Central regulation of energy metabolism by estrogens

Mauvais-Jarvis, 2013, The role of estrogens in control of energy balance and glucose homeostasis, Endocrine Reviews, 34, 309, 10.1210/er.2012-1055 Lopez, 2015, Estrogens and the control of energy homeostasis: a brain perspective, Trends in Endocrinology and Metabolism, 26, 411, 10.1016/j.tem.2015.06.003 Palmer, 2015, The sexual dimorphism of obesity, Molecular and Cellular Endocrinology, 402, 113, 10.1016/j.mce.2014.11.029 Flegal, 2010, Prevalence and trends in obesity among US adults, 1999-2008, Journal of the American Medical Association, 303, 235, 10.1001/jama.2009.2014 Billeci, 2008, Hormone replacement therapy and stroke, Current Vascular Pharmacology, 6, 112, 10.2174/157016108783955338 Canonico, 2007, Hormone therapy and venous thromboembolism among postmenopausal women: impact of the route of estrogen administration and progestogens: the ESTHER study, Circulation, 115, 840, 10.1161/CIRCULATIONAHA.106.642280 Saito, 2015, Progress in the molecular understanding of central regulation of body weight by estrogens, Obesity (Silver Spring), 23, 919, 10.1002/oby.21099 Lopez, 2016, Estradiol and brown fat, Best Practice and Research Clinical Endocrinology and Metabolism, 30, 527, 10.1016/j.beem.2016.08.004 Asarian, 2013, Sex differences in the physiology of eating, American Journal of Physiology - Regulatory, Integrative and Comparative Physiology, 305, R1215, 10.1152/ajpregu.00446.2012 Blaustein, 1976, Ovarian influences on the meal patterns of female rats, Physiology and Behavior, 17, 201, 10.1016/0031-9384(76)90064-0 Drewett, 1973, Sexual behaviour and sexual motivation in the female rat, Nature, 242, 476, 10.1038/242476a0 Geary, 2001, Deficits in E2-dependent control of feeding, weight gain, and cholecystokinin satiation in ER-alpha null mice, Endocrinology, 142, 4751, 10.1210/endo.142.11.8504 Wallen, 2001, Sex hormones and the selective estrogen receptor modulator tamoxifen modulate weekly body weights and food intakes in adolescent and adult rats, Journal of Nutrition, 131, 2351, 10.1093/jn/131.9.2351 Roesch, 2006, Effects of selective estrogen receptor agonists on food intake and body weight gain in rats, Physiology and Behavior, 87, 39, 10.1016/j.physbeh.2005.08.035 Rogers, 2009, Reduced energy expenditure and increased inflammation are early events in the development of ovariectomy-induced obesity, Endocrinology, 10.1210/en.2008-1405 Komm, 2012, Incorporating bazedoxifene/conjugated estrogens into the current paradigm of menopausal therapy, International Journal of Womens Health, 4, 129, 10.2147/IJWH.S29346 Kim, 2014, Tissue-selective estrogen complexes with bazedoxifene prevent metabolic dysfunction in female mice, Molecular Metabolism, 3, 177, 10.1016/j.molmet.2013.12.009 Cedars, 1987, Nonoral routes of estrogen administration, Obstetrics and Gynecology Clinics of North America, 14, 269, 10.1016/S0889-8545(21)00583-0 Okura, 2003, Association of polymorphisms in the estrogen receptor alpha gene with body fat distribution, International Journal of Obesity and Related Metabolic Disorders, 27, 1020, 10.1038/sj.ijo.0802378 Heine, 2000, Increased adipose tissue in male and female estrogen receptor-alpha knockout mice, Proceedings of the National Academy of Sciences of the United States of America, 97, 12729, 10.1073/pnas.97.23.12729 Butera, 1989, Central implants of diluted estradiol: independent effects on ingestive and reproductive behaviors of ovariectomized rats, Brain Research, 491, 266, 10.1016/0006-8993(89)90062-0 Palmer, 1986, Central vs. peripheral effects of estrogen on food intake and lipoprotein lipase activity in ovariectomized rats, Physiology and Behavior, 37, 187, 10.1016/0031-9384(86)90404-X Xu, 2011, Distinct hypothalamic neurons mediate estrogenic effects on energy homeostasis and reproduction, Cell Metabolism, 14, 453, 10.1016/j.cmet.2011.08.009 Merchenthaler, 2004, Distribution of estrogen receptor alpha and beta in the mouse central nervous system: in vivo autoradiographic and immunocytochemical analyses, Journal of Comparative Neurology, 473, 270, 10.1002/cne.20128 Morton, 2006, Central nervous system control of food