Pituitary leptin-A paracrine regulator of gonadotropes: A review

Open Neuroendocrinology Journal

Volumn 4, Issue SPEC. ISSUE 1, 2011, Pages 25-42

  Akhter, Noor ^a   Crane, Christopher ^a   Childs, Gwen V ^a  

^a UNIVERSITY OF ARKANSAS FOR MEDICAL SCIENCES   (United States)

Author keywords

Gonadotropes; Gonadotropins; Growth hormone; Leptin; Paracrine; Pituitary; Somatotropes

Indexed keywords

GHRELIN; GONADORELIN; GROWTH HORMONE RELEASING FACTOR; LEPTIN; LUTEINIZING HORMONE; NEUROPEPTIDE Y;

ADENOHYPOPHYSIS; CELL DENSITY; CELL FUNCTION; GONADOTROPIN SECRETING CELL; GROWTH HORMONE SECRETING CELL; HORMONE ACTION; HORMONE SYNTHESIS; HUMAN; INTRACELLULAR SIGNALING; NONHUMAN; NUTRITIONAL STATUS; PARACRINE SIGNALING; PRIORITY JOURNAL; PROTEIN DETERMINATION; PROTEIN EXPRESSION; PROTEIN FUNCTION; PROTEIN INTERACTION; PROTEIN LOCALIZATION; PROTEIN SECRETION; QUANTITATIVE ANALYSIS; REPRODUCTION; REVERSE TRANSCRIPTION POLYMERASE CHAIN REACTION; REVIEW;

EID: 80455150338     PISSN: None     EISSN: 18765289     Source Type: Journal    
DOI: 10.2174/1876528901104010025     Document Type: Review

References (189)

1. Zhang Y, Proenca R, Maffei M, Barone M, Leopold L, Friedman JM. Positional cloning of the mouse obese gene and its human homologue. Nature 1994; 372: 425-432.
2. Rowland NE, Morien A, Li BH. The physiology and brain mechanisms of feeding. Nutrition 1996; 12: 626-639.
3. Mizuno T, Bergen H, Kleopoulos S, Bauman WA, Mobbs CV. Effects of nutritional status and aging on leptin gene expression in mice: importance of glucose. Horm Metab Res 1996; 28: 679-684.
4. Baranowska B, Chmielowska M, Wolinska-Witort E, Roguski K, Wasilewska-Dziubinska E. The relationship between neuropeptides and hormones in starvation. Neuro Endocrinol Lett 2001; 22: 349-355.
5. Neary NM, Goldstone AP, Bloom SR. Appetite regulation: from the gut to the hypothalamus. Clin Endocrinol (Oxf) 2004; 60: 153-160.
6. Vasselli JR. Behavioral and biological determinants of leptin resistance. Appetite 2001; 37(2): 115-117.
7. Mizuno TM, Makimura H, Silverstein J, Roberts JL, Lopingco T, Mobbs CV. Fasting regulates hypothalamic neuropeptide Y, agouti-related peptide, and proopiomelanocortin in diabetic mice independent of changes in leptin or insulin. Endocrinology 1999; 140: 4551-4557.
8. Mizuno TM, Mobbs CV. Hypothalamic agouti-related protein messenger ribonucleic acid is inhibited by leptin and stimulated by fasting. Endocrinology 1999; 140(2): 814-817.
9. Chua SC, Jr., Leibel RL, Hirsch J. Food deprivation and age modulate neuropeptide gene expression in the murine hypothalamus and adrenal gland. Brain Res Mol Brain Res 1991 Jan; 9(1/2): 95-101.
10. Ebihara K, Ogawa Y, Katsuura G, et al. Involvement of agoutirelated protein, an endogenous antagonist of hypothalamic melanocortin receptor, in leptin action. Diabetes 1999; 48: 2028-2033.
11. Korner J, Savontaus E, Chua SC, Jr., Leibel RL, Wardlaw SL. Leptin regulation of Agrp and Npy mRNA in the rat hypothalamus. J Neuroendocrinol 2001; 13: 959-966.
12. Mizuno TM, Bergen H, Funabashi T, et al. Obese gene expression: reduction by fasting and stimulation by insulin and glucose in lean mice, and persistent elevation in acquired (diet-induced) and genetic (yellow agouti) obesity. Proc Natl Acad Sci U S A 1996 16; 93: 3434-3438.
13. Fruhbeck G. Intracellular signalling pathways activated by leptin. Biochem J 2006; 393: 7-20.
14. Schneider JE, Blum RM, Wade GN. Metabolic control of food intake and estrous cycles in syrian hamsters. I. Plasma insulin and leptin. Am J Physiol Regul Integr Comp Physiol 2000; 278(2): 476-485.
15. Schneider JE, Buckley CA, Blum RM, et al. Metabolic signals, hormones and neuropeptides involved in control of energy balance and reproductive success in hamsters. Eur J Neurosci 2002; 16(3): 377-379.
16. Schneider JE, Goldman MD, Tang S, Bean B, Ji H, Friedman MI. Leptin indirectly affects estrous cycles by increasing metabolic fuel oxidation. Horm Behav 1998; 33(3): 217-228.
17. Schneider JE, Zhou D. Interactive effects of central leptin and peripheral fuel oxidation on estrous cyclicity. Am J Physiol 1999; 277: 1020-1024.
18. Schneider JE, Wade GN. Availability of metabolic fuels controls estrous cyclicity of Syrian hamsters. Science 1989; 244(4910): 1326-128.
19. Casanueva FF, Dieguez C. Neuroendocrine regulation and actions of leptin. Front Neuroendocrinol 1999; 20(4): 317-363.
