O neuropeptídeo kisspeptina e a reprodução animal: uma revisão
DOI:
https://doi.org/10.33837/msj.v1i2.76Resumo
A kisspeptina (Kp) é um neuropeptídeo que tem uma importante ação na regulação da fertilidade de mamíferos, devido a sua potente ação estimulatória sobre a secreção dos hormônios luteinizante (LH) e folículo estimulante (FSH). O objetivo com esta revisão é abordar a síntese da Kisspeptina e a sua sinalização via receptor GPR54, assim como a sua localização no SNC, funções fisiológicas e mecanismos de ação na fisiologia reprodutiva de mamíferos.Referências
ADACHI, S. et al. Involvement of anteroventral periventricular metastin/kisspeptin neurons in estrogen positive feedback action on luteinizing hormone release in female rats. The Journal of Reproduction and Development, v. 53, n. 2, p. 367–378, 2007.
ANDERSON, G. M. et al. Evidence that thyroid hormones act in the ventromedial preoptic area and the premammillary region of the brain to allow the termination of the breeding season in the ewe. Endocrinology, v. 144, n. 7, p. 2892–2901, 2003.
ANSEL, L. et al. Differential regulation of kiss1 expression by melatonin and gonadal hormones in male and female Syrian hamsters. Journal of Biological Rhythms, v. 25, n. 2, p. 81–91, 2010.
ANSEL, L. et al. Peripheral kisspeptin reverses short photoperiod-induced gonadal regression in Syrian hamsters by promoting GNRH release. Reproduction, v. 142, n. 3, p. 417–425, 2011.
BACKHOLER, K.; SMITH, J.; CLARKE, I. J. Melanocortins may stimulate reproduction by activating orexin neurons in the dorsomedial hypothalamus and kisspeptin neurons in the preoptic area of the ewe. Endocrinology, v. 150, n. 12, p. 5488–5497, 2009.
BACKHOLER, K. et al. Kisspeptin cells in the ewe brain respond to leptin and communicate with neuropeptide Y and proopiomelanocortin cells. Endocrinology, v. 151, n. 5, p. 2233–2243, 2010.
BALASCH, J. et al. Ovarian luteinizing hormone priming preceding follicle-stimulating hormone stimulation: clinical and endocrine effects in women with long-term hypogonadotropic hypogonadism. The Journal of Clinical Endocrinology and Metabolism, v. 94, n. 7, p. 2367–2373, 2009.
BENTLEY, G. E. et al. Gonadotropin-inhibitory hormone and its receptor in the avian reproductive system. General and Comparative Endocrinology, v. 156, n. 1, p. 34–43, 2008.
BIANCO, S. D. C. et al. KISS1R intracellular trafficking and degradation: effect of the Arg386Pro disease-associated Mutation. Endocrinology, v. 152, n. 4, p. 1616–1626, 2011.
BILBAN, M. et al. Kisspeptin-10, a Kiss-1/metastin-derived decapeptide, is a physiological invasion inhibitor of primary human trophoblasts. Journal of Cell Science, v. 11, n. 8, p. 1319–1328, 2004.
BURGER, L. L. et al. GnRH pulse frequency modulation of gonadotropin subunit gene transcription in normal gonadotropes-assessment by primary transcript assay provides evidence for roles of GnRH and follistatin. Endocrinology, v. 143, n. 9, p. 3243–3249, 2002.
CARATY, A. et al. Kisspeptin synchronizes preovulatory surges in cyclical ewes and causes ovulation in seasonally acyclic ewes. Endocrinology, v. 148, n. 11, p. 5258–5267, 2007.
CASANUEVA, F. F.; DIEGUEZ, C. Neuroendocrine regulation and actions of leptin. Frontiers in Neuroendocrinology, v. 20, n. 4, p. 317–63, 1999.
CASTELLANO, J. M. et al. Changes in hypothalamic KiSS-1 system and restoration of pubertal activation of the reproductive axis by kisspeptin in undernutrition. Endocrinology, v. 146, p. 3917–3925, 2005.
CASTELLANO, J. M. et al. Ontogeny and mechanisms of action for the stimulatory effect of kisspeptin on gonadotropin-releasing hormone system of the rat. Molecular and Cellular Endocrinology, v. 257-258, p. 75–83, 2006a.