intake and body weight, Nature, 443, 289, 10.1038/nature05026 Miller, 1995, Effects of age and long-term ovariectomy on the estrogen-receptor containing subpopulations of beta-endorphin-immunoreactive neurons in the arcuate nucleus of female C57BL/6J mice, Neuroendocrinology, 61, 542, 10.1159/000126878 de Souza, 2011, The estrogen receptor alpha colocalizes with proopiomelanocortin in hypothalamic neurons and binds to a conserved motif present in the neuron-specific enhancer nPE2, European Journal of Pharmacology, 660, 181, 10.1016/j.ejphar.2010.10.114 Gao, 2007, Anorectic estrogen mimics leptin's effect on the rewiring of melanocortin cells and Stat3 signaling in obese animals, Nature Medicine, 13, 89, 10.1038/nm1525 Malyala, 2008, PI3K signaling effects in hypothalamic neurons mediated by estrogen, Journal of Comparative Neurology, 506, 895, 10.1002/cne.21584 Zhu, 2015, The eralpha-PI3K cascade in proopiomelanocortin Progenitor neurons regulates feeding and glucose balance in female mice, Endocrinology, 156, 4474, 10.1210/en.2015-1660 Osterlund, 1998, Differential distribution and regulation of estrogen receptor-alpha and -beta mRNA within the female rat brain, Brain Research Molecular Brain Research, 54, 175, 10.1016/S0169-328X(97)00351-3 Schlenker, 2006, Sex-specific densities of estrogen receptors alpha and beta in the subnuclei of the nucleus tractus solitarius, hypoglossal nucleus and dorsal vagal motor nucleus weanling rats, Brain Research, 1123, 89, 10.1016/j.brainres.2006.09.035 Asarian, 2007, Estradiol enhances cholecystokinin-dependent lipid-induced satiation and activates estrogen receptor-alpha-expressing cells in the nucleus tractus solitarius of ovariectomized rats, Endocrinology, 148, 5656, 10.1210/en.2007-0341 Thammacharoen, 2008, Hindbrain administration of estradiol inhibits feeding and activates estrogen receptor-alpha-expressing cells in the nucleus tractus solitarius of ovariectomized rats, Endocrinology, 149, 1609, 10.1210/en.2007-0340 Cao, 2014, Estrogens stimulate serotonin neurons to inhibit binge-like eating in mice, Journal of Clinical Investigation, 124, 4351, 10.1172/JCI74726 Dalmasso, 2011, Serotonergic system involvement in the inhibitory action of estrogen on induced sodium appetite in female rats, Physiology and Behavior, 104, 398, 10.1016/j.physbeh.2011.04.029 Robichaud, 2005, Oestrogen and testosterone modulate the firing activity of dorsal raphe nucleus serotonergic neurones in both male and female rats, Journal of Neuroendocrinology, 17, 179, 10.1111/j.1365-2826.2005.01292.x Santollo, 2011, Estradiol acts in the medial preoptic area, arcuate nucleus, and dorsal raphe nucleus to reduce food intake in ovariectomized rats, Hormones and Behavior, 60, 86, 10.1016/j.yhbeh.2011.03.009 Butera, 1992, Effects of PVN lesions on the responsiveness of female rats to estradiol, Brain Research, 576, 304, 10.1016/0006-8993(92)90694-5 Hrupka, 2002, Hypothalamic implants of dilute estradiol fail to reduce feeding in ovariectomized rats, Physiology and Behavior, 77, 233, 10.1016/S0031-9384(02)00857-0 Shimizu, 1993, Effects of castration, estrogen replacement and estrus cycle on monoamine metabolism in the nucleus accumbens, measured by microdialysis, Brain Research, 621, 200, 10.1016/0006-8993(93)90107-X Cannon, 2004, Brown adipose tissue: function and physiological significance, Physiological Reviews, 84, 277, 10.1152/physrev.00015.2003 Silva, 2006, Thermogenic mechanisms and their hormonal regulation, Physiological Reviews, 86, 435, 10.1152/physrev.00009.2005 Nedergaard, 2007, Unexpected evidence for active brown adipose tissue in adult humans, American Journal of Physiology Endocrinology and Metabolism, 293, E444, 10.1152/ajpendo.00691.2006 Contreras, 2015, The brain and brown fat, Annals of Medicine, 47, 150, 10.3109/07853890.2014.919727 Cypess, 2009, Identification and importance of brown adipose tissue in adult humans, New England Journal of Medicine, 360, 1509, 10.1056/NEJMoa0810780 van Marken Lichtenbelt, 2009, Cold-activated brown adipose tissue in healthy men, New England Journal of Medicine, 360, 1500, 10.1056/NEJMoa0808718 Virtanen, 2009, Functional brown adipose tissue