20. Ahima RS, Prabakaran D, Mantzoros C, et al. Role of leptin in the neuroendocrine response to fasting. Nature 1996; 382(6588): 250-252.
21. Moschos S, Chan JL, Mantzoros CS. Leptin and reproduction: a review. Fertil Steril 2002; 77(3): 433-444.
22. Ahima RS, Dushay J, Flier SN, Prabakaran D, Flier JS. Leptin accelerates the onset of puberty in normal female mice. J Clin Invest 1997; 99(3): 391-395.
23. Casanueva FF, Dieguez C. Neuroendocrine Regulation and Actions of Leptin. Front Neuroendocrinol 1999; 20(4): 317-363.
24. Chehab FF, Lim ME, Lu R. Correction of the sterility defect in homozygous obese female mice by treatment with the human recombinant leptin. Nat Genet 1996; 12(3): 318-320.
25. Cunningham MJ, Clifton DK, Steiner RA. Leptin's actions on the reproductive axis: perspectives and mechanisms. Biol Reprod 1999; 60(2): 216-222.
26. Mannucci E, Ognibene A, Becorpi A, et al. Relationship between leptin and oestrogens in healthy women. Eur J Endocrinol 1998; 139(2): 198-201.
27. McMinn JE, Liu SM, Liu H, et al. Neuronal deletion of Lepr elicits diabesity in mice without affecting cold tolerance or fertility. Am J Physiol Endocrinol Metab 2005; 289(3): E403-E411.
28. Yura S, Ogawa Y, Sagawa N, et al. Accelerated puberty and lateonset hypothalamic hypogonadism in female transgenic skinny mice overexpressing leptin. J Clin Invest 2000; 105(6): 749-755.
29. Isozaki O, Tsushima T, Miyakawa M, Nozoe Y, Demura H, Seki H. Growth hormone directly inhibits leptin gene expression in visceral fat tissue in fatty Zucker rats. J Endocrinol 1999; 161(3): 511-516.
30. Popovic V, Damjanovic S, Dieguez C, Casanueva FF. Leptin and the pituitary. Pituitary 2001; 4(1/2): 7-14.
31. Lloyd RV, Jin L, Tsumanuma I, et al. Leptin and leptin receptor in anterior pituitary function. Pituitary. 2001; 4(1/2): 33-47.
32. Mann DR, Plant TM. Leptin and pubertal development. Semin Reprod Med 2002; 20(2): 93-102.
33. Urbanski HF. Leptin and puberty. Trends Endocrinol Metab 2001; 12(10): 428-429.
34. Farooqi IS, Jebb SA, Langmack G, et al. Effects of recombinant leptin therapy in a child with congenital leptin deficiency. N Engl J Med 1999; 341(12): 879-884.
35. Montague CT, Farooqi IS, Whitehead JP, et al. Congenital leptin deficiency is associated with severe early-onset obesity in humans. Nature 1997; 387(6636): 903-908.
36. Strobel A, Issad T, Camoin L, Ozata M, Strosberg AD. A leptin issense mutation associated with hypogonadism and morbid obesity. Nat Genet 1998; 18(3): 213-215.
37. Clement K, Vaisse C, Lahlou N, et al. A mutation in the human leptin receptor gene causes obesity and pituitary dysfunction. Nature 1998; 392(6674): 398-401.
38. Kaufman BA, Warren MP, Dominguez JE, Wang J, Heymsfield SB, Pierson RN. Bone density and amenorrhea in ballet dancers are related to a decreased resting metabolic rate and lower leptin levels. J Clin Endocrinol Metab 2002; 87(6): 2777-2783.
39. Laughlin GA, Yen SS. Hypoleptinemia in women athletes: absence of a diurnal rhythm with amenorrhea. J Clin Endocrinol Metab 1997; 82(1): 318-321.
40. Miller KK, Parulekar MS, Schoenfeld E, et al. Decreased leptin levels in normal weight women with hypothalamic amenorrhea: the effects of body composition and nutritional intake. J Clin Endocrinol Metab 1998; 83(7): 2309-2312.
41. Thong FS, Graham TE. Leptin and reproduction: is it a critical link between adipose tissue, nutrition, and reproduction? Can J Appl Physiol 1999; 24(4): 317-336.
42. Thong FS, McLean C, Graham TE. Plasma leptin in female athletes: relationship with body fat, reproductive, nutritional, and endocrine factors. J Appl Physiol 2000; 88(6): 2037-2044.
43. Warren MP, Voussoughian F, Geer EB, Hyle EP, Adberg CL, Ramos RH. Functional hypothalamic amenorrhea: hypoleptinemia and disordered eating. J Clin Endocrinol Metab 1999; 84(3): 873-877.
44. Welt CK. Will leptin become the treatment of choice for functional hypothalamic amenorrhea? Nat Clin Pract Endocrinol Metab 2007; 3(8): 556-557.
45. Welt CK, Chan JL, Bullen J, et al. Recombinant human leptin in women with hypothalamic amenorrhea. N Engl J Med 2004; 351(10): 987-997.
46. Chan JL, Heist K, DePaoli AM, Veldhuis JD, Mantzoros CS. The role of falling leptin levels in the neuroendocrine and metabolic adaptation to short-term starvation in healthy men. J Clin Invest 2003; 111(9): 1409-1421.
47. Barash IA, Cheung CC, Weigle DS, et al. Leptin is a metabolic signal to the reproductive system. Endocrinology 1996; 137(7): 3144-3147.
48. Chehab FF, Mounzih K, Lu R, Lim ME. Early onset of reproductive function in normal female mice treated with leptin. Science 1997; 275(5296): 88-90.