CASTELLANO, J. M. et al. Expression of KiSS-1 in rat ovary: putative local regulator of ovulation? Endocrinology, v. 147, n. 10, p. 4852–4862, 2006b.
CHAN, Y. M. et al. Kisspeptin resets the hypothalamic GnRH clock in men. The Journal of Clinical Endocrinology and Metabolism, v. 96, n. 6, p. E908–915, 2011.
CHRISTIAN, C. A.; MOENTER, S. M. The neurobiology of preovulatory and estradiol-induced gonadotropin-releasing hormone surges. Endocrine Reviews, v. 31, n. 4, p. 544–577, 2010.
CLARKE, I. J. et al. Potent action of RFamide-related peptide-3 on pituitary gonadotropes indicative of a hypophysiotropic role in the negative regulation of gonadotropin secretion. Endocrinology, v. 149, n. 11, p. 5811–5821, 2008.
CLARKSON, J.; HERBISON, A. E. Postnatal development of kisspeptin neurons in mouse hypothalamus; sexual dimorphism and projections to gonadotropin-releasing hormone neurons. Endocrinology, v. 147, n. 12, p. 5817–5825, 2006.
CLARKSON, J. et al. Distribution of kisspeptin neurones in the adult female mouse brain. Journal of Neuroendocrinology, v. 21, n. 8, p. 6736–82, 2009.
DAHL, G. E. et al. The thyroid gland is required for reproductive neuroendocrine responses to photoperiod in the ewe. Endocrinology, v.135, p.10-15, 1994.
DAHL, G. E. et al. Thyroxine is permissive to seasonal transitions in reproductive neuroendocrine activity in the ewe. Biology of Reproduction, v. 52, n. 3, p. 690–696, 1995.
DARDENTE, H. et al. RFamide-related peptide and its cognate receptor in the sheep: cDNA cloning, mRNA distribution in the hypothalamus and the effect of photoperiod. Journal of Neuroendocrinology, v. 20, p.1252-1259, 2008.
DHILLO, W. S. et al. Kisspeptin-54 stimulates the hypothalamic-pituitary gonadal axis in human males. The Journal of Clinical Endocrinology and Metabolism, v. 90, n. 12, p. 6609–6615, 2005.
DUNGAN, H. M.; CLIFTON, D. K.; STEINER, R. A. Minireview: kisspeptin neurons as central processors in the regulation of gonadotropin-releasing hormone secretion. Endocrinology, v. 147, n. 3, p. 1154–1158, 2006.
EGHLIDI, D. H. et al. Influence of age and 17 beta-estradiol on kisspeptin, neurokinin B, and prodynorphin gene expression in the arcuate-median eminence of female rhesus macaques. Endocrinology, v. 151, n. 8, p. 3783–3794, 2010.
ESTRADA, K. M. et al. Neuropeptide Y (NPY) delays the oestrogen-induced luteinizing hormone (LH) surge in the ovariectomized ewe: further evidence that NPY has a predominant negative effect on LH secretion in the ewe. Journal Neuroendocrinol, v.15, p. 1011–1020, 2003.
FERNANDEZ-FERNANDEZ, R. et al. Novel signals for the integration of energy balance and reproduction. Molecular and Cellular Endocrinology, v. 254-255, p. 127–32, 2006.
FORBES, S. et al. Effects of ghrelin on Kisspeptin mRNA expression in the hypothalamic medial preoptic area and pulsatile luteinising hormone secretion in the female rat. Neuroscience Letters, v. 460, n. 2, p. 143–147, 2009.
FRANCESCHINI, I. et al. Kisspeptin immunoreactive cells of the ovine preoptic area and arcuate nucleus co-express estrogen receptor alpha. Neuroscience Letters, v. 401, n. 3, p. 225–230, 2006.
GAYTÁN, F. et al. KiSS-1 in the mammalian ovary: distribution of kisspeptin in human and marmoset and alterations in KiSS-1 mRNA levels in a rat model of ovulatory dysfunction. American Journal of Physiology Endocrinology and Metabolism, v. 296, n. 3, p. E520–E531, 2009.
GEORGE, J. T. et al. Kisspeptin-10 is a potent stimulator of LH and increases pulse frequency in men. The Journal of Clinical Endocrinology and Metabolism, v. 96, n. 8, p. E1228–E1236, 2011.
GOLDMAN, B. D. Mammalian Photoperiodic System: formal properties and neuroendocrine mechanisms of photoperiodic time measurement. Journal of Biological Rhythms, v. 16, n. 4, p. 283–301, 2001.