in healthy adults, New England Journal of Medicine, 360, 1518, 10.1056/NEJMoa0808949 Nedergaard, 2014, The browning of white adipose tissue: some burning issues, Cell Metabolism, 20, 396, 10.1016/j.cmet.2014.07.005 Futai, 1989, ATP synthase (H+-ATPase): results by combined biochemical and molecular biological approaches, Annual Review of Biochemistry, 58, 111, 10.1146/annurev.bi.58.070189.000551 Boyer, 1997, The ATP synthase–a splendid molecular machine, Annual Review of Biochemistry, 66, 717, 10.1146/annurev.biochem.66.1.717 von Ballmoos, 2009, Essentials for ATP synthesis by F1F0 ATP synthases, Annual Review of Biochemistry, 78, 649, 10.1146/annurev.biochem.78.081307.104803 Nicholls, 1984, Thermogenic mechanisms in brown fat, Physiological Reviews, 64, 1, 10.1152/physrev.1984.64.1.1 Whittle, 2011, Using brown adipose tissue to treat obesity - the central issue, Trends in Molecular Medicine, 17, 405, 10.1016/j.molmed.2011.04.001 Morrison, 2012, Central control of brown adipose tissue thermogenesis, Frontiers in Endocrinology, 3, 10.3389/fendo.2012.00005 Morrison, 2014, Central neural regulation of brown adipose tissue thermogenesis and energy expenditure, Cell Metabolism, 19, 741, 10.1016/j.cmet.2014.02.007 Wade, 1978, Cytoplasmic 17 beta-[3H]estradiol binding in rat adipose tissues, Endocrinology, 103, 1695, 10.1210/endo-103-5-1695 Edens, 1983, Effects of estradiol on tissue distribution of newly-synthesized fatty acids in rats and hamsters, Physiology and Behavior, 31, 703, 10.1016/S0031-9384(83)80007-9 Bartness, 1984, Effects of interscapular brown adipose tissue denervation on body weight and energy metabolism in ovariectomized and estradiol-treated rats, Behavioral Neuroscience, 98, 674, 10.1037/0735-7044.98.4.674 Schneider, 1986, Effects of estrous cycles and ovarian steroids on body weight and energy expenditure in Syrian hamsters, Physiology and Behavior, 38, 119, 10.1016/0031-9384(86)90141-1 Richard, 1986, Effects of ovarian hormones on energy balance and brown adipose tissue thermogenesis, American Journal of Physiology, 250, R245 Kamei, 2005, Ovariectomy in mice decreases lipid metabolism-related gene expression in adipose tissue and skeletal muscle with increased body fat, Journal of Nutritional Science and Vitaminology, 51, 110, 10.3177/jnsv.51.110 Yoshida, 1987, Reduced norepinephrine turnover in interscapular brown adipose tissue of obese rats after ovariectomy, Metabolism, 36, 1, 10.1016/0026-0495(87)90054-0 Lopez, 2013, Energy balance regulation by thyroid hormones at central level, Trends in Molecular Medicine, 19, 418, 10.1016/j.molmed.2013.04.004 Ribeiro, 2000, Evidence of UCP1-independent regulation of norepinephrine-induced thermogenesis in brown fat, American Journal of Physiology Endocrinology and Metabolism, 279, E314, 10.1152/ajpendo.2000.279.2.E314 de Jesus, 2001, The type 2 iodothyronine deiodinase is essential for adaptive thermogenesis in brown adipose tissue, Journal of Clinical Investigation, 108, 1379, 10.1172/JCI200113803 Martinez-Sanchez, 2014, Hypothalamic effects of thyroid hormones on metabolism, Best Practice and Research Clinical Endocrinology and Metabolism, 28, 703, 10.1016/j.beem.2014.04.004 Rodriguez, 2002, Opposite actions of testosterone and progesterone on UCP1 mRNA expression in cultured brown adipocytes, Cellular and Molecular Life Sciences, 59, 1714, 10.1007/PL00012499 Monjo, 2003, Direct effects of testosterone, 17 beta-estradiol, and progesterone on adrenergic regulation in cultured brown adipocytes: potential mechanism for gender-dependent thermogenesis, Endocrinology, 144, 4923, 10.1210/en.2003-0537 Rodriguez-Cuenca, 2007, Expression of mitochondrial biogenesis-signaling factors in brown adipocytes is influenced specifically by 17beta-estradiol, testosterone, and progesterone, American Journal of Physiology Endocrinology and Metabolism, 292, E340, 10.1152/ajpendo.00175.2006 Rodriguez-Cuenca, 2007, Sex steroid receptor expression profile in brown adipose tissue. Effects of hormonal status, Cell Physiology Biochemistry, 20, 877, 10.1159/000110448 Cooke, 2001, The role of estrogen and estrogen receptor-alpha in male adipose tissue, Molecular and