49. Cheung CC, Thornton JE, Kuijper JL, Weigle DS, Clifton DK, Steiner RA. Leptin is a metabolic gate for the onset of puberty in the female rat. Endocrinology 1997; 138(2): 855-858.
50. Bronson FH. Food-restricted, prepubertal, female rats: rapid recovery of luteinizing hormone pulsing with excess food, and full recovery of pubertal development with gonadotropin-releasing hormone. Endocrinology 1986; 118(6): 2483-2487.
51. Bronson FH. Puberty in female mice is not associated with increases in either body fat or leptin. Endocrinology 2001; 142(11): 4758-47561.
52. Cheung CC, Thornton JE, Nurani SD, Clifton DK, Steiner RA. A reassessment of leptin's role in triggering the onset of puberty in the rat and mouse. Neuroendocrinology 2001; 74(1): 12-21.
53. Mann DR, Akinbami MA, Gould KG, Castracane VD. Leptin and thyroxine during sexual development in male monkeys: effect of neonatal gonadotropin-releasing hormone antagonist treatment and delayed puberty on the developmental pattern of leptin and thyroxine secretion. Eur J Endocrinol 2002; 146(6): 891-898.
54. Mann DR, Bhat GK, Ramaswamy S, Stah CD, Plant TM. Regulation of circulating leptin and its soluble receptor during pubertal development in the male rhesus monkey (Macaca mulatta). Endocrine 2007; 31(2): 125-129.
55. Plant TM, Durrant AR. Circulating leptin does not appear to provide a signal for triggering the initiation of puberty in the male rhesus monkey (Macaca mulatta). Endocrinology 1997; 138(10): 4505-4508.
56. Urbanski HF, Pau KY. A biphasic developmental pattern of circulating leptin in the male rhesus macaque (Macaca mulatta). Endocrinology 1998; 139(5): 2284-2286.
57. Zamorano PL, Mahesh VB, De Sevilla LM, Chorich LP, Bhat GK, Brann DW. Expression and localization of the leptin receptor in endocrine and neuroendocrine tissues of the rat. Neuroendocrinology 1997; 65(3): 223-228.
58. Finn PD, Cunningham MJ, Pau KY, Spies HG, Clifton DK, Steiner RA. The stimulatory effect of leptin on the neuroendocrine reproductive axis of the monkey. Endocrinology 1998; 139(11): 4652-4662.
59. Hakansson ML, Brown H, Ghilardi N, Skoda RC, Meister B. Leptin receptor immunoreactivity in chemically defined target neurons of the hypothalamus. J Neurosci 1998; 18(1): 559-572.
60. Magni P, Vettor R, Pagano C, et al. Expression of a leptin receptor in immortalized gonadotropin-releasing hormone-secreting neurons. Endocrinology 1999; 140(4): 1581-1585.
61. Chan JL, Mantzoros CS. Leptin and the hypothalamic-pituitary regulation of the gonadotropin-gonadal axis. Pituitary 2001; 4(1/2): 87-92.
62. Lebrethon MC, Vandersmissen E, Gerard A, Parent AS, Bourguignon JP. Cocaine and amphetamine-regulated-transcript peptide mediation of leptin stimulatory effect on the rat gonadotropin-releasing hormone pulse generator in vitro. J Neuroendocrinol 2000; 12(5): 383-385.
63. Lebrethon MC, Vandersmissen E, Gerard A, Parent AS, Junien JL, Bourguignon JP. In vitro stimulation of the prepubertal rat gonadotropin-releasing hormone pulse generator by leptin and neuropeptide Y through distinct mechanisms. Endocrinology 2000; 141(4): 1464-1499.
64. Nagatani S, Guthikonda P, Thompson RC, Tsukamura H, Maeda KI, Foster DL. Evidence for GnRH regulation by leptin: leptin administration prevents reduced pulsatile LH secretion during fasting. Neuroendocrinology 1998; 67(6): 370-376.
65. Spicer LJ. Leptin: a possible metabolic signal affecting reproduction. Domest Anim Endocrinol 2001; 21(4): 251-270.
66. Watanobe H. Leptin directly acts within the hypothalamus to stimulate gonadotropin-releasing hormone secretion in vivo in rats. J Physiol 2002; 545(Pt 1): 255-268.
67. Gonzalez LC, Pinilla L, Tena-Sempere M, Aguilar E. Leptin(116- 130) stimulates prolactin and luteinizing hormone secretion in fasted adult male rats. Neuroendocrinology 1999; 70(3): 213-220.
68. Nagatani S, Zeng Y, Keisler DH, Foster DL, Jaffe CA. Leptin regulates pulsatile luteinizing hormone and growth hormone secretion in the sheep. Endocrinology 2000; 141(11): 3965-3975.
69. Tezuka M, Irahara M, Ogura K, et al. Effects of leptin on gonadotropin secretion in juvenile female rat pituitary cells. Eur J Endocrinol 2002; 146(2): 261-266.
70. De Biasi SN, Apfelbaum LI, Apfelbaum ME. In vitro effect of leptin on LH release by anterior pituitary glands from female rats at the time of spontaneous and steroid-induced LH surge. Eur J Endocrinol 2001; 145(5): 659-665.
71. Ogura K, Irahara M, Kiyokawa M, et al. Effects of leptin on secretion of LH and FSH from primary cultured female rat pituitary cells. Eur J Endocrinol 2001; 144(6): 653-658.
72. Walczewska A, Yu WH, Karanth S, McCann SM. Estrogen and leptin have differential effects on FSH and LH release in female rats. Proc Soc Exp Biol Med 1999; 222(2): 170-177.
