fbpx
Wikipedia

Gonadotropin-inhibitory hormone

Gonadotropin-inhibitory hormone (GnIH) is a RFamide-related peptide coded by the NPVF gene in mammals.

Discovery edit

GnIH was discovered in 2000. It is an RFamide peptide that significantly reduced luteinizing hormone release in Coturnix Japonica (Japanese quail). This peptide emerged as the first tropic hormone known to inhibit gonadotropin secretion in the hypothalamic-pituitary-gonadal axis of vertebrates.[1] Subsequent research identified GnIH peptide homologs in variety of mammals, including humans.[2]

Structure edit

GnIH is a neurohormone classified as an RFamide (RFa) or RFamide-related peptide (RFRP), coded by the NPVF gene in mammals. The complete amino acid sequence varies by species, but all RFa and RFRP peptides contain an arginine-phenylalanine-amine sequence at the C-terminal. This is seen in both Coturnix Japonica GnIH RFa (Ser-Ile-Lys-Pro-Ser-Ala-Tyr-Leu-Pro-Leu-Arg-Phe-NH2), and the human homolog, RFRP-3 (Val-Pro-Asp-Leu-Pro-Glu-Arg-Phe-NH2).[1][3]

Production edit

GnIH neurons reside primarily in the dorsomedial nucleus of the hypothalamus (humans and rodents) and the paraventricular nucleus of the hypothalamus (avian species). Some GnIH neuron terminals in both mammalian and avian species project to the median eminence.[4][5][6] GnIH and GnRH (gonadotropin releasing hormone) neurons exist in close proximity in the hypothalamus, which may enable the direct inhibition of GnRH neurons by GnIH.[7] GnIH enters the bloodstream via the hypothalamo-hypophyseal portal system, the vascular network supplying both the hypothalamus and the pituitary.[8]

GnIH and GnIH receptor (GnIH-R) mRNA is expressed in the hypothalamus, pituitary, and ovaries.[5] GnIH expression is highest during proestrus and lowest during estrus, suggesting the estrus cycle influences release of the hormone. Furthermore, GnIH neuronal cell counts in multiple vertebrates fluctuate with an organism’s parental status.[9] GnIH cell count may also vary with breeding season in some species. For instance, European starlings (Sturnus vulgaris) with greater reproductive success exhibited higher quantities of GnIH-producing cells than did those that were less successful, but this effect did not appear until mid-breeding season.[9][10][11]

Receptor action edit

GnIH binds to the Gαi protein coupled receptor GPR147 to suppress adenylyl cyclase formation of cAMP and inhibit protein kinase cascades affecting gene expression. GnIH inhibits the same signaling pathway that GnRH activates to promote follicle stimulating hormone (FSH) and luteinizing hormone (LH) expression.[12][13] The compound RF9 is a known GPR147 receptor antagonist.[14]

Effects and physiological function edit

GnIH-R expression in the pituitary and other brain regions implies GnIH acts directly on the pituitary to downregulate gonadotropin production, impacting reproductive behaviors.[6][15][16][17] This neurohormone also acts on the hypothalamus to inhibit the expression of GnRH, which may further inhibit gonadotropin secretion, and kisspeptin, which may inhibit kisspeptin-mediated stimulation of GnRH neurons prior to the preovulatory hormonal surge. GnIH also spurs the production of cytochrome P450 aromatase, promoting the synthesis of neuroestrogen in the brains of quails and reducing aggressivity in reproductive behaviors.[7][18][19]

In male vertebrates, GnIH reduces testis size, lowers testosterone secretion, and increases the incidence of apoptosis in germ cells and Sertoli cells of the seminiferous tubules.[20][21] These gonadal changes, in addition to GnIH and GnIH-R mRNA expression in the seminiferous tubules, Sertoli cells, and spermatogonia, implicate function in spermatogenesis. In female vertebrates, high doses of GnIH increases ovarian mass and produce follicle irregularities, such as vacuole formation in nuclei and distorted morphology.[22] Ovarian changes in response to GnIH administration, as well as GnIH/GnIH-R mRNA expression in granulosa cells and luteal cells in different stages of the estrus cycle, implicate function in development of follicles and atresia.[21]