Cellular Endocrinology, 178, 147, 10.1016/S0303-7207(01)00414-2 Foryst-Ludwig, 2008, Metabolic actions of estrogen receptor beta (ERbeta) are mediated by a negative cross-talk with PPARgamma, PLoS Genetics, 4, e1000108, 10.1371/journal.pgen.1000108 Hewitt, 2004, Estrogen replacement reverses the hepatic steatosis phenotype in the male aromatase knockout mouse, Endocrinology, 145, 1842, 10.1210/en.2003-1369 Davis, 2014, Sexually dimorphic role of G protein-coupled estrogen receptor (GPER) in modulating energy homeostasis, Hormones and Behavior, 66, 196, 10.1016/j.yhbeh.2014.02.004 Velickovic, 2014, Expression and subcellular localization of estrogen receptors alpha and beta in human fetal brown adipose tissue, The Journal of Cinical Endocrinology and Metabolism, 99, 151, 10.1210/jc.2013-2017 Shughrue, 1997, Comparative distribution of estrogen receptor-alpha and -beta mRNA in the rat central nervous system, Journal of Comparative Neurology, 388, 507, 10.1002/(SICI)1096-9861(19971201)388:4<507::AID-CNE1>3.0.CO;2-6 Martinez de Morentin, 2015, Pregnancy induces resistance to the anorectic effect of hypothalamic malonyl-CoA and the thermogenic effect of hypothalamic AMPK inhibition in female rats, Endocrinology, 156, 947, 10.1210/en.2014-1611 Perkins, 1981, Activation of brown adipose tissue thermogenesis by the ventromedial hypothalamus, Nature, 289, 401, 10.1038/289401a0 Yoshimatsu, 1993, Sympathetic nerve activity after discrete hypothalamic injections of L-glutamate, Brain Research, 601, 121, 10.1016/0006-8993(93)91702-T Lopez, 2010, Hypothalamic AMPK and fatty acid metabolism mediate thyroid regulation of energy balance, Nature Medicine, 16, 1001, 10.1038/nm.2207 Whittle, 2012, BMP8B increases brown adipose tissue thermogenesis through both central and peripheral actions, Cell, 149, 871, 10.1016/j.cell.2012.02.066 Martinez de Morentin, 2012, Nicotine induces negative energy balance through hypothalamic AMP-activated protein kinase, Diabetes, 61, 807, 10.2337/db11-1079 Seoane-Collazo, 2014, Nicotine improves obesity and hepatic steatosis and ER stress in diet-induced obese male rats, Endocrinology, 155, 1679, 10.1210/en.2013-1839 Martinez de Morentin, 2014, Estradiol regulates Brown adipose tissue thermogenesis via hypothalamic AMPK, Cell Metabolism, 20, 41, 10.1016/j.cmet.2014.03.031 Contreras, 2017, Reduction of hypothalamic endoplasmic reticulum stress activates browning of white fat and ameliorates obesity, Diabetes, 66, 87, 10.2337/db15-1547 Minami, 1990, 17 beta-estradiol depolarization of hypothalamic neurons is mediated by cyclic AMP, Brain Research, 519, 301, 10.1016/0006-8993(90)90092-P Musatov, 2007, Silencing of estrogen receptor alpha in the ventromedial nucleus of hypothalamus leads to metabolic syndrome, Proceedings of the National Academy of Sciences of the United States of America, 104, 2501, 10.1073/pnas.0610787104 Yepuru, 2010, Estrogen receptor-{beta}-selective ligands alleviate high-fat diet- and ovariectomy-induced obesity in mice, Journal of Biological Chemistry, 285, 31292, 10.1074/jbc.M110.147850 Correa, 2015, An estrogen-responsive module in the ventromedial hypothalamus selectively drives sex-specific activity in females, Cell Reports, 10, 62, 10.1016/j.celrep.2014.12.011 Lopez, 2016, Hypothalamic AMPK: a canonical regulator of whole-body energy balance, Nature Reviews Endocrinology, 12, 421, 10.1038/nrendo.2016.67 Tanida, 2011, Central AMP-activated protein kinase affects sympathetic nerve activity in rats, Neuroscience Letters, 503, 167, 10.1016/j.neulet.2011.08.013 Tanida, 2013, Involvement of hypothalamic AMP-activated protein kinase in leptin-induced sympathetic nerve activation, PLoS One, 8, e56660, 10.1371/journal.pone.0056660 Beiroa, 2014, GLP-1 agonism stimulates brown adipose tissue thermogenesis and browning through hypothalamic AMPK, Diabetes, 63, 3346, 10.2337/db14-0302 Alvarez-Crespo, 2016, Essential role of UCP1 modulating the central effects of thyroid hormones on energy balance, Molecular Metabolism, 5, 271, 10.1016/j.molmet.2016.01.008 Martins, 2016, A functional link between AMPK and orexin mediates the effect of BMP8B on energy balance, Cell Reports, 16, 