73. Yu WH, Kimura M, Walczewska A, Karanth S, McCann SM. Role of leptin in hypothalamic-pituitary function. Proc Natl Acad Sci USA 1997; 94(3): 1023-1028.
74. Yu WH, Walczewska A, Karanth S, McCann SM. Nitric oxide mediates leptin-induced luteinizing hormone-releasing hormone (LHRH) and LHRH and leptin-induced LH release from the pituitary gland. Endocrinology 1997; 138(11): 5055-5058.
75. Aubert ML, Pierroz DD, Gruaz NM, et al. Metabolic control of sexual function and growth: role of neuropeptide Y and leptin. Mol Cell Endocrinol 1998; 140(1/2): 107-113.
76. Pombo M, Pombo CM, Astorga R, et al. Regulation of growth hormone secretion by signals produced by the adipose tissue. J Endocrinol Invest 1999; 22(5 Suppl): 22-26.
77. Roh SG, Nie GY, Loneragan K, Gertler A, Chen C. Direct modification of somatotrope function by long-term leptin treatment of primary cultured ovine pituitary cells. Endocrinology 2001; 142(12): 5167-51671.
78. Vuagnat BA, Pierroz DD, Lalaoui M, et al. Evidence for a leptinneuropeptide Y axis for the regulation of growth hormone secretion in the rat. Neuroendocrinology 1998; 67(5): 291-300.
79. Chen C, Roh SG, Nie GY, et al. The in vitro effect of leptin on growth hormone secretion from primary cultured ovine somatotrophs. Endocrine 2001; 14(1): 73-78.
80. Luque RM, Huang ZH, Shah B, Mazzone T, Kineman RD. Effects of leptin replacement on hypothalamic-pituitary growth hormone axis function and circulating ghrelin levels in ob/ob mice. Am J Physiol Endocrinol Metab 2007; 292(3): E891-E899.
81. Ahima RS, Prabakaran D, Flier JS. Postnatal leptin surge and regulation of circadian rhythm of leptin by feeding. Implications for energy homeostasis and neuroendocrine function. J Clin Invest 1998; 101(5): 1020-1027.
82. Devaskar SU, Ollesch C, Rajakumar RA, Rajakumar PA. Developmental changes in ob gene expression and circulating leptin peptide concentrations. Biochem Biophys Res Commun 1997; 238(1): 44-47.
83. Clancy B, Kersh B, Hyde J, Darlington RB, Anand KJ, Finlay BL. Web-based method for translating neurodevelopment from laboratory species to humans. Neuroinformatics 2007 Spring; 5(1): 79-94.
84. Terasawa E, Fernandez DL. Neurobiological mechanisms of the onset of puberty in primates. Endocr Rev 2001; 22(1): 111-151.
85. Reitman ML, Bi S, Marcus-Samuels B, Gavrilova O. Leptin and its role in pregnancy and fetal development--an overview. Biochem Soc Trans 2001; 29: 68-72.
86. Ong KK, Ahmed ML, Dunger DB. The role of leptin in human growth and puberty. Acta Paediatr Suppl 1999; 88(433): 95-98.
87. Kaplan SL, Grumbach MM, Shepard TH. The ontogenesis of human fetal hormones. I. Growth hormone and insulin. J Clin Invest 1972; 51(12): 3080-3093.
88. Shimon I, Yan X, Magoffin DA, Friedman TC, Melmed S. Intact leptin receptor is selectively expressed in human fetal pituitary and pituitary adenomas and signals human fetal pituitary growth hormone secretion. J Clin Endocrinol Metab 1998; 83(11): 4059-4064.
89. Castillo RH, Matteri RL, Dumesic DA. Luteinizing hormone synthesis in cultured fetal human pituitary cells exposed to gonadotropin-releasing hormone. J Clin Endocrinol Metab 1992; 75(1): 318-322.
90. Kaplan SL, Grumbach MM. The ontogenesis of human foetal hormones. II. Luteinizing hormone (LH) and follicle stimulating hormone (FSH). Acta Endocrinol (Copenh) 1976; 81(4): 808-829.
91. Li JY, Claustrat B, Begeot M, Dubois PM. Human fetal antehypophysis in vitro. II. Immunocytological study and radioimmunoassay of LH and FSH (author's transl). Ann Endocrinol (Paris). 1977; 38(6): 389-390.
92. Siler-Khodr TM, Khodr GS. Studies in human fetal endocrinology: II. LH and FSH content and concentration in the pituitary. Obstet Gynecol 1980; 56(2): 176-181.
93. Akhter N, Johnson BW, Crane C, et al. Anterior pituitary leptin expression changes in different reproductive states: stimulation, In vitro, by gonadotropin releasing hormone (GnRH). J Histochem Cytochem 2007; 55: 151-166.
94. Bronson FH, Heideman PD. Short-term hormonal responses to food intake in peripubertal female rats. Am J Physiol 1990; 259: R25-R31.
95. Cameron JL. Regulation of reproductive hormone secretion in primates by short-term changes in nutrition. Rev Reprod 1996; 1(2): 117-1126.
96. Jin L, Burguera BG, Couce ME, et al. Leptin and leptin receptor expression in normal and neoplastic human pituitary: evidence of a regulatory role for leptin on pituitary cell proliferation. J Clin Endocrinol Metab 1999; 84(8): 2903-2911.
97. Jin L, Zhang S, Burguera BG, et al. Leptin and leptin receptor expression in rat and mouse pituitary cells. Endocrinology 2000; 141(1): 333-9.