Additional biological roles edit

Stress-induced adrenal hormone increase may upregulate GnIH release, as some GnIH neurons have adrenal glucocorticoid receptors. GnIH may therefore mediate interactions between the HPG and HPA (hypothalamic-pituitary-adrenal) axes and play a role in stress-related infertility.[23] GnIH neurons of the paraventricular nucleus in the hypothalamus also express melatonin receptors. Because melatonin secretion is modulated by environmental light patterns, melatonin influence on GnIH production may enable photoperiodic regulation of reproduction in seasonally breeding birds, rodents, and sheep.[24]

GnIH increases food consumption, implying a role in appetite. This finding is consistent with the location of most GnIH neurons, as the dorsomedial nucleus of the hypothalamus is involved in appetite regulation. GnIH may allow the energy reserves of an organism to modulate reproduction.[6]

Higher levels of thyroid hormone suppress GnIH expression, and lower levels of thyroid hormone are associated with higher GnIH levels. The inactivation of GnIH expression prevents delayed puberty caused by hypothyroidism, demonstrating that GnIH mediates interactions between the HPG and HPT (hypothalamic-pituitary-thyroid) axes.[25] Furthermore, thyroid hormone may function in a pathway for photoperiodic regulation of reproduction involving GnIH and energy status. Melatonin modulates thyroid-stimulating hormone (TSH) production in the anterior pituitary, and TSH promotes thyroid hormone production. Thyroid hormone production influences metabolism and GnIH production, both of which impact reproduction.[26]