GOODMAN, R. L. et al. Kisspeptin neurons in the arcuate nucleus of the ewe express both dynorphin A and neurokinin B. Endocrinology, v. 148, n. 12, p. 5752–5760, 2007.
GOODMAN, R. L. et al. Neural systems mediating seasonal breeding in the ewe. Journal of Neuroendocrinology, v. 22, p. 674–81, 2010.
GOODMAN, R. L. et al. Neural systems mediating seasonal breeding in the ewe. Journal of Neuroendocrinology, v. 22, n. 7, p. 674–681, 2011.
GOTTSCH, M. L. et al. A role for kisspeptins in the regulation of gonadotropin secretion in the mouse. Endocrinology, v. 145, n. 9, p. 4073–4077, 2004.
GREIVES, T. J. et al. Photoperiod and testosterone interact to drive seasonal changes in kisspeptin expression in Siberian hamsters (Phodopus sungorus). Journal of Neuroendocrinology, v. 20, n. 12, p. 1339–1347, 2008.
GUTIÉRREZ-PASCUAL, E. et al. Direct pituitary effects of kisspeptin: activation of gonadotrophs and somatotrophs and stimulation of luteinising hormone and growth hormone secretion. Journal of Neuroendocrinology, v. 19, n. 7, p. 521–530, 2007.
HAN, S. K. et al. Activation of Gonadotropin-Releasing Hormone Neurons by Kisspeptin as a Neuroendocrine Switch for the Onset of Puberty. The Journal of Neuroscience, v. 25, n. 49, p. 11349–11356, 2005.
HASHIZUME, T. et al. Characteristics of stimulation of gonadotropin secretion by kisspeptin-10 in female goats. Animal Reproduction Science, v. 118, p. 37–41, 2010.
HAVERN, R. L.; WHISNANT, C. S.; GOODMAN, R. L. Hypothalamic sites of catecholamine inhibition of luteinizing hormone in the anestrous ewe. Biology of Reproduction, v. 44, n. 3, p. 476–482, 1991.
HERBISON, A. E.; THEODOSIS, D. T. Localization of oestrogen receptors in preoptic neurons containing neurotensin but not tyrosine hydroxylase , cholecystokinin or luteinizing hormone-releasing hormone in the male and female rat. Neuroscience, v. 50, n. 2, p. 283–298, 1992.
HERBISON, A. E. et al. Distribution and postnatal development of Gpr54 gene expression in mouse brain and gonadotropin-releasing hormone neurons. Endocrinology, v. 151, n. 1, p. 312–321, 2010.
HERBISON, A. E.; MOENTER, S. M. Depolarising and hyperpolarising actions of GABA receptor activation on GnRH neurons: towards an emerging consensus. Journal of Neuroendocrinology, v. 23, n. 7, p. 557–569, 2011.
HILL, J. W.; ELMQUIST, J. K.; ELIAS, C. F. Hypothalamic pathways linking energy balance and reproduction. American Journal of Physiology Endocrinology and Metabolism, v. 294, n. 5, p. E827–E832, 2008.
HINEY, J. K. et al. Insulin-like growth factor-I activates KiSS-1 gene expression in the brain of the prepubertal female rat. Endocrinology, v. 150, n. 1, p. 376–384, 2009.
HOFFMAN, G. E. et al. Expression of fos and in vivo median eminence release of LHRH identifies an active role for preoptic area kisspeptin neurons in synchronized surges of LH and LHRH in the ewe. Endocrinology, v. 152, n. 1, p. 214–222, 2011.
HRABOVSZKY, E. et al. The kisspeptin system of the human hypothalamus: sexual dimorphism and relationship with gonadotropin-releasing hormone and neurokinin B neurons. The European Journal of Neuroscience, v. 31, n. 11, p. 1984–1998, 2010.
IRWIG, M. S. et al. Kisspeptin activation of gonadotropin releasing hormone neurons and regulation of KiSS-1 mRNA in the male rat. Neuroendocrinology, v. 80, n. 4, p. 264–272, 2004.
IWASA, T. et al. Delayed puberty in prenatally glucocorticoid administered female rats occurs independently of the hypothalamic Kiss1-Kiss1r-GnRH system. International Journal of Developmental Neuroscience, v. 29, n. 2, p. 183–188, 2011.