98. McDuffie IA, Akhter N, Childs GV. Regulation of leptin Mrna and protein expression in pituitary somatotropes. J Histochem Cytochem 2004; 52(2): 263-273.
99. Sone M, Nagata H, Takekoshi S, Osamura RY. Expression and localization of leptin receptor in the normal rat pituitary gland. Cell Tissue Res 2001; 305(3): 351-356.
100. Sone M, Osamura RY. Leptin and the pituitary. Pituitary 2001; 4(1/2): 15-23.
101. Vidal S, Cohen SM, Horvath E, et al. Subcellular localization of leptin in non-tumorous and adenomatous human pituitaries: an immuno-ultrastructural study. J Histochem Cytochem 2000 Aug; 48(8): 1147-1152.
102. Korbonits M, Chitnis MM, Gueorguiev M, et al. Leptin in pituitary adenomas--a novel paracrine regulatory system. Pituitary 2001; 4(1/2): 49-55.
103. Korbonits M, Chitnis MM, Gueorguiev M, et al. The release of leptin and its effect on hormone release from human pituitary adenomas. Clin Endocrinol (Oxf) 2001; 54(6): 781-789.
104. Scarpace PJ, Matheny M, Zhang Y, Cheng KY, Tumer N. Leptin antagonist reveals an uncoupling between leptin receptor STAT3 signaling and metabolic responses with central leptin resistance. J Pharmacol Exp Ther 2007; 320(2): 706-712.
105. Zhang J, Matheny MK, Tumer N, Mitchell MK, Scarpace PJ. Leptin antagonist reveals that the normalization of caloric intake and the thermic effect of food following high-fat feeding are leptin dependent. Am J Physiol Regul Integr Comp Physiol 2007; 292(2): R868-R874.
106. Iruthayanathan M, Zhou YH, Childs GV. Dehydroepiandrosterone restoration of growth hormone gene expression in aging female rats, in vivo and in vitro: evidence for actions via estrogen receptors. Endocrinology 2005; 146(12): 5176-5187.
107. Schulz LC, Widmaier EP. Leptin receptors. In: Castracane VD, Henson MC, Eds. Leptin. New York, NY: Springer Science; 2006; Chp. 2: 11-33.
108. Baumann H, Morella KK, White DW, et al. The full-length leptin receptor has signaling capabilities of interleukin 6-type cytokine receptors. Proc Natl Acad Sci U S A. 1996; 93(16): 8374-8378.
109. Chen H, Charlat O, Tartaglia LA, et al. Evidence that the diabetes gene encodes the leptin receptor: identification of a mutation in the leptin receptor gene in db/db mice. Cell 1996; 84(3): 491-495.
110. Ghilardi N, Ziegler S, Wiestner A, Stoffel R, Heim MH, Skoda RC. Defective STAT signaling by the leptin receptor in diabetic mice. Proc Natl Acad Sci U S A 1996; 93(13): 6231-5.
111. Vaisse C, Halaas JL, Horvath CM, Darnell JE, Jr., Stoffel M, Friedman JM. Leptin activation of Stat3 in the hypothalamus of wild-type and ob/ob mice but not db/db mice. Nat Genet 1996; 14(1): 95-97.
112. Cai A, Hyde JF. Upregulation of leptin receptor gene expression in the anterior pituitary of human growth hormone-releasing hormone transgenic mice. Endocrinology 1998; 139(1): 420-423.
113. Cai A, Hyde JF. The human growth hormone-releasing hormone transgenic mouse as a model of modest obesity: differential changes in leptin receptor (OBR) gene expression in the anterior pituitary and hypothalamus after fasting and OBR localization in somatotrophs. Endocrinology 1999; 140(8): 3609-36014.
114. Giusti M, Bocca L, Florio T, et al. In vitro effect of human recombinant leptin and expression of leptin receptors on growth hormone-secreting human pituitary adenomas. Clin Endocrinol (Oxf) 2002; 57(4): 449-455.
115. Lin J, Barb CR, Matteri RL, et al. Long form leptin receptor mRNA expression in the brain, pituitary, and other tissues in the pig. Domest Anim Endocrinol 2000; 19(1): 53-61.
116. Iqbal J, Pompolo S, Considine RV, Clarke IJ. Localization of leptin receptor-like immunoreactivity in the corticotropes, somatotropes, and gonadotropes in the ovine anterior pituitary. Endocrinology 2000; 141(4): 1515-1520.
117. Crane C, Akhter N, Johnson BW, et al. Fasting and Glucose Effects on Pituitary Leptin Expression: Is Leptin a Local Signal for Nutrient Status? J Histochem Cytochem 2007; 55(10): 1059-1074.
118. Kaminski T, Smolinska N, Gajewska A, et al. Leptin and long form of leptin receptor genes expression in the hypothalamus and pituitary during the luteal phase and early pregnancy in pigs. J Physiol Pharmacol 2006; 57(1): 95-108.
119. Childs GV, Iruthayanathan M, Akhter N, Unabia G, Whitehead- Johnson B. Bipotential effects of estrogen on growth hormone synthesis and storage in vitro. Endocrinology 2005; 146(4): 1780-1788.
120. Gonzalez LC, Pinilla L, Tena-Sempere M, Dieguez C, Casanueva FF, Aguilar E. Effect of acute immunoneutralization of endogenous leptin on prolactin and LH secretion during the afternoon of prooestrus or in steroid-treated ovariectomized female rats. J Reprod Fertil 2000; 118(1): 39-45.
121. Schoeller DA, Cella LK, Sinha MK, Caro JF. Entrainment of the diurnal rhythm of plasma leptin to meal timing. J Clin Invest 1997 Oct 1; 100(7): 1882-1887.
