cheviot ewes with different levels of intake before and after mating, and associated effects on reproductive performance [J]. Anim Reprod Sci, 1985, 8:301-313.
[184] GOMBE S, HANSEL W. Plasma luteinizing hormone (LH) and progesterone levels in heifers on restricted energy intakes [J].Anim Sci, 1973, 37:728-733.
[185] MCCANN J P, HANSEL W. Relationships between insulin and glucose metabolism and pituitary-ovarian functions in fasted heifers [J]. Biol. Reprod, 1986, 34:630-641.
[186] SCHRICK F N, SPITZER J C, JENKINS T C, et al. Effect of dietary energy restriction on metabolic and endocrine responses during the estrous cycle of the suckled beef cow [J]. Anim Sci, 1990, 68:3313-3321.
[187] COOPER K, BROOKSR P H, COLE D J A, et al. The effect of feed level during the estrous cycle on ovulation, embryo survival and anterior pituitary LH potency in the gilt [J]. J Reprod Fertil, 1973, 32:71-78.
[188] KIRKPATRICK, R L, HOWLAND B E, FIRST N L, et al. Ovarian and pituitary gland changes in gilts on two nutrient energy levels [J]. J Anita Sci, 1967, 26:358-364.
[189] KEMP B, SOEDE N M, HELMOND F A, et al. Effects of energy source in the diet on reproductive hormones and insulin during lactation and subsequent estrus in multiparous sows [J]. Anim Sci, 1995, 73:3022-3029.
[190] KOKETSU Y, DIAL G D, PELLIGREW J E, et al. Influence of imposed feed intake patterns during lactation on reproductive performance and on circulating levels of glucose, insulin, and luteinizing hormone in primiparous sows [J]. Anim Sci, 1996, 74:1036-1046.
[191] BAIDOO S K, LYTHGOEy E S, KIRKWOOD R N, et al. Effect of lactation feed intake on endocrine status and metabolite levels in sows [J]. Anim Sci, 1992, 72:799-807.
[192] DOMINGUEZ M M. Effects of body condition, reproductive status and breed on follicular population and oocyte quality in cows [J].Theriogenology, 1995, 43:1405-1418.
[193] GHARIB S D, WIERMAN M E, SHUPNIK M A, et al. Molecular biology of the pituitary Gonadotropins [J]. Endocr Rev, 1990, 11:177-199.
[194] MARSHALL JC M, CHRISTINE A E, CHRISTOPER R M. Hypothalamic dysfunction [J]. Mol Cell Endocrinol, 2001, 183: 29-32.
[195] NETT T M, TURZILLO A M, BARATTA M, et al. Pituitary effects of steroid hormones on secretion of follicle-stimulating hormone and luteinizing hormone [J]. Dourest Anim Endocrinol, 2002, 23: 33-42.
[196] WADE G N, SCHNEIDER J E. Metabolic fuels and reproduction in female animal [J]. Neurosci, Biobehav, 1992, 16: 235-272.
[197] MONGET P, MARTIN G B. Involvement of insulin-like growth factor in the interactions between nutrition and reproduction in female mammals [J]. Human Reprod, (Oxf.), 1997, 12: 33-52.
[198] MOUNIZH K, LU R , CHEHAB F F. Leptin treatmeant rescues the terility of genetically ob/ob males [J]. Endocrinology, 1997, 138: 1190-1193.
[199] VIGUIE C, BATTAGLIA D F, KRASA H A, et al. Thyroid hormones act primarily within the brain to promote the seasonal inhibition of luteinizing hormone secretion in the ewe [J]. Endocrinology, 1999, 140: 1111-1117.
[200] LANDAU, B R, TAKAOKA Y, ABRAMS M A, et al. Binding of insulin by monkey and pig hypothalamus [J]. Diabetes, 1983, 32:284-291.
[201] HAVRANKOVA J, SCHMECHEL D, ROTH J, et al. Identification of insulin in rat brain [J]. Proc Natl Acad Sci, USA 1978, 75:5737-5742.
[202] ADASHI E Y HSUEHA J W, YENS S C. Insulin enhancement of luteinizing hormone and follicle-stimulating hormone release by cultured pituitary cell [J]. Endocrinology, 1981, 108:1441-1449.
[203] ARMSTRONG D G, MCEVOY T G, BAXTER G J. et al. Effect of dietary energy and protein on bovine follicular dynamics and embryo production in vitro: associations with the ovarian insulin-like growth factor system [J]. Biol Reprod, 2001, 64:1624-1632.