References edit

  1. ^ a b Tsutsui K, Saigoh E, Ukena K, Teranishi H, Fujisawa Y, Kikuchi M, Ishii S, Sharp PJ (August 2000). "A novel avian hypothalamic peptide inhibiting gonadotropin release". Biochemical and Biophysical Research Communications. 275 (2): 661–7. doi:10.1006/bbrc.2000.3350. PMID 10964719.
  2. ^ Fukusumi S, Habata Y, Yoshida H, Iijima N, Kawamata Y, Hosoya M, Fujii R, Hinuma S, Kitada C, Shintani Y, Suenaga M, Onda H, Nishimura O, Tanaka M, Ibata Y, Fujino M (September 2001). "Characteristics and distribution of endogenous RFamide-related peptide-1". Biochimica et Biophysica Acta (BBA) - Molecular Cell Research. 1540 (3): 221–32. doi:10.1016/S0167-4889(01)00135-5. PMID 11583817.
  3. ^ Findeisen M, Rathmann D, Beck-Sickinger AG (2011). "RFamide Peptides: Structure, Function, Mechanisms and Pharmaceutical Potential". Pharmaceuticals. 4 (9): 1248–1280. doi:10.3390/ph4091248. PMC 4058657.
  4. ^ Kriegsfeld LJ, Mei DF, Bentley GE, Ubuka T, Mason AO, Inoue K, Ukena K, Tsutsui K, Silver R (February 2006). "Identification and characterization of a gonadotropin-inhibitory system in the brains of mammals". Proceedings of the National Academy of Sciences of the United States of America. 103 (7): 2410–5. Bibcode:2006PNAS..103.2410K. doi:10.1073/pnas.0511003103. PMC 1413747. PMID 16467147.
  5. ^ a b Ubuka T, Morgan K, Pawson AJ, Osugi T, Chowdhury VS, Minakata H, Tsutsui K, Millar RP, Bentley GE (December 2009). "Identification of human GnIH homologs, RFRP-1 and RFRP-3, and the cognate receptor, GPR147 in the human hypothalamic pituitary axis". PLOS ONE. 4 (12): e8400. Bibcode:2009PLoSO...4.8400U. doi:10.1371/journal.pone.0008400. PMC 2791420. PMID 20027225.
  6. ^ a b c Smith JT, Clarke IJ (April 2010). "Gonadotropin inhibitory hormone function in mammals". Trends in Endocrinology and Metabolism. 21 (4): 255–60. doi:10.1016/j.tem.2009.11.010. PMID 20060314. S2CID 41737276.
  7. ^ a b Smith JT, Shahab M, Pereira A, Pau KY, Clarke IJ (October 2010). "Hypothalamic expression of KISS1 and gonadotropin inhibitory hormone genes during the menstrual cycle of a non-human primate". Biology of Reproduction. 83 (4): 568–77. doi:10.1095/biolreprod.110.085407. PMC 2957156. PMID 20574054.
  8. ^ Smith JT, Young IR, Veldhuis JD, Clarke IJ (July 2012). "Gonadotropin-inhibitory hormone (GnIH) secretion into the ovine hypophyseal portal system". Endocrinology. 153 (7): 3368–75. doi:10.1210/en.2012-1088. PMC 3380300. PMID 22549225.
  9. ^ a b Li X, Su J, Lei Z, Zhao Y, Jin M, Fang R, Zheng L, Jiao Y (August 2012). "Gonadotropin-inhibitory hormone (GnIH) and its receptor in the female pig: cDNA cloning, expression in tissues and expression pattern in the reproductive axis during the estrous cycle". Peptides. 36 (2): 176–85. doi:10.1016/j.peptides.2012.05.008. PMID 22664321. S2CID 902250.
  10. ^ Calisi RM, Díaz-Muñoz SL, Wingfield JC, Bentley GE (July 2011). "Social and breeding status are associated with the expression of GnIH". Genes, Brain and Behavior. 10 (5): 557–64. doi:10.1111/j.1601-183X.2011.00693.x. PMID 21466656.
  11. ^ Calisi RM, Geraghty AC, Avila A, Kaufer D, Bentley GE, Wingfield JC (October 2016). "Patterns of hypothalamic GnIH change over the reproductive period in starlings and rats". General and Comparative Endocrinology. 237: 140–146. doi:10.1016/j.ygcen.2016.08.015. PMID 27591072.
  12. ^ Ubuka T, Son YL, Tobari Y, Tsutsui K (2012). "Gonadotropin-inhibitory hormone action in the brain and pituitary". Frontiers in Endocrinology. 3: 148. doi:10.3389/fendo.2012.00148. PMC 3515997. PMID 23233850.
  13. ^ Ubuka T, Son YL, Bentley GE, Millar RP, Tsutsui K (September 2013). "Gonadotropin-inhibitory hormone (GnIH), GnIH receptor and cell signaling". General and Comparative Endocrinology. 10th International Symposium on Avian Endocrinology. 190: 10–7. doi:10.1016/j.ygcen.2013.02.030. PMID 23499786.