KIM, G. L.; DHILLON, S. S.; BELSHAM, D. D. Kisspeptin directly regulates neuropeptide y synthesis and secretion via the ERK1/2 and p38 mitogen-activated protein kinase signaling pathways in NPY-secreting hypothalamic neurons. Endocrinology, v. 151, p. 5038–5047, 2010.
KOTANI, M. et al. The metastasis suppressor gene KiSS-1 encodes kisspeptins , the natural ligands of the orphan G protein-coupled receptor GPR54. The Journal of Biological Chemistry, v. 276, p. 34631–34636, 2001.
KRETSER, D. M. DE et al. Inhibins, activins and follistatin in reproduction. Human Reproduction Update, v. 8, n. 6, p. 529–541, 2002.
KRIEGSFELD, L. J. et al. 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, v. 103, n. 7, p. 2410–2415, 2006.
LEE, J. H. et al. KiSS-1, a novel human malignant melanoma metastasis-suppressor gene. Journal of the National Cancer Institute, v. 88, p. 1731–1737, 1996.
LEHMAN, M. N.; COOLEN, L. M.; GOODMAN, R. L. Minireview: kisspeptin/neurokinin B/dynorphin (KNDy) cells of the arcuate nucleus: a central node in the control of gonadotropin-releasing hormone secretion. Endocrinology, v. 151, n. 8, p. 3479–3489, 2010.
LI, Q. et al. Kisspeptin cells in the ovine arcuate nucleus express prolactin receptor but not melatonin receptor. Journal of Neuroendocrinology, v.23, p. 871-882, 2011.
LIU, X.; LEE, K.; HERBISON, A. E. Kisspeptin excites gonadotropin-releasing hormone neurons through a phospholipase C/Calcium-dependent pathway regulating multiple ion channels. Endocrinology, v. 149, n. 9, p. 4605–4614, 2008.
LOPES, R. A. Papel da prolactina na regulação da expressão de kisspeptina e secreção do hormônio luteinizante em fêmeas. 2012. 70 f. Dissertação (Mestrado em Ciências Biológicas) - Universidade Federal de Minas Gerais, 2012.
LUQUE, R. M.; KINEMAN, R. D.; TENA-SEMPERE, M. Regulation of hypothalamic expression of KiSS-1 and GPR54 genes by metabolic factors: analyses using mouse models and a cell line. Endocrinology, v. 148, n. 10, p. 4601–4611, 2007.
MAGEE, C. et al. Biological and anatomical evidence for kisspeptin regulation of the hypothalamic-pituitary-gonadal axis of estrous horse mares. Endocrinology, v. 150, n. 6, p. 2813–2821, 2009.
MESSAGER, S. et al. Kisspeptin directly stimulates gonadotropin-releasing hormone release via G protein-coupled receptor 54. Proceedings of the National Academy of Sciences of the United States of America, v. 102, n. 5, p. 1761–1766, 2005.
MIKKELSEN, J. D. et al. Hypocretin (orexin) in the rat pineal gland: A central transmitter with effects on noradrenaline-induced release of melatoni. European Journal of Neuroscience, v. 14, p. 419-425, 2001.
MIKKELSEN, J. D. et al. Comparison of the effects of peripherally administered kisspeptins. Regulatory Peptides, v. 152, n. 1-3, p. 95–100, 2009.
MOENTER, S. M.; WOODFILL, C. J.; KARSCH, F. J. Role of the thyroid gland in seasonal reproduction: thyroidectomy blocks seasonal suppression of reproductive neuroendocrine activity in ewes. Endocrinology, v. 128, p. 1337-1344, 1991.
MORTON, G. J. et al. Central nervous system control of food intake and body weight. Nature, v. 443, n. 21, p. 289–295, 2006.
MUIR, A. I. et al. AXOR 12, a novel human G protein-coupled receptor, activated by the peptide KISS-1. The Journal of Biological Chemistry, v. 276, p. 28969-28975, 2001.
NAVARRO, V. M. et al. Developmental and hormonally regulated messenger ribonucleic acid expression of KiSS-1 and its putative receptor , GPR54 , in rat hypothalamus and of KiSS-1 peptide. Endocrinology, v. 145, n. 10, p. 4565–4574, 2004.