122. Gui Y, Silha JV, Mishra S, Murphy LJ. Changes in adipokine expression during food deprivation in the mouse and the relationship to fasting-induced insulin resistance. Can J Physiol Pharmacol 2003; 81(10): 979-985.
123. Kowalska I, Straczkowski M, Gorski J, Kinalska I. The effect of fasting and physical exercise on plasma leptin concentrations in high-fat fed rats. J Physiol Pharmacol 1999; 50(2): 309-320.
124. Igel M, Kainulainen H, Brauers A, Becker W, Herberg L, Joost HG. Long-term and rapid regulation of ob mRNA levels in adipose tissue from normal (Sprague Dawley rats) and obese (db/db mice, fa/fa rats) rodents. Diabetologia 1996; 39(7): 7587-7565.
125. MacDougald OA, Hwang CS, Fan H, Lane MD. Regulated expression of the obese gene product (leptin) in white adipose tissue and 3T3-L1 adipocytes. Proc Natl Acad Sci USA 1995; 92(20): 9034-9037.
126. Childs GV, Unabia G, Miller BT, Collins TJ. Differential expression of gonadotropin and prolactin antigens by GHRH target cells from male and female rats. J Endocrinol 1999; 162(2): 177-187.
127. Kojima M, Hosoda H, Date Y, Nakazato M, Matsuo H, Kangawa K. Ghrelin is a growth-hormone-releasing acylated peptide from stomach. Nature 1999; 402(6762): 656-660.
128. Kojima M, Hosoda H, Matsuo H, Kangawa K. Ghrelin: discovery of the natural endogenous ligand for the growth hormone secretagogue receptor. Trends Endocrinol Metab 2001; 12(3): 118-122.
129. Takaya K, Ariyasu H, Kanamoto N, et al. Ghrelin strongly stimulates growth hormone release in humans. J Clin Endocrinol Metab 2000; 85(12): 4908-4911.
130. Wren AM, Small CJ, Ward HL, et al. The novel hypothalamic peptide ghrelin stimulates food intake and growth hormone secretion. Endocrinology 2000; 141(11): 4325-4328.
131. Malagon MM, Luque RM, Ruiz-Guerrero E, et al. Intracellular signaling mechanisms mediating ghrelin-stimulated growth hormone release in somatotropes. Endocrinology 2003; 144(12): 5372-5380.
132. Pinilla L, Barreiro ML, Tena-Sempere M, Aguilar E. Role of ghrelin in the control of growth hormone secretion in prepubertal rats: interactions with excitatory amino acids. Neuroendocrinology 2003; 77(2): 83-90.
133. Kappeler L, Zizzari P, Grouselle D, Epelbaum J, Bluet-Pajot MT. Plasma and hypothalamic peptide-hormone levels regulating somatotroph function and energy balance in fed and fasted states: a comparative study in four strains of rats. J Neuroendocrinol 2004; 16(12): 980-988.
134. Korbonits M, Goldstone AP, Gueorguiev M, Grossman AB. Ghrelin--a hormone with multiple functions. Front Neuroendocrinol 2004; 25(1): 27-68.
135. van der Lely AJ, Tschop M, Heiman ML, Ghigo E. Biological, physiological, pathophysiological, and pharmacological aspects of ghrelin. Endocr Rev 2004; 25(3): 426-457.
136. Chen C. Growth hormone secretagogue actions on the pituitary gland: multiple receptors for multiple ligands? Clin Exp Pharmacol Physiol 2000; 27(5-6): 323-329.
137. Frohman LA, Kineman RD, Kamegai J, et al. Secretagogues and the somatotrope: signaling and proliferation. Recent Prog Horm Res 2000; 55: 269-290; discussion 90-91.
138. Gracia-Navarro F, Castano JP, Malagon MM, et al. Research progress in the stimulatory inputs regulating growth hormone (GH) secretion. Comp Biochem Physiol B Biochem Mol Biol 2002; 132(1): 141-150.
139. McKee KK, Palyha OC, Feighner SD, et al. Molecular analysis of rat pituitary and hypothalamic growth hormone secretagogue receptors. Mol Endocrinol 1997; 11(4): 415-423.
140. Park S, Peng XD, Frohman LA, Kineman RD. Expression analysis of hypothalamic and pituitary components of the growth hormone axis in fasted and streptozotocin-treated neuropeptide Y (NPY)- intact (NPY+/+) and NPY-knockout (NPY-/-) mice. Neuroendocrinology 2005; 81(6): 360-371.
141. Fernandez-Fernandez R, Navarro VM, Barreiro ML, et al. Effects of chronic hyperghrelinemia on puberty onset and pregnancy outcome in the rat. Endocrinology 2005; 146(7): 3018-3025.
142. Fernandez-Fernandez R, Tena-Sempere M, Navarro VM, et al. Effects of ghrelin upon gonadotropin-releasing hormone and gonadotropin secretion in adult female rats: in vivo and in vitro studies. Neuroendocrinology 2005; 82(5-6): 245-255.
143. Fernandez-Fernandez R, Tena-Sempere M, Navarro VM, et al. Effects of Ghrelin upon Gonadotropin-Releasing Hormone and Gonadotropin Secretion in Adult Female Rats: In vivo and in vitro Studies. Neuroendocrinology 2006; 82(5-6): 245-255.
144. Furuta M, Funabashi T, Kimura F. Intracerebroventricular administration of ghrelin rapidly suppresses pulsatile luteinizing hormone secretion in ovariectomized rats. Biochem Biophys Res Commun 2001; 288(4): 780-785.