  14. ^ Caraty A, Blomenröhr M, Vogel GM, Lomet D, Briant C, Beltramo M (May 2012). "RF9 powerfully stimulates gonadotrophin secretion in the ewe: evidence for a seasonal threshold of sensitivity". Journal of Neuroendocrinology. 24 (5): 725–36. doi:10.1111/j.1365-2826.2012.02283.x. PMID 22283564. S2CID 25770293.
  15. ^ Ubuka T, Ukena K, Sharp PJ, Bentley GE, Tsutsui K (March 2006). "Gonadotropin-inhibitory hormone inhibits gonadal development and maintenance by decreasing gonadotropin synthesis and release in male quail". Endocrinology. 147 (3): 1187–94. doi:10.1210/en.2005-1178. PMID 16293662.
  16. ^ Tsutsui K, Bentley GE, Ubuka T, Saigoh E, Yin H, Osugi T, Inoue K, Chowdhury VS, Ukena K, Ciccone N, Sharp PJ, Wingfield JC (2007-08-01). "The general and comparative biology of gonadotropin-inhibitory hormone (GnIH)". General and Comparative Endocrinology. Proceedings of the 23rd Conference of European Comparative Endocrinologists: Part 2. 153 (1–3): 365–70. doi:10.1016/j.ygcen.2006.10.005. PMID 17141777.
  17. ^ Bentley GE, Jensen JP, Kaur GJ, Wacker DW, Tsutsui K, Wingfield JC (April 2006). "Rapid inhibition of female sexual behavior by gonadotropin-inhibitory hormone (GnIH)". Hormones and Behavior. 49 (4): 550–5. doi:10.1016/j.yhbeh.2005.12.005. PMID 16460739. S2CID 1801090.
  18. ^ Paullada-Salmerón JA, Cowan M, Aliaga-Guerrero M, Morano F, Zanuy S, Muñoz-Cueto JA (June 2016). "Gonadotropin Inhibitory Hormone Down-Regulates the Brain-Pituitary Reproductive Axis of Male European Sea Bass (Dicentrarchus labrax)". Biology of Reproduction. 94 (6): 121. doi:10.1095/biolreprod.116.139022. PMC 6322450. PMID 26984999.
  19. ^ Tsutsui K, Ubuka T (2016). "GnIH Control of Feeding and Reproductive Behaviors". Frontiers in Endocrinology. 7: 170. doi:10.3389/fendo.2016.00170. PMC 5186799. PMID 28082949.
  20. ^ Ubuka T, Ukena K, Sharp P, Bentley G, Tsutsui K. Gonadotropin-inhibitory hormone inhibits gonadal development and maintenance by decreasing gonadotropin synthesis and release in male quail. Endocrinology. 2006; 147, 1187-1194. doi: 10.1210/en.2005-1178
  21. ^ a b Tsutsui K, Ubuka T, Bentley GE, Kriegsfeld LJ (July 2012). "Gonadotropin-inhibitory hormone (GnIH): discovery, progress and prospect". General and Comparative Endocrinology. Profiles in Comparative Endocrinology: Eric Roubos. 177 (3): 305–14. doi:10.1016/j.ygcen.2012.02.013. PMC 3378827. PMID 22391238.
  22. ^ Singh P, Krishna A, Tsutsui K (March 2011). "Effects of gonadotropin-inhibitory hormone on folliculogenesis and steroidogenesis of cyclic mice". Fertility and Sterility. 95 (4): 1397–404. doi:10.1016/j.fertnstert.2010.03.052. PMID 20452585.
  23. ^ Kirby ED, Geraghty AC, Ubuka T, Bentley GE, Kaufer D (July 2009). "Stress increases putative gonadotropin inhibitory hormone and decreases luteinizing hormone in male rats". Proceedings of the National Academy of Sciences of the United States of America. 106 (27): 11324–9. Bibcode:2009PNAS..10611324K. doi:10.1073/pnas.0901176106. PMC 2698887. PMID 19541621.
  24. ^ Tsutsui K, Bentley GE, Bedecarrats G, Osugi T, Ubuka T, Kriegsfeld LJ (July 2010). "Gonadotropin-inhibitory hormone (GnIH) and its control of central and peripheral reproductive function". Frontiers in Neuroendocrinology. 31 (3): 284–95. doi:10.1016/j.yfrne.2010.03.001. PMID 20211640. S2CID 10120758.
  25. ^ Kiyohara M, Son YL, Tsutsui K (April 2017). "Involvement of gonadotropin-inhibitory hormone in pubertal disorders induced by thyroid status". Scientific Reports. 7 (1): 1042. Bibcode:2017NatSR...7.1042K. doi:10.1038/s41598-017-01183-8. PMC 5430760. PMID 28432332.
  26. ^ Shinomiya A, Shimmura T, Nishiwaki-Ohkawa T, Yoshimura T (2014). "Regulation of seasonal reproduction by hypothalamic activation of thyroid hormone". Frontiers in Endocrinology. 5: 12. doi:10.3389/fendo.2014.00012. PMC 3930870. PMID 24600435.