NAVARRO, V. M. et al. Effects of KiSS-1 peptide, the natural ligand of GPR54, on follicle-stimulating hormone secretion in the rat. Endocrinology, v. 146, n. 4, p. 1689–1697, 2005a.
NAVARRO, V. M. et al. Characterization of the potent luteinizing hormone-releasing activity of KiSS-1 peptide, the natural ligand. Endocrinology, v. 146, n. 1, p. 156–163, 2005b.
NAVARRO, V. M. et al. Regulation of gonadotropin-releasing hormone secretion by kisspeptin/dynorphin/neurokinin B neurons in the arcuate nucleus of the mouse. The Journal of Neuroscience, v. 29, n. 38, p. 11859–11866, 2009.
NICHOLLS, T. J. et al. Possible homologies between photorefractoriness in sheep and birds: the effect of thyroidectomy on the length of the ewe's breeding season. Resultados da pesquisa. Reproduction Nutrition Development, v. 28, p. 375-385, 1988.
OAKLEY, A. E.; CLIFTON, D. K.; STEINER, R. A. Kisspeptin Signaling in the Brain. Endocrine Reviews, v. 30, n. 6, p. 713–743, 2009.
OHTAKI, T. et al. Metastasis suppressor gene KiSS-1 encodes peptide ligand of a G-protein-coupled receptor. Nature, v. 411 (6837), p. 613–617, 2001.
OJEDA, S. R. et al. Minireview: the neuroendocrine regulation of puberty: is the time ripe for a systems biology approach? Endocrinology, v. 147, n. 3, p. 1166–1174, 2006.
OJEDA, S. R.; LOMNICZI, A.; SANDAU, U. S. Glial-gonadotrophin hormone (GnRH) neurone interactions in the median eminence and the control of GnRH secretion. Journal of Neuroendocrinology, v. 20, n. 6, p. 732–742, 2008.
OJEDA, S. R. et al. New concepts on the control of the onset of puberty. Endocrine Development, v. 17, p. 44-51, 2010.
PAMPILLO, M. et al. Regulation of GPR54 signaling by GRK2 and B-arrestin. Molecular Endocrinology, v. 23, n. 12, p. 2060–2074, 2009.
PARKINSON, T. J.; FOLLETT, B. K. Effect of thyroidectomy upon seasonality in rams. Journal of Reproduction and Fertility, v. 101, p. 51-58, 1994.
PATTERSON, M. et al. Administration of kisspeptin-54 into discrete regions of the hypothalamus potently increases plasma luteinising hormone and testosterone in male adult rats. Journal of Neuroendocrinology, v. 18, n. 5, p. 349–354, 2006.
PAUL, M. J. et al. Photic and non-photic seasonal cues differentially engage hypothalamic kisspeptin and RFamide-related peptide mRNA expression in Siberian hamsters. Journal of Neuroendocrinology, v. 21, n. 12, p. 1007–1014, 2010.
PHENG, V. et al. Potencies of centrally- or peripherally-injected full-length kisspeptin or its C-terminal decapeptide on LH release in intact male rats. The Journal of Reproduction and Development, v. 55, n. 4, p. 378–382, 2009.
PIELECKA-FORTUNA, J.; CHU, Z.; MOENTER, S. M. Kisspeptin acts directly and indirectly to increase gonadotropin-releasing hormone neuron activity and its effects are modulated by estradiol. Endocrinology, v. 149, n. 4, p. 1979–1986, 2008.
PIELECKA-FORTUNA, J.; MOENTER, S. M. Kisspeptin increases gamma-aminobutyric acidergic and glutamatergic transmission directly to gonadotropin-releasing hormone neurons in an estradiol-dependent manner. Endocrinology, v. 151, n. 1, p. 291–300, 2010.
PINEDA, R. et al. Critical roles of kisspeptins in female puberty and preovulatory gonadotropin surges as revealed by a novel antagonist. Endocrinology, v. 151, n. 2, p. 722–730, 2010.
PINILLA, L. et al. Kisspeptins and reproduction: physiological roles and regulatory mechanisms. Physiological reviews, v. 92, n. 3, p. 1235–316, 2012.
PINTO, F. M. et al. Characterization of the kisspeptin system in human spermatozoa. International Journal of Andrology, v. 35, n. 1, p. 63–73, 2012.
POMPOLO, S.; RAWSON, J. A.; CLARKE, I. J. Projections from the arcuate/ventromedial region of the hypothalamus to the preoptic area and bed nucleus of stria terminalis in the brain of the ewe; lack of direct input to gonadotropin-releasing hormone neurons. Brain Research, v. 904, p. 1–12, 2001.