[204] BOSSIS I R. WETTEMANN P, WELTY S D J. et al. Nutritionally induced anovulation in beef heifers: ovarian and endocrine function during realimentation and resumptions of ovulation [J]. Biol Reprod, 2000, 62:1436-1444.
[205] GRIGSBY M E, TRENKLE A. Plasma growth hormone, insulin, glucocorticoids and thyroid hormones in large, medium and small breeds of steers with and without an estradiol implant [J]. Domest Anim Endocrinol, 1986, 3:261-267.
[206] RICHARDS M W, WELLEMAN R P, SPICER L J, et al. Nutritional anestrus in beef cows: Effect of body condition and ovariectomy on serum luteinizing hormone and insulin-likegrowth factor-I [J]. Biol Reprod, 1991, 44:961-966.
[207] SIMPSON R B, CHASE C C, SPICER L J, et al. Effect of exogenous estradiol on plasma concentrations of somatotropin, insulin-like growth factor-I, insulin- like growth factor binding protein activity, and metabolites in ovariectomized Angus and Brahman cows [J]. Domest Anim Endocrinol, 1997, 14:367-380.
[208] ARMSTRONG, D G, GONG J G, WEBB R. Interactions between nutrition and ovarian activity in cattle: Physiological, cellular and molecular mechanisms [J]. Repro duction Suppl, 2003, 61:403-414.
[209] ZAMORANO P L, DE SEVILLA L, MAHESH V B, et al. Production and refolding of recombinant leptin [J]. Biotechniques, 1997, 23(5):800-804.
[210] LONG J, BARB C R, KRAELELING R R, et al. Growth hormone releaseing factor decrease long form Leptin receptor expression in porcine anterior pituitary cells [J]. Domestic Anim Endocrinology, 2003, 141:333-339.
[211] SANJAY K, AGARW A L, KLARA L, et al. Leptin Antagonizes the Insulin-Like Growth Factor-I Augmentation of Steroidogenesis in Granulosa and Theca Cells of the Human Ovary [J].Clin Endocrinol Metab, 1999, 84: 1072-1076.
[212] WISHANNT C S, HARELL R J. Leptin and luteinnizing homone concentration in pigs. College of agriculture and life sciences department of animal science annual swine report, 2001. 27:367-378.
[213] AMSTALDEN M, GARCIA M R, WILLIAMS S W, et al. Leptin gene expression, circulating leptin, and luteinizing hormone pulsatility are acutely responsive to short-term fasting in prepubertal heifers: Relationships to circulating insulin and insulin-like growth factor-I [J]. Biol Reprod, 2000, 63:127-133.
[214] BRYAN K A. Effect of doses of porcine somatotropin (pST) on growth, carcass traits and granulosa cell function of gilts [J]. Anim Sci, 1988, 66 (1): 379-385.
[215] BRYAN K A, HAMMOND J M, CANNING S, et al. Reproductive and growth responses of gilts to exogenous porcine pituitary omatotropin [J]. Anim Sci, 1989, 67: 196-205.
[216] ADASHI, E Y, HSUEH AJ W, Yen S S C. Insulin enhancement of luteinizing hormone and follicle-stimulating hormone release by cultured pituitary cell [J]. Endocrinology, 1981, 108:1441-1449.
[217] WANDJI S A, WOOD T L, CRAWFORD J, et al . Expression of mouse ovarian insulin-likegrowth factor system components during follicular development and atresia [J]. Endocrinol, 1998, 139 (12): 5205-5214.
[218]葛立军,芮荣.促黄体素受体在母畜生殖道的分布与表达[J].动物医学进展, 2007, 28 (11) :92-95.
[219] MINEGISHI T, HIRAKAWA T, ABE K. A Role of Insulin-Like Growth Factor I in Luteinizing Hormone Receptor Expression in Granulosa Cells [J]. Endocrinology, 1999, 11: 4965-4971.
[220] ZHOU J, KUMAR T R, MATZUK M M, et al. Insulin-like growth factor I regulates gonadotropin responsiveness in the murine ovary [J]. Mol Endocrinol, 1997, 11:1924-1933.
[221] BAKER J, HARDY M P, ZHOU J, et al. Effects of an Igf1 gene null mutation on mouse reproduction [J]. Mol Endocrinol, 1996, 10:903-918.
[222] TILLY J L, LAPOLT P S, HSUECH J W. Hormonal regulation of follicle stimulating hormone receptor messenger ribonucleic acid level in cultured rat granulose cells [J]. Endocrinology, 1992, 130(3) : 1296-1302.