gonadotropin, inhibitory, hormone, gnih, rfamide, related, peptide, coded, npvf, gene, mammals, contents, discovery, structure, production, receptor, action, effects, physiological, function, additional, biological, roles, referencesdiscovery, editgnih, discov. Gonadotropin inhibitory hormone GnIH is a RFamide related peptide coded by the NPVF gene in mammals Contents 1 Discovery 2 Structure 3 Production 4 Receptor action 5 Effects and physiological function 6 Additional biological roles 7 ReferencesDiscovery editGnIH was discovered in 2000 It is an RFamide peptide that significantly reduced luteinizing hormone release in Coturnix Japonica Japanese quail This peptide emerged as the first tropic hormone known to inhibit gonadotropin secretion in the hypothalamic pituitary gonadal axis of vertebrates 1 Subsequent research identified GnIH peptide homologs in variety of mammals including humans 2 Structure editGnIH is a neurohormone classified as an RFamide RFa or RFamide related peptide RFRP coded by the NPVF gene in mammals The complete amino acid sequence varies by species but all RFa and RFRP peptides contain an arginine phenylalanine amine sequence at the C terminal This is seen in both Coturnix Japonica GnIH RFa Ser Ile Lys Pro Ser Ala Tyr Leu Pro Leu Arg Phe NH2 and the human homolog RFRP 3 Val Pro Asp Leu Pro Glu Arg Phe NH2 1 3 Production editGnIH neurons reside primarily in the dorsomedial nucleus of the hypothalamus humans and rodents and the paraventricular nucleus of the hypothalamus avian species Some GnIH neuron terminals in both mammalian and avian species project to the median eminence 4 5 6 GnIH and GnRH gonadotropin releasing hormone neurons exist in close proximity in the hypothalamus which may enable the direct inhibition of GnRH neurons by GnIH 7 GnIH enters the bloodstream via the hypothalamo hypophyseal portal system the vascular network supplying both the hypothalamus and the pituitary 8 GnIH and GnIH receptor GnIH R mRNA is expressed in the hypothalamus pituitary and ovaries 5 GnIH expression is highest during proestrus and lowest during estrus suggesting the estrus cycle influences release of the hormone Furthermore GnIH neuronal cell counts in multiple vertebrates fluctuate with an organism s parental status 9 GnIH cell count may also vary with breeding season in some species For instance European starlings Sturnus vulgaris with greater reproductive success exhibited higher quantities of GnIH producing cells than did those that were less successful but this effect did not appear until mid breeding season 9 10 11 Receptor action editGnIH binds to the Gai protein coupled receptor GPR147 to suppress adenylyl cyclase formation of cAMP and inhibit protein kinase cascades affecting gene expression GnIH inhibits the same signaling pathway that GnRH activates to promote follicle stimulating hormone FSH and luteinizing hormone LH expression 12 13 The compound RF9 is a known GPR147 receptor antagonist 14 Effects and physiological function editGnIH R expression in the pituitary and other brain regions implies GnIH acts directly on the pituitary to downregulate gonadotropin production impacting reproductive behaviors 6 15 16 17 This neurohormone also acts on the hypothalamus to inhibit the expression of GnRH which may further inhibit gonadotropin secretion and kisspeptin which may inhibit kisspeptin mediated stimulation of GnRH neurons prior to the preovulatory hormonal surge GnIH also spurs the production of cytochrome P450 aromatase promoting the synthesis of neuroestrogen in the brains of quails and reducing aggressivity in reproductive behaviors 7 18 19 In male vertebrates GnIH reduces testis size lowers testosterone secretion and increases the incidence of apoptosis in germ cells and Sertoli cells of the seminiferous tubules 20 21 These gonadal changes in addition to GnIH and GnIH R mRNA expression in the seminiferous tubules Sertoli cells and spermatogonia implicate function in spermatogenesis In female vertebrates high doses of GnIH increases ovarian mass and produce follicle irregularities such as vacuole formation in nuclei and distorted morphology 22 Ovarian changes in response to GnIH administration as well as GnIH GnIH R mRNA expression in granulosa cells and luteal cells in different stages of the estrus cycle implicate function in development of follicles and atresia 21 Additional biological roles editStress induced