PRALONG, F. P. Insulin and NPY pathways and the control of GnRH function and puberty onset. Molecular and Cellular Endocrinology, v. 324, n. 1-2, p. 82–86, 2010.
QUENNELL, J. H. et al. Leptin indirectly regulates gonadotropin-releasing hormone neuronal function. Endocrinology, v. 150, n. 6, p. 2805–2812, 2009.
QUENNELL, J. H. et al. Developmental and steroidogenic effects on the gene expression of RFamide related peptides and their receptor in the rat brain and pituitary gland. Journal of Neuroendocrinology, v. 22, n. 4, p. 309–316, 2010.
RAMASWAMY, S. et al. Effect of continuous intravenous administration of human metastin 45-54 on the neuroendocrine activity of the hypothalamic-pituitary-testicular axis in the adult male rhesus monkey (Macaca mulatta). Endocrinology, v. 148, n. 7, p. 3364–3370, 2007.
RAMASWAMY, S. et al. Structural interactions between kisspeptin and GnRH neurons in the mediobasal hypothalamus of the male rhesus monkey (Macaca mulatta) as revealed by double immunofluorescence and confocal microscopy. Endocrinology, v. 149, n. 9, p. 4387–4395, 2008.
RAMASWAMY, S. et al. Neurokinin B stimulates GnRH release in the male monkey (Macaca mulatta) and is colocalized with kisspeptin in the arcuate nucleus. Endocrinology, v. 151, n. 9, p. 4494–4503, 2010.
RANCE, N. E. et al. Neurokinin B and the hypothalamic regulation of reproduction. Brain Research, v. 1364, p. 116–128, 2010.
REVEL, F. G. et al. Kisspeptin mediates the photoperiodic control of reproduction in hamsters. Current Biology, v. 16, p. 1730–1735, 2006.
REVEL, F. G. et al. RFamide-related peptide gene is a melatonin-driven photoperiodic gene. Endocrinology, v. 149, n. 3, p. 902–912, 2008.
RICHARD, N. et al. KiSS-1 and GPR54 genes are co-expressed in rat gonadotrophs and differentially regulated in vivo by oestradiol and gonadotrophin-releasing hormone. Journal of Neuroendocrinology, v. 20, n. 3, p. 381–393, 2008.
ROA, J. et al. New frontiers in kisspeptin/GPR54 physiology as fundamental gatekeepers of reproductive function. Frontiers in Neuroendocrinology, v. 29, p. 48–69, 2008.
ROA, J.; TENA-SEMPERE, M. Energy balance and puberty onset: emerging role of central mTOR signaling. Trends in Endocrinology and Metabolism, v. 21, n. 9, p. 519–528, 2010.
ROUX, N. DE et al. Hypogonadotropic hypogonadism due to loss of function of the KiSS1-derived peptide receptor GPR54. Proceedings of the National Academy of Sciences of the United States, v. 100, n. 19, p. 10972–10976, 2003.
SARI, I. P. et al. Effect of RF-amide-related peptide-3 on luteinizing hormone and follicle-stimulating hormone synthesis and secretion in ovine pituitary gonadotropes. Endocrinology, v. 150, n. 12, p. 5549–5556, 2009.
SCHNEIDER, J. E. Energy balance and reproduction. Physiology & Behavior, v. 81, n. 2, p. 289–317, 2004.
SCHWARTZ, M. W. et al. Identification of targets of leptin action in rat hypothalamus. The Journal of Clinical Investigation, v. 98, n. 5, p. 1101–1106, 1996.
SCHWARTZ, M. W. et al. Central nervous system control of food intake. Nature, v. 404, p. 661–671, 2000.
SEMINARA, S. B. et al. The GPR54 gene as a regulator of puberty. The New England Journal of Medicine, v. 349, n. 17, p. 1614–1627, 2003.
SEMINARA, S. B. et al. Continuous human metastin 45–54 infusion desensitizes G protein-coupled receptor 54-induced gonadotropin-releasing hormone release monitored indirectly in the juvenile male Rhesus monkey (Macaca mulatta): a finding with therapeutic implications. Endocrinology, v. 147, p. 2122–2126, 2006.