2231, 10.1016/j.celrep.2016.07.045 Martinez-Sanchez, 2017, Thyroid hormones induce browning of white fat, Journal of Endocrinology, 232, 351, 10.1530/JOE-16-0425 Finan, 2012, Targeted estrogen delivery reverses the metabolic syndrome, Nature Medicine, 18, 1847, 10.1038/nm.3009 Lopez, 2017, Estradiol effects on hypothalamic AMPK and BAT thermogenesis: a gateway for obesity treatment?, Pharmacology and Therapeutics, 178, 109, 10.1016/j.pharmthera.2017.03.014 Lopez, 2017, 2017: hypothalamic AMPK: a golden target against obesity?, European Journal of Endocrinology, 176, R235, 10.1530/EJE-16-0927 Yonezawa, 2012, Central versus peripheral impact of estradiol on the impaired glucose metabolism in ovariectomized mice on a high-fat diet, American Journal of Physiology Endocrinology and Metabolism, 303, E445, 10.1152/ajpendo.00638.2011 McCrimmon, 2004, Potential role for AMP-activated protein kinase in hypoglycemia sensing in the ventromedial hypothalamus, Diabetes, 53, 1953, 10.2337/diabetes.53.8.1953 Han, 2005, Hypothalamic AMP-activated protein kinase mediates counter-regulatory responses to hypoglycaemia in rats, Diabetologia, 48, 2170, 10.1007/s00125-005-1913-1 McCrimmon, 2006, Activation of AMP-activated protein kinase within the ventromedial hypothalamus amplifies counterregulatory hormone responses in rats with defective counterregulation, Diabetes, 55, 1755, 10.2337/db05-1359 McCrimmon, 2008, Key role for AMP-activated protein kinase in the ventromedial hypothalamus in regulating counterregulatory hormone responses to acute hypoglycemia, Diabetes, 57, 444, 10.2337/db07-0837 Tamrakar, 2015, Estradiol regulation of hypothalamic astrocyte adenosine 5'-monophosphate-activated protein kinase activity: role of hindbrain catecholamine signaling, Brain Research Bulletin, 110, 47, 10.1016/j.brainresbull.2014.12.002 Briski, 2014, Energy metabolism and hindbrain AMPK: regulation by estradiol, Hormone Molecular Biology and Clinical Investigation, 17, 129, 10.1515/hmbci-2013-0067 Park, 2011, Genetic rescue of nonclassical ERalpha signaling normalizes energy balance in obese Eralpha-null mutant mice, Journal of Clinical Investigation, 121, 604, 10.1172/JCI41702 Jakacka, 2002, An estrogen receptor (ER)alpha deoxyribonucleic acid-binding domain knock-in mutation provides evidence for nonclassical ER pathway signaling in vivo, Molecular Endocrinology, 16, 2188, 10.1210/me.2001-0174 Handgraaf, 2013, Prevention of obesity and insulin resistance by estrogens requires ERalpha activation function-2 (ERalphaAF-2), whereas ERalphaAF-1 is dispensable, Diabetes, 62, 4098, 10.2337/db13-0282 Saito, 2016, PI3K in the ventromedial hypothalamic nucleus mediates estrogenic actions on energy expenditure in female mice, Scientific Reports, 6, 23459, 10.1038/srep23459 Pedram, 2008, Developmental phenotype of a membrane only estrogen receptor alpha (MOER) mouse, Journal of Biological Chemistry Leibel, 1997, The molecular genetics of rodent single gene obesities, Journal of Biological Chemistry, 272, 31937, 10.1074/jbc.272.51.31937 Alingh Prins, 1986, Daily rhythms of feeding in the genetically obese and lean Zucker rats, Physiology and Behavior, 38, 423, 10.1016/0031-9384(86)90115-0 McLaughlin, 1981, Ontogeny of feeding behavior in the Zucker obese rat, Physiology and Behavior, 26, 607, 10.1016/0031-9384(81)90132-3 Trayhurn, 1977, Thermogenic defect in pre-obese ob/ob mice, Nature, 266, 60, 10.1038/266060a0 Dauncey, 1986, Activity-induced thermogenesis in lean and genetically obese (ob/ob) mice, Experientia, 42, 547, 10.1007/BF01946696 Dauncey, 1987, Role of activity-induced thermogenesis in twenty-four hour energy expenditure of lean and genetically obese (ob/ob) mice, Quarterly Journal of Experimental Physiology, 72, 549, 10.1113/expphysiol.1987.sp003096 Tartaglia, 1997, The leptin receptor, Journal of Biological Chemistry, 272, 6093, 10.1074/jbc.272.10.6093 Halaas, 1997, Physiological response to long-term peripheral and central leptin infusion in lean and obese mice, Proceedings of the National Academy of Sciences of the United States of America, 94, 8878, 10.1073/pnas.94.16.8878 Diano, 1998, Leptin receptors in estrogen receptor-containing