adrenal hormone increase may upregulate GnIH release as some GnIH neurons have adrenal glucocorticoid receptors GnIH may therefore mediate interactions between the HPG and HPA hypothalamic pituitary adrenal axes and play a role in stress related infertility 23 GnIH neurons of the paraventricular nucleus in the hypothalamus also express melatonin receptors Because melatonin secretion is modulated by environmental light patterns melatonin influence on GnIH production may enable photoperiodic regulation of reproduction in seasonally breeding birds rodents and sheep 24 GnIH increases food consumption implying a role in appetite This finding is consistent with the location of most GnIH neurons as the dorsomedial nucleus of the hypothalamus is involved in appetite regulation GnIH may allow the energy reserves of an organism to modulate reproduction 6 Higher levels of thyroid hormone suppress GnIH expression and lower levels of thyroid hormone are associated with higher GnIH levels The inactivation of GnIH expression prevents delayed puberty caused by hypothyroidism demonstrating that GnIH mediates interactions between the HPG and HPT hypothalamic pituitary thyroid axes 25 Furthermore thyroid hormone may function in a pathway for photoperiodic regulation of reproduction involving GnIH and energy status Melatonin modulates thyroid stimulating hormone TSH production in the anterior pituitary and TSH promotes thyroid hormone production Thyroid hormone production influences metabolism and GnIH production both of which impact reproduction 26 References edit a b Tsutsui K Saigoh E Ukena K Teranishi H Fujisawa Y Kikuchi M Ishii S Sharp PJ August 2000 A novel avian hypothalamic peptide inhibiting gonadotropin release Biochemical and Biophysical Research Communications 275 2 661 7 doi 10 1006 bbrc 2000 3350 PMID 10964719 Fukusumi S Habata Y Yoshida H Iijima N Kawamata Y Hosoya M Fujii R Hinuma S Kitada C Shintani Y Suenaga M Onda H Nishimura O Tanaka M Ibata Y Fujino M September 2001 Characteristics and distribution of endogenous RFamide related peptide 1 Biochimica et Biophysica Acta BBA Molecular Cell Research 1540 3 221 32 doi 10 1016 S0167 4889 01 00135 5 PMID 11583817 Findeisen M Rathmann D Beck Sickinger AG 2011 RFamide Peptides Structure Function Mechanisms and Pharmaceutical Potential Pharmaceuticals 4 9 1248 1280 doi 10 3390 ph4091248 PMC 4058657 Kriegsfeld LJ Mei DF Bentley GE Ubuka T Mason AO Inoue K Ukena K Tsutsui K Silver R February 2006 Identification and characterization of a gonadotropin inhibitory system in the brains of mammals Proceedings of the National Academy of Sciences of the United States of America 103 7 2410 5 Bibcode 2006PNAS 103 2410K doi 10 1073 pnas 0511003103 PMC 1413747 PMID 16467147 a b Ubuka T Morgan K Pawson AJ Osugi T Chowdhury VS Minakata H Tsutsui K Millar RP Bentley GE December 2009 Identification of human GnIH homologs RFRP 1 and RFRP 3 and the cognate receptor GPR147 in the human hypothalamic pituitary axis PLOS ONE 4 12 e8400 Bibcode 2009PLoSO 4 8400U doi 10 1371 journal pone 0008400 PMC 2791420 PMID 20027225 a b c Smith JT Clarke IJ April 2010 Gonadotropin inhibitory hormone function in mammals Trends in Endocrinology and Metabolism 21 4 255 60 doi 10 1016 j tem 2009 11 010 PMID 20060314 S2CID 41737276 a b Smith JT Shahab M Pereira A Pau KY Clarke IJ October 2010 Hypothalamic expression of KISS1 and gonadotropin inhibitory hormone genes during the menstrual cycle of a non human primate Biology of Reproduction 83 4 568 77 doi 10 1095 biolreprod 110 085407 PMC 2957156 PMID 20574054 Smith JT Young IR Veldhuis JD Clarke IJ July 2012 Gonadotropin inhibitory hormone GnIH secretion into the ovine hypophyseal portal system Endocrinology 153 7 3368 75 doi 10 1210 en 2012 1088 PMC 3380300 PMID 22549225 a b Li X Su J Lei Z Zhao Y Jin M Fang R Zheng L Jiao Y August 2012 Gonadotropin inhibitory hormone GnIH and its receptor in the female pig cDNA cloning expression in tissues and expression pattern in the reproductive axis during the estrous cycle Peptides 36 2 176 85 doi 10 1016 j peptides 2012 05 008 PMID 22664321 S2CID 902250 Calisi RM Diaz Munoz SL Wingfield JC Bentley GE July 2011 Social and breeding status are associated with the expression of GnIH Genes Brain and Behavior 10 5 557 64 doi 10 1111 j 1601 183X 2011 00693 x PMID 21466656 Calisi RM Geraghty AC Avila A Kaufer D Bentley GE Wingfield JC October 2016 Patterns of