SHIBATA, M. et al. Evidence that down regulation of hypothalamic KiSS-1 expression is involved in the negative feedback action of testosterone to regulate luteinising hormone secretion in the adult male rhesus monkey (Macaca mulatta). Journal of Neuroendocrinology, v. 19, n. 6, p. 432–438, 2007.
SKINNER, D. C.; CARATY, A.; ALLINGHAM, R. Unmasking the progesterone receptor in the preoptic area and hypothalamus of the ewe: no colocalization with gonadotropin-releasing neurons. Endocrinology, v. 142, n. 2, p. 573–579, 2001.
SMITH, J. T. et al. Differential regulation of KiSS-1 mRNA expression by sex steroids in the brain of the male mouse. Endocrinology, v. 146, n. 7, p. 2976–2984, 2005a.
SMITH, J. T. et al. Regulation of Kiss1 gene expression in the brain of the female mouse. Endocrinology, v. 146, n. 9, p. 3686–3692, 2005b.
SMITH, J. T. et al. KiSS-1 neurones are direct targets for leptin in the ob/ob mouse. Journal of Neuroendocrinology, v. 18, n. 4, p. 298–303, 2006a.
SMITH, J. T. et al. Kiss1 neurons in the forebrain as central processors for generating the preovulatory luteinizing hormone surge. The Journal of Neuroscience, v. 26, n. 25, p. 6687–6694, 2006b.
SMITH, J. T. et al. KiSS-1 messenger ribonucleic acid expression in the hypothalamus of the ewe is regulated by sex steroids and season. Endocrinology, v. 148, n. 3, p. 1150–1157, 2007.
SMITH, J. T. et al. Variation in kisspeptin and RFamide-related peptide (RFRP) expression and terminal connections to gonadotropin-releasing hormone neurons in the brain: a novel medium for seasonal breeding in the sheep. Endocrinology, v. 149, n. 11, p. 5770–5782, 2008a.
SMITH, J. T. et al. Kisspeptin is present in ovine hypophysial portal blood but does not increase during the preovulatory luteinizing hormone surge: evidence that gonadotropes are not direct targets of kisspeptin in vivo. Endocrinology, v. 149, n. 4, p. 1951–1959, 2008b.
SMITH, J. T. et al. Sex steroid control of hypothalamic Kiss 1 expression in sheep and rodents: comparative aspects. Peptides, v. 30, n. 1, p. 94-102, 2009a.
SMITH, J. T. et al. Kisspeptin neurons in the ovine arcuate nucleus and preoptic area are involved in the preovulatory luteinizing hormone surge. Endocrine Reviews, v. 150, n. 12, p. 5530–5538, 2009b.
SMITH, J. T.; CLARKE, I. J. Gonadotropin inhibitory hormone function in mammals. Trends in Endocrinology and Metabolism, v. 21, n. 4, p. 255–260, 2010a.
SMITH, J. T. et al. Kisspeptin is essential for the full preovulatory LH surge and stimulates GnRH release from the isolated ovine median eminence. Endocrinology, v. 152, n. 3, p. 1001–1012, 2011.
SMITH, J. T. The role of kisspeptin and gonadotropin inhibitory hormone in the seasonal regulation of reproduction in sheep. Domestic Animal Endocrinology, v. 43, n. 2, p. 75–84, 2012.
TASSIGNY, X. A. et al. Kisspeptin can stimulate gonadotropin-releasing hormone (GnRH) release by a direct action at GnRH nerve terminals. Endocrinology, v. 149, n. 8, p. 3926–3932, 2008.
TASSIGNY, X. A.; COLLEDGE, W. H. The role of kisspeptin signaling in reproduction. Physiology, v. 25, n. 4, p. 207-217, 2010.
TENA-SEMPERE, M. GPR54 and kisspeptin in reproduction. Human Reproduction Update, v. 12, n. 5, p. 631–639, 2006.
TENA-SEMPERE, M. Kisspeptin signaling in the brain: recent developments and future challenges. Molecular and Cellular Endocrinology, v. 314, p. 164–169, 2010.
THOMPSON, E. L. et al. Central and peripheral administration of Kisspeptin-10 stimulates the hypothalamic-pituitary-gonadal axis. Journal of Neuroendocrinology, v. 16, n. 10, p. 850–858, 2004.
THRUN, L. A. et al. Effect of thyroidectomy on maintenance of seasonal reproductive suppression in the ewe. Biology of Reproduction, v. 56, n. 4, p. 1035–1040, 1997.