neurons of the female rat hypothalamus, Brain Research, 812, 256, 10.1016/S0006-8993(98)00936-6 Lindell, 2001, Leptin receptor 5'untranslated regions in the rat: relative abundance, genomic organization and relation to putative response elements, Molecular and Cellular Endocrinology, 172, 37, 10.1016/S0303-7207(00)00382-8 Clegg, 2006, Gonadal hormones determine sensitivity to central leptin and insulin, Diabetes, 55, 978, 10.2337/diabetes.55.04.06.db05-1339 Raybould, 2007, Mechanisms of CCK signaling from gut to brain, Current Opinion in Pharmacology, 7, 570, 10.1016/j.coph.2007.09.006 Huang, 1993, Effect of cholecystokinin on food intake at different stages of the estrous cycle in female rats, Journal of the Association for Academic Minority Physicians, 4, 56 Asarian, 1999, Cyclic estradiol treatment phasically potentiates endogenous cholecystokinin's satiating action in ovariectomized rats, Peptides, 20, 445, 10.1016/S0196-9781(99)00024-8 Eckel, 1999, Endogenous cholecystokinin's satiating action increases during estrus in female rats, Peptides, 20, 451, 10.1016/S0196-9781(99)00025-X Geary, 1996, The increased satiating potency of CCK-8 by estradiol is not mediated by upregulation of NTS CCK receptors, Brain Research, 719, 179, 10.1016/0006-8993(96)00099-6 Butera, 1993, Modulation of the satiety effect of cholecystokinin by estradiol, Physiology and Behavior, 53, 1235, 10.1016/0031-9384(93)90387-U DiNardo, 1997, Distribution of fos-like immunoreactivity in the medullary reticular formation of the rat after gustatory elicited ingestion and rejection behaviors, Journal of Neuroscience, 17, 3826, 10.1523/JNEUROSCI.17-10-03826.1997 Eckel, 2001, Estradiol treatment increases feeding-induced c-Fos expression in the brains of ovariectomized rats, American Journal of Physiology Regulatory Integrative and Comparative Physiology, 281, R738, 10.1152/ajpregu.2001.281.3.R738 Rinaman, 1998, Medullary c-Fos activation in rats after ingestion of a satiating meal, American Journal of Physiology, 275, R262 Park, 1998, Neuroanatomical patterns of fos-like immunoreactivity induced by a palatable meal and meal-paired environment in saline- and naltrexone-treated rats, Brain Research, 805, 169, 10.1016/S0006-8993(98)00719-7 Rinaman, 1995, Cholecystokinin activates catecholaminergic neurons in the caudal medulla that innervate the paraventricular nucleus of the hypothalamus in rats, Journal of Comparative Neurology, 360, 246, 10.1002/cne.903600204 Li, 1994, Cholecystokinin- and dexfenfluramine-induced anorexia compared using devazepide and c-fos expression in the rat brain, Regulatory Peptides, 50, 223, 10.1016/0167-0115(94)90003-5 Eckel, 2002, Estradiol treatment increases CCK-induced c-Fos expression in the brains of ovariectomized rats, American Journal of Physiology Regulatory Integrative and Comparative Physiology, 283, R1378, 10.1152/ajpregu.00300.2002 Nakazato, 2001, A role for ghrelin in the central regulation of feeding, Nature, 409, 194, 10.1038/35051587 Tschop, 2000, Ghrelin induces adiposity in rodents, Nature, 407, 908, 10.1038/35038090 Wren, 2001, Ghrelin causes hyperphagia and obesity in rats, Diabetes, 50, 2540, 10.2337/diabetes.50.11.2540 Wren, 2001, Ghrelin enhances appetite and increases food intake in humans, The Journal of Cinical Endocrinology and Metabolism, 86, 5992, 10.1210/jcem.86.12.8111 Arnold, 2006, Gut vagal afferents are not necessary for the eating-stimulatory effect of intraperitoneally injected ghrelin in the rat, Journal of Neuroscience, 26, 11052, 10.1523/JNEUROSCI.2606-06.2006 Davidson, 2005, The interoceptive cue properties of ghrelin generalize to cues produced by food deprivation, Peptides, 26, 1602, 10.1016/j.peptides.2005.02.014 Clegg, 2007, Estradiol-dependent decrease in the orexigenic potency of ghrelin in female rats, Diabetes, 56, 1051, 10.2337/db06-0015 Elmquist, 1999, From lesions to leptin: hypothalamic control of food intake and body weight, Neuron, 22, 221, 10.1016/S0896-6273(00)81084-3 Cone, 2005, Anatomy and regulation of the central melanocortin system, Nature Neuroscience, 8, 571, 10.1038/nn1455 Williams, 2005, The melanocortin system as a central integrator