hypothalamic GnIH change over the reproductive period in starlings and rats General and Comparative Endocrinology 237 140 146 doi 10 1016 j ygcen 2016 08 015 PMID 27591072 Ubuka T Son YL Tobari Y Tsutsui K 2012 Gonadotropin inhibitory hormone action in the brain and pituitary Frontiers in Endocrinology 3 148 doi 10 3389 fendo 2012 00148 PMC 3515997 PMID 23233850 Ubuka T Son YL Bentley GE Millar RP Tsutsui K September 2013 Gonadotropin inhibitory hormone GnIH GnIH receptor and cell signaling General and Comparative Endocrinology 10th International Symposium on Avian Endocrinology 190 10 7 doi 10 1016 j ygcen 2013 02 030 PMID 23499786 Caraty A Blomenrohr M Vogel GM Lomet D Briant C Beltramo M May 2012 RF9 powerfully stimulates gonadotrophin secretion in the ewe evidence for a seasonal threshold of sensitivity Journal of Neuroendocrinology 24 5 725 36 doi 10 1111 j 1365 2826 2012 02283 x PMID 22283564 S2CID 25770293 Ubuka T Ukena K Sharp PJ Bentley GE Tsutsui K March 2006 Gonadotropin inhibitory hormone inhibits gonadal development and maintenance by decreasing gonadotropin synthesis and release in male quail Endocrinology 147 3 1187 94 doi 10 1210 en 2005 1178 PMID 16293662 Tsutsui K Bentley GE Ubuka T Saigoh E Yin H Osugi T Inoue K Chowdhury VS Ukena K Ciccone N Sharp PJ Wingfield JC 2007 08 01 The general and comparative biology of gonadotropin inhibitory hormone GnIH General and Comparative Endocrinology Proceedings of the 23rd Conference of European Comparative Endocrinologists Part 2 153 1 3 365 70 doi 10 1016 j ygcen 2006 10 005 PMID 17141777 Bentley GE Jensen JP Kaur GJ Wacker DW Tsutsui K Wingfield JC April 2006 Rapid inhibition of female sexual behavior by gonadotropin inhibitory hormone GnIH Hormones and Behavior 49 4 550 5 doi 10 1016 j yhbeh 2005 12 005 PMID 16460739 S2CID 1801090 Paullada Salmeron JA Cowan M Aliaga Guerrero M Morano F Zanuy S Munoz Cueto JA June 2016 Gonadotropin Inhibitory Hormone Down Regulates the Brain Pituitary Reproductive Axis of Male European Sea Bass Dicentrarchus labrax Biology of Reproduction 94 6 121 doi 10 1095 biolreprod 116 139022 PMC 6322450 PMID 26984999 Tsutsui K Ubuka T 2016 GnIH Control of Feeding and Reproductive Behaviors Frontiers in Endocrinology 7 170 doi 10 3389 fendo 2016 00170 PMC 5186799 PMID 28082949 Ubuka T Ukena K Sharp P Bentley G Tsutsui K Gonadotropin inhibitory hormone inhibits gonadal development and maintenance by decreasing gonadotropin synthesis and release in male quail Endocrinology 2006 147 1187 1194 doi 10 1210 en 2005 1178 a b Tsutsui K Ubuka T Bentley GE Kriegsfeld LJ July 2012 Gonadotropin inhibitory hormone GnIH discovery progress and prospect General and Comparative Endocrinology Profiles in Comparative Endocrinology Eric Roubos 177 3 305 14 doi 10 1016 j ygcen 2012 02 013 PMC 3378827 PMID 22391238 Singh P Krishna A Tsutsui K March 2011 Effects of gonadotropin inhibitory hormone on folliculogenesis and steroidogenesis of cyclic mice Fertility and Sterility 95 4 1397 404 doi 10 1016 j fertnstert 2010 03 052 PMID 20452585 Kirby ED Geraghty AC Ubuka T Bentley GE Kaufer D July 2009 Stress increases putative gonadotropin inhibitory hormone and decreases luteinizing hormone in male rats Proceedings of the National Academy of Sciences of the United States of America 106 27 11324 9 Bibcode 2009PNAS 10611324K doi 10 1073 pnas 0901176106 PMC 2698887 PMID 19541621 Tsutsui K Bentley GE Bedecarrats G Osugi T Ubuka T Kriegsfeld LJ July 2010 Gonadotropin inhibitory hormone GnIH and its control of central and peripheral reproductive function Frontiers in Neuroendocrinology 31 3 284 95 doi 10 1016 j yfrne 2010 03 001 PMID 20211640 S2CID 10120758 Kiyohara M Son YL Tsutsui K April 2017 Involvement of gonadotropin inhibitory hormone in pubertal disorders induced by thyroid status Scientific Reports 7 1 1042 Bibcode 2017NatSR 7 1042K doi 10 1038 s41598 017 01183 8 PMC 5430760 PMID 28432332 Shinomiya A Shimmura T Nishiwaki Ohkawa T Yoshimura T 2014 Regulation of seasonal reproduction by hypothalamic activation of thyroid hormone Frontiers in Endocrinology 5 12 doi 10 3389 fendo 2014 00012 PMC 3930870 PMID 24600435 Retrieved from https en wikipedia org w index php title Gonadotropin inhibitory hormone amp oldid 1105594518, wikipedia, wiki, book, books, library,

article

, read, download, free, free download, mp3, video, mp4, 3gp, jpg, jpeg, gif, png, picture, music, song, movie, book, game, games.