TOVAR, S. et al. Effects of single or repeated intravenous administration of kisspeptin upon dynamic LH secretion in conscious male rats. Endocrinology, v. 147, n. 6, p. 2696–2704, 2006.
UBUKA, T. et al. Gonadotropin-inhibitory hormone neurons interact directly with gonadotropin-releasing hormone-I and -II neurons in European starling brain. Endocrinology, v. 149, n. 1, p. 268–278, 2008.
UBUKA, T. et al. Gonadotropin-inhibitory hormone identification, cDNA cloning, and distribution in rhesus macaque brain. Journal of Comparative Neurology, v. 517, p. 841-855, 2009.
UENOYAMA, Y. et al. Ultrastructural evidence of kisspeptin-gonadotrophin-releasing hormone (GnRH) interaction in the median eminence of female rats: implication of axo-axonal regulation of GnRH release. Journal of Neuroendocrinology, v. 23, n. 10, p. 863–870, 2011.
VAN DER LELY, A. J. et al. Biological, physiological, pathophysiological, and pharmacological aspects of ghrelin. Endocrine Reviews, v. 25, n. 3, p. 426–457, 2004.
WAGNER, G. C. et al. Redefining the limits of day length responsiveness in a seasonal mammal. Endocrinology, v. 149, n. 1, p. 32–39, 2008.
WAKABAYASHI, Y. et al. Neurokinin B and dynorphin A in kisspeptin neurons of the arcuate nucleus participate in generation of periodic oscillation of neural activity driving pulsatile gonadotropin-releasing hormone secretion in the goat. The Journal of neuroscience, v. 30, n. 8, p. 3124–32, 2010.
WINTERMANTEL, T. M. et al. Definition of estrogen receptor pathway critical for estrogen positive feedback to gonadotropin-releasing hormone neurons and fertility. Neuron, v. 52, n. 2, p. 271–280, 2006.
XU, J. et al. Regulation of food intake and gonadotropinreleasing hormone/luteinizing hormone during lactation: role of insul and leptin. Endocrinology, v. 150, n. 9, p. 4231–4240, 2009.
YAMADA, S. et al. Inhibition of metastin (kisspeptin-54)-GPR54 signaling in the arcuate nucleusmedian
eminence region during lactation in rats. Endocrinology, v. 148, n. 5, p. 2226-2232, 2007.
YEO, S. H.; HERBISON, A. E. Projections of arcuate nucleus and rostral periventricular kisspeptin neurons in the adult female mouse brain. Endocrinology, v. 152, n. 6, p. 2387–2399, 2011.
YEN, S. S. et al. Neuroendocrinology of opioid peptides and their role in the control of gonadotropin and prolactin secretion. American Journal of Obstetrics and Gynecology, v. 152, p. 485–493, 1985.
ZHANG, C. et al. Kisspeptin depolarizes gonadotropin-releasing hormone neurons through activation of TRPC-like cationic channels. The Journal of Neuroscience, v. 28, n. 17, p. 4423–4434, 2008.
ZHANG, C. et al. Gamma-aminobutyric acid B receptor mediated inhibition of gonadotropin-releasing hormone neurons is suppressed by kisspeptin-G protein-coupled receptor 54 signaling. Endocrinology, v. 150, n. 5, p. 2388–2394, 2009.
Downloads
Publicado
Como Citar
Edição
Seção
Licença
Autores que publicam nesta revista concordam com os seguintes termos:- Autores mantém os direitos autorais e concedem à revista o direito de primeira publicação, com o trabalho simultaneamente licenciado sob a Licença Creative Commons Attribution que permite o compartilhamento do trabalho com reconhecimento da autoria e publicação inicial nesta revista.
- Autores têm autorização para assumir contratos adicionais separadamente, para distribuição não-exclusiva da versão do trabalho publicada nesta revista (ex.: publicar em repositório institucional ou como capítulo de livro), com reconhecimento de autoria e publicação inicial nesta revista.
- Autores têm permissão e são estimulados a publicar e distribuir seu trabalho online (ex.: em repositórios institucionais ou na sua página pessoal) a qualquer ponto antes ou durante o processo editorial, já que isso pode gerar alterações produtivas, bem como aumentar o impacto e a citação do trabalho publicado (Veja O Efeito do Acesso Livre).