145. Vulliemoz NR, Xiao E, Xia-Zhang L, Germond M, Rivier J, Ferin M. Decrease in luteinizing hormone pulse frequency during a fivehour peripheral ghrelin infusion in the ovariectomized rhesus monkey. J Clin Endocrinol Metab 2004; 89(11): 5718-5723.
146. Iqbal J, Kurose Y, Canny B, Clarke IJ. Effects of central infusion of ghrelin on food intake and plasma levels of growth hormone, luteinizing hormone, prolactin, and cortisol secretion in sheep. Endocrinology 2006; 147(1): 510-519.
147. Clayton RN. Gonadotrophin-releasing hormone: its actions and receptors. J Endocrinol 1989; 120(1): 11-19.
148. Conn PM, editor. The molecular mechanism of gonadotropinreleasing hormone action in the pituitary. 2nd ed. New York: New York: Raven Press 1994.
149. Conn PM, Huckle WR, Andrews WV, McArdle CA. The molecular mechanism of action of gonadotropin releasing hormone (GnRH) in the pituitary. Recent Prog Horm Res 1987; 43: 29-68.
150. Belchetz PE, Plant TM, Nakai Y, Keogh EJ, Knobil E. Hypophysial responses to continuous and intermittent delivery of hypothalamic gonadotropin-releasing hormone. Science 1978; 202(4368): 631-633.
151. Burger LL, Dalkin AC, Aylor KW, Haisenleder DJ, Marshall JC. 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 2002; 143(9): 3243-3249.
152. Crowley WF, Jr., Filicori M, Spratt DI, Santoro NF. The physiology of gonadotropin-releasing hormone (GnRH) secretion in men and women. Recent Prog Horm Res 1985; 41: 473-531.
153. Haisenleder DJ, Dalkin AC, Ortolano GA, Marshall JC, Shupnik MA. A pulsatile gonadotropin-releasing hormone stimulus is required to increase transcription of the gonadotropin subunit genes: evidence for differential regulation of transcription by pulse frequency in vivo. Endocrinology 1991; 128(1): 509-517.
154. Kaiser UB, Jakubowiak A, Steinberger A, Chin WW. Differential effects of gonadotropin-releasing hormone (GnRH) pulse frequency on gonadotropin subunit and GnRH receptor messenger ribonucleic acid levels in vitro. Endocrinology 1997; 138(3): 1224-1231.
155. Kaiser UB, Sabbagh E, Katzenellenbogen RA, Conn PM, Chin WW. A mechanism for the differential regulation of gonadotropin subunit gene expression by gonadotropin-releasing hormone. Proc Natl Acad Sci USA 1995; 92(26): 12280-12284.
156. Levine JE, Ramirez VD. Luteinizing hormone-releasing hormone release during the rat estrous cycle and after ovariectomy, as estimated with push-pull cannulae. Endocrinology 1982; 111(5): 1439-1448.
157. Lloyd JM, Childs GV. Changes in the number of GnRH-receptive cells during the rat estrous cycle: biphasic effects of estradiol. Neuroendocrinology 1988; 48(2): 138-146.
158. Loumaye E, Catt KJ. Homologous regulation of gonadotropinreleasing hormone receptors in cultured pituitary cells. Science 1982; 215(4535): 983-985.
159. Savoy-Moore RT, Schwartz NB, Duncan JA, Marshall JC. Pituitary gonadotropin-releasing hormone receptors during the rat estrous cycle. Science 1980; 209(4459): 942-944.
160. Carro E, Pinilla L, Seoane LM, et al. Influence of endogenous leptin tone on the estrous cycle and luteinizing hormone pulsatility in female rats. Neuroendocrinology 1997; 66(6): 375-377.
161. Childs GV, Naor Z, Hazum E, Tibolt R, Westlund KN, Hancock MB. Localization of biotinylated gonadotropin releasing hormone on pituitary monolayer cells with avidin-biotin-peroxidase complexes. J Histochem Cytochem 1983; 31(12): 1422-1425.
162. Childs GV, Naor Z, Hazum E, Tibolt R, Westlund KN, Hancock MB. Cytochemical characterization of pituitary target cells for biotinylated gonadotropin releasing hormone. Peptides 1983; 4(4): 549-555.
163. Childs GV, Unabia G. Cytochemical studies of the effects of activin on gonadotropin-releasing hormone (GnRH) binding by pituitary gonadotropes and growth hormone cells. J Histochem Cytochem 1997; 45(12): 1603-1610.
164. Childs GV, Unabia G, Miller BT. Cytochemical detection of gonadotropin-releasing hormone-binding sites on rat pituitary cells with luteinizing hormone, follicle-stimulating hormone, and growth hormone antigens during diestrous up-regulation. Endocrinology 1994; 134(4): 1943-1951.
165. Childs GV, Unabia G, Rougeau D. Cells that express luteinizing hormone (LH) and follicle-stimulating hormone (FSH) beta-subunit messenger ribonucleic acids during the estrous cycle: the major contributors contain LH beta, FSH beta, and/or growth hormone. Endocrinology 1994; 134(2): 990-997.
166. Childs GV. Growth hormone cells as co-gonadotropes: partners in the regulation of the reproductive system. Trends Endocrinol Metab 2000; 11(5): 168-1675.
167. Sutton SW, Toyama TT, Otto S, Plotsky PM. Evidence that neuropeptide Y (NPY) released into the hypophysial-portal circulation participates in priming gonadotropes to the effects of gonadotropin releasing hormone (GnRH). Endocrinology 1988; 123(2): 1208-1210.