of direct and indirect controls of food intake, American Journal of Physiology Regulatory Integrative and Comparative Physiology, 289, R2, 10.1152/ajpregu.00226.2005 Cone, 1999, The central melanocortin system and energy homeostasis, Trends in Endocrinology and Metabolism, 10, 211, 10.1016/S1043-2760(99)00153-8 Huszar, 1997, Targeted disruption of the melanocortin-4 receptor results in obesity in mice, Cell, 88, 131, 10.1016/S0092-8674(00)81865-6 Pelletier, 2007, Oestrogenic regulation of pro-opiomelanocortin, neuropeptide Y and corticotrophin-releasing hormone mRNAs in mouse hypothalamus, Journal of Neuroendocrinology, 19, 426, 10.1111/j.1365-2826.2007.01548.x Hirosawa, 2008, Ablation of estrogen receptor alpha (ERalpha) prevents upregulation of POMC by leptin and insulin, Biochemical and Biophysical Research Communications, 371, 320, 10.1016/j.bbrc.2008.04.073 Qiu, 2003, Rapid signaling of estrogen in hypothalamic neurons involves a novel G-protein-coupled estrogen receptor that activates protein kinase C, Journal of Neuroscience, 23, 9529, 10.1523/JNEUROSCI.23-29-09529.2003 Paez, 1991, Insatiable feeding evoked in rats by recurrent perfusion of neuropeptide Y in the hypothalamus, Peptides, 12, 609, 10.1016/0196-9781(91)90109-3 Pierroz, 1996, Chronic administration of neuropeptide Y into the lateral ventricle inhibits both the pituitary-testicular axis and growth hormone and insulin-like growth factor I secretion in intact adult male rats, Endocrinology, 137, 3, 10.1210/endo.137.1.8536627 Olofsson, 2009, Functional requirement of AgRP and NPY neurons in ovarian cycle-dependent regulation of food intake, Proceedings of the National Academy of Sciences of the United States of America, 106, 15932, 10.1073/pnas.0904747106 Lechin, 2006, Dorsal raphe vs. median raphe serotonergic antagonism. Anatomical, physiological, behavioral, neuroendocrinological, neuropharmacological and clinical evidences: relevance for neuropharmacological therapy, Progress In Neuro-Psychopharmacology and Biological Psychiatry, 30, 565, 10.1016/j.pnpbp.2005.11.025 De Fanti, 2001, Meal-induced changes in extracellular 5-HT in medial hypothalamus of lean (Fa/Fa) and obese (fa/fa) Zucker rats, Brain Research, 902, 164, 10.1016/S0006-8993(01)02371-X Rowland, 1986, Neurobiology of an anorectic drug: fenfluramine, Progress in Neurobiology, 27, 13, 10.1016/0301-0082(86)90011-0 Foltin, 1989, Food intake in baboons: effects of a long-acting cholecystokinin analog, Appetite, 12, 145, 10.1016/0195-6663(89)90103-7 McGuirk, 1991, Differential effects of d-fenfluramine, l-fenfluramine and d-amphetamine on the microstructure of human eating behaviour, Behavioural Pharmacology, 2, 113, 10.1097/00008877-199104000-00004 Rogers, 1979, Effect of anorexic drugs on food intake and the micro-structure of eating in human subjects, Psychopharmacology (Berl), 66, 159, 10.1007/BF00427624 Blundell, 1974, Central action of anorexic agents: effects of amphetamine and fenfluramine in rats with lateral hypothalamic lesions, European Journal of Pharmacology, 28, 81, 10.1016/0014-2999(74)90115-0 Geyer, 1976, Behavioral studies following lesions of the mesolimbic and mesostriatal serotonergic pathways, Brain Research, 106, 257, 10.1016/0006-8993(76)91024-6 Ghosh, 1973, The effect of cyproheptadine on water and food intake and on body weight in the fasted adult and weanling rats, British Journal of Pharmacology, 48 Saller, 1976, Hyperphagia and increased growth in rats after intraventricular injection of 5,7-dihydroxytryptamine, Science, 192, 385, 10.1126/science.1257774 Rivera, 2009, Estradiol increases Pet-1 and serotonin transporter mRNA in the midbrain raphe nuclei of ovariectomized rats, Brain Research, 1259, 51, 10.1016/j.brainres.2008.12.067 Rivera, 2012, Estradiol increases the anorexia associated with increased 5-HT(2C) receptor activation in ovariectomized rats, Physiology and Behavior, 105, 188, 10.1016/j.physbeh.2011.08.018 Callewaert, 2009, Differential regulation of bone and body composition in male mice with combined inactivation of androgen and estrogen receptor-alpha, The FASEB Journal, 23, 232, 10.1096/fj.08-113456 Grumbach, 1999, Estrogen: consequences and implications of human mutations in synthesis and action, Journal of Clinics in Endocrinology and Metabolism, 84, 4677 Smith, 1994, Estrogen resistance caused by a mutation in the estrogen-receptor gene in a man, New England Journal of Medicine, 331, 1056, 10.1056/NEJM199410203311604 Jones, 2000, Aromatase-deficient (ArKO) mice have a phenotype of increased adiposity, Proceedings of the National Academy of Sciences of the United States of America, 97, 12735, 10.1073/pnas.97.23.12735 Simpson, 2003, Sources of estrogen and their importance, The Journal of Steroid Biochemistry and Molecular Biology, 86, 225, 10.1016/S0960-0760(03)00360-1 Nelson, 2001, Estrogen production and action, Journal of the American Academy of Dermatology, 45, S116, 10.1067/mjd.2001.117432 Simpson, 2005, Estrogen–the good, the bad, and the unexpected, Endocrine Reviews, 26, 322, 10.1210/er.2004-0020 Wu, 2009, Estrogen masculinizes neural pathways and sex-specific behaviors, Cell, 139, 61, 10.1016/j.cell.2009.07.036 Xu, 2015, Estrogen receptor-alpha in medial amygdala neurons regulates body weight, Journal of Clinical Investigation, 125, 2861, 10.1172/JCI80941 Markakis, 2002, Development of the neuroendocrine hypothalamus, Frontiers in Neuroendocrinology, 23, 257, 10.1016/S0091-3022(02)00003-1 Grove, 2003, Ontogeny of the hypothalamic neuropeptide Y system, Physiology and Behavior, 79, 47, 10.1016/S0031-9384(03)00104-5 Rinaman, 2006, Ontogeny of hypothalamic-hindbrain feeding control circuits, Developmental Psychobiology, 48, 389, 10.1002/dev.20146 Rinaman, 2007, Visceral sensory inputs to the endocrine hypothalamus, Frontiers in Neuroendocrinology, 28, 50, 10.1016/j.yfrne.2007.02.002 McNay, 2011, Remodeling of the arcuate nucleus energy-balance circuit is inhibited in obese mice, Journal of Clinical Investigation Remmers, 2011, Developmental programming of energy balance and its hypothalamic regulation, Endocrine Reviews, 32, 272, 10.1210/er.2009-0028 MacLusky, 1979, The development of estrogen receptor systems in the rat brain: perinatal development, Brain Research, 178, 129, 10.1016/0006-8993(79)90093-3 Yokosuka, 1997, Postnatal development and sex difference in neurons containing estrogen receptor-alpha immunoreactivity in the preoptic brain, the diencephalon, and the amygdala in the rat, Journal of Comparative Neurology, 389, 81, 10.1002/(SICI)1096-9861(19971208)389:1<81::AID-CNE6>3.0.CO;2-A Rhoda, 1984, Gonadal steroid concentrations in serum and hypothalamus of the rat at birth: aromatization of testosterone to 17 beta-estradiol, Endocrinology, 114, 1754, 10.1210/endo-114-5-1754 Amateau, 2004, Brain estradiol content in newborn rats: sex differences, regional heterogeneity, and possible de novo synthesis by the female telencephalon, Endocrinology, 145, 2906, 10.1210/en.2003-1363 Bidlingmaier, 1987, Estrone and estradiol concentrations in human ovaries, testes, and adrenals during the first two years of life, The Journal of Cinical Endocrinology and Metabolism, 65, 862, 10.1210/jcem-65-5-862 Corpechot, 1981, Testosterone, dihydrotestosterone and androstanediols in plasma, testes and prostates of rats during development, Acta Endocrinologica (Copenh), 96, 127, 10.1530/acta.0.0960127 Seiki, 1989, Hormonal events surrounding spontaneous onset of puberty in female rats, Tokai Journal of Experimental and Clinical Medicine, 14, 45 Matsumoto, 1980, Sexual dimorphism in 'wiring pattern' in the hypothalamic arcuate nucleus and its modification by neonatal hormonal environment, Brain Research, 190, 238, 10.1016/0006-8993(80)91173-7 Nohara, 2011, Early-life exposure to testosterone programs the hypothalamic melanocortin system, Endocrinology, 152, 1661, 10.1210/en.2010-1288 Cederroth, 2009, Fetal programming of adult glucose homeostasis in mice, PLoS One, 4, e7281, 10.1371/journal.pone.0007281 Klump, 2017, The significant effects of puberty on the genetic diathesis of binge eating in girls, International Journal of Eating Disorders, 50, 984, 10.1002/eat.22727 Klump, 2012, The effects of puberty on genetic risk for disordered eating: evidence for a sex difference, Psychological Medicine, 42, 627, 10.1017/S0033291711001541