168. Bauer-Dantoin AC, McDonald JK, Levine JE. Neuropeptide Y potentiates luteinizing hormone (LH)-releasing hormone-stimulated LH surges in pentobarbital-blocked proestrous rats. Endocrinology 1991; 129(1): 402-408.
169. Watanobe H, Takebe K. Evidence that neuropeptide Y secretion in the median eminence increases prior to the luteinizing hormone surge in ovariectomized steroid-primed rats: estimation by pushpull perfusion. Neurosci Lett 1992; 146(1): 57-59.
170. Woller MJ, McDonald JK, Reboussin DM, Terasawa E. Neuropeptide Y is a neuromodulator of pulsatile luteinizing hormone-releasing hormone release in the gonadectomized rhesus monkey. Endocrinology 1992; 130(4): 2333-2342.
171. Xu M, Hill JW, Levine JE. Attenuation of luteinizing hormone surges in neuropeptide Y knockout mice. Neuroendocrinology 2000; 72(5): 263-271.
172. Xu M, Urban JH, Hill JW, Levine JE. Regulation of hypothalamic neuropeptide Y Y1 receptor gene expression during the estrous cycle: role of progesterone receptors. Endocrinology 2000; 141(9): 3319-3327.
173. Morrison CD, Morton GJ, Niswender KD, Gelling RW, Schwartz MW. Leptin inhibits hypothalamic Npy and Agrp gene expression via a mechanism that requires phosphatidylinositol 3-OH-kinase signaling. Am J Physiol Endocrinol Metab 2005; 289(6): E1051-E1057.
174. Schwartz MW, Baskin DG, Bukowski TR, et al. Specificity of leptin action on elevated blood glucose levels and hypothalamic neuropeptide Y gene expression in ob/ob mice. Diabetes 1996; 45(4): 531-535.
175. Schwartz MW, Seeley RJ, Campfield LA, Burn P, Baskin DG. Identification of targets of leptin action in rat hypothalamus. J Clin Invest. 1996; 98(5): 1101-1106.
176. Hill JW, Urban JH, Xu M, Levine JE. Estrogen Induces Neuropeptide Y (NPY) Y1 receptor gene expression and responsiveness to NPY in gonadotrope-enriched pituitary cell cultures. Endocrinology 2004; 145(5): 2283-2290.
177. Bauer-Dantoin AC, Tabesh B, Norgle JR, Levine JE. RU486 administration blocks neuropeptide Y potentiation of luteinizing hormone (LH)-releasing hormone-induced LH surges in proestrous rats. Endocrinology 1993; 133(6): 2418-2423.
178. Childs GV, Iruthayanathan M, Akhter N, Johnson BW. Estrogen mediated cross talk between the ovary and pituitary somatotrope. Pre-ovulatory support for reproductive activity. Mol Cell Endocrinol 2006; 247(1/2): 60-63.
179. Isozaki O, Tsushima T, Miyakawa M, Demura H, Seki H. Interaction between leptin and growth hormone (GH)/IGF-I axis. Endocr J 1999; 46(Suppl): S17-S24.
180. Baratta M, Saleri R, Mainardi GL, Valle D, Giustina A, Tamanini C. Leptin regulates GH gene expression and secretion and nitric oxide production in pig pituitary cells. Endocrinology 2002; 143(2): 551-557.
181. Saleri R, Giustina A, Tamanini C, et al. Leptin stimulates growth hormone secretion via a direct pituitary effect combined with a decreased somatostatin tone in a median eminence-pituitary perifusion study. Neuroendocrinology 2004; 79(4): 221-228.
182. Saleri R, Grasselli F, Tamanini C. Effects of different culture conditions and leptin on GH mRNA expression and GH secretion by pig pituitary cells. Horm Metab Res 2005; 37(4): 214-219.
183. Mizuno I, Okimura Y, Takahashi Y, Kaji H, Abe H, Chihara K. Leptin stimulates basal and GHRH-induced GH release from cultured rat anterior pituitary cells in vitro. Kobe J Med Sci 1999; 45(5): 221-227.
184. Zieba DA, Amstalden M, Morton S, et al. Effects of leptin on basal and GHRH-stimulated GH secretion from the bovine adenohypophysis are dependent upon nutritional status. J Endocrinol 2003; 178(1): 83-89.
185. Hardie LJ, Rayner DV, Holmes S, Trayhurn P. Circulating leptin levels are modulated by fasting, cold exposure and insulin administration in lean but not Zucker (fa/fa) rats as measured by ELISA. Biochem Biophys Res Commun 1996; 223(3): 660-665.
186. Luque RM, Park S, Kineman RD. Severity of the catabolic condition differentially modulates hypothalamic expression of growth hormone-releasing hormone in the fasted mouse: potential role of neuropeptide Y and corticotropin-releasing hormone. Endocrinology 2007; 148(1): 300-309.
187. Maciel MN, Zieba DA, Amstalden M, Keisler DH, Neves JP, Williams GL. Leptin prevents fasting-mediated reductions in pulsatile secretion of luteinizing hormone and enhances its gonadotropin-releasing hormone-mediated release in heifers. Biol Reprod 2004; 70(1): 229-235.
188. Sorenson RL, Stout LE, Brelje TC, Jetton TL, Matschinsky FM. Immunohistochemical evidence for the presence of glucokinase in the gonadotropes and thyrotropes of the anterior pituitary gland of rat and monkey. J Histochem Cytochem 2007; 55(6): 555-566.
189. Zelent D, Golson ML, Koeberlein B, et al. A glucose sensor role for glucokinase in anterior pituitary cells. Diabetes 2006; 55(7): 1923-1929.