{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T17:00:48Z","timestamp":1785517248024,"version":"3.56.0"},"reference-count":86,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100017618","name":"Division of Diabetes, Endocrinology, and Metabolic Diseases","doi-asserted-by":"publisher","award":["DK092587"],"award-info":[{"award-number":["DK092587"]}],"id":[{"id":"10.13039\/100017618","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000135","name":"NIH Blueprint for Neuroscience Research","doi-asserted-by":"publisher","award":["AT011683"],"award-info":[{"award-number":["AT011683"]}],"id":[{"id":"10.13039\/100000135","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Comp Physiol B"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Energy expenditure (EE) is the dynamic physiological process that balances energy intake to maintain body weight and body temperature. Environmental changes induce EE adaptations that limit fluctuations in body weight or body temperature by balancing between body temperature defense and energy conservation. The central nervous system (CNS) orchestrates these adaptations by integrating multiple external and internal cues to coordinate thermogenesis together with other behavioral and metabolic processes. Key hypothalamic and brainstem nuclei form a distributed network that modulates autonomic output to peripheral organs, including sympathetic tone to brown and white adipose tissues. Here, we review known neural circuits and neurotransmitter systems, their integration with sensory pathways and functionally opposing output discrepancies that reveal knowledge gaps how the CNS switches between body temperature defense and energy preservation.<\/jats:p>","DOI":"10.1007\/s00360-026-01688-7","type":"journal-article","created":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T09:14:51Z","timestamp":1785143691000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Central circuits in control of energy expenditure: connecting energy and temperature homeostasis"],"prefix":"10.1007","author":[{"given":"Clint","family":"Kinney","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura","family":"Kaiser","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher D.","family":"Morrison","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hans-Rudolf","family":"Berthoud","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1152-4884","authenticated-orcid":false,"given":"Heike","family":"M\u00fcnzberg","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,27]]},"reference":[{"key":"1688_CR1","doi-asserted-by":"publisher","first-page":"6569","DOI":"10.1113\/JP274667","volume":"595","author":"SBG Abbott","year":"2017","unstructured":"Abbott SBG, Saper CB (2017) Median preoptic glutamatergic neurons promote thermoregulatory heat loss and water consumption in mice. J Physiol 595:6569\u20136583. https:\/\/doi.org\/10.1113\/JP274667","journal-title":"J Physiol"},{"key":"1688_CR2","doi-asserted-by":"publisher","first-page":"5813","DOI":"10.1073\/pnas.1015551108","volume":"108","author":"G Acosta-Galvan","year":"2011","unstructured":"Acosta-Galvan G, Yi C-X, van der Vliet J et al (2011) Interaction between hypothalamic dorsomedial nucleus and the suprachiasmatic nucleus determines intensity of food anticipatory behavior. Proc Natl Acad Sci U S A 108:5813\u20135818. https:\/\/doi.org\/10.1073\/pnas.1015551108","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1688_CR3","doi-asserted-by":"publisher","first-page":"313","DOI":"10.12688\/wellcomeopenres.17379.2","volume":"6","author":"M Ambler","year":"2021","unstructured":"Ambler M, Hitrec T, Pickering A (2021) Turn it off and on again: characteristics and control of torpor. Wellcome Open Res 6:313. https:\/\/doi.org\/10.12688\/wellcomeopenres.17379.2","journal-title":"Wellcome Open Res"},{"key":"1688_CR4","doi-asserted-by":"publisher","first-page":"4267","DOI":"10.1523\/JNEUROSCI.2102-21.2022","volume":"42","author":"M Ambler","year":"2022","unstructured":"Ambler M, Hitrec T, Wilson A et al (2022) Neurons in the dorsomedial hypothalamus promote, prolong, and deepen torpor in the mouse. J Neurosci off J Soc Neurosci 42:4267\u20134277. https:\/\/doi.org\/10.1523\/JNEUROSCI.2102-21.2022","journal-title":"J Neurosci off J Soc Neurosci"},{"key":"1688_CR5","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1038\/s41593-024-01830-0","volume":"28","author":"W Ambroziak","year":"2025","unstructured":"Ambroziak W, Nencini S, Pohle J et al (2025) Thermally induced neuronal plasticity in the hypothalamus mediates heat tolerance. Nat Neurosci 28:346\u2013360. https:\/\/doi.org\/10.1038\/s41593-024-01830-0","journal-title":"Nat Neurosci"},{"key":"1688_CR6","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1016\/j.cell.2005.08.035","volume":"123","author":"N Balthasar","year":"2005","unstructured":"Balthasar N, Dalgaard LT, Lee CE et al (2005) Divergence of melanocortin pathways in the control of food intake and energy expenditure. Cell 123:493\u2013505. https:\/\/doi.org\/10.1016\/j.cell.2005.08.035","journal-title":"Cell"},{"key":"1688_CR7","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1038\/nn.3737","volume":"17","author":"ED Berglund","year":"2014","unstructured":"Berglund ED, Liu T, Kong X et al (2014) Melanocortin 4 receptors in autonomic neurons regulate thermogenesis and glycemia. Nat Neurosci 17:911\u2013913. https:\/\/doi.org\/10.1038\/nn.3737","journal-title":"Nat Neurosci"},{"key":"1688_CR8","doi-asserted-by":"publisher","first-page":"R255","DOI":"10.1152\/ajpregu.2000.279.1.R255","volume":"279","author":"CL Buck","year":"2000","unstructured":"Buck CL, Barnes BM (2000) Effects of ambient temperature on metabolic rate, respiratory quotient, and torpor in an arctic hibernator. Am J Physiol Regul Integr Comp Physiol 279:R255-262. https:\/\/doi.org\/10.1152\/ajpregu.2000.279.1.R255","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"key":"1688_CR9","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1002\/cne.10643","volume":"460","author":"G Cano","year":"2003","unstructured":"Cano G, Passerin AM, Schiltz JC et al (2003) Anatomical substrates for the central control of sympathetic outflow to interscapular adipose tissue during cold exposure. J Comp Neurol 460:303\u2013326. https:\/\/doi.org\/10.1002\/cne.10643","journal-title":"J Comp Neurol"},{"key":"1688_CR10","doi-asserted-by":"publisher","first-page":"R33","DOI":"10.1016\/j.cub.2023.11.053","volume":"34","author":"M Cerri","year":"2024","unstructured":"Cerri M (2024) Torpor: a whole-brain view of the underlying neural network. Curr Biol CB 34:R33\u2013R35. https:\/\/doi.org\/10.1016\/j.cub.2023.11.053","journal-title":"Curr Biol CB"},{"key":"1688_CR11","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1016\/j.cmet.2011.02.019","volume":"13","author":"P-T Chao","year":"2011","unstructured":"Chao P-T, Yang L, Aja S et al (2011) Knockdown of NPY expression in the dorsomedial hypothalamus promotes development of brown adipocytes and prevents diet-induced obesity. Cell Metab 13:573\u2013583. https:\/\/doi.org\/10.1016\/j.cmet.2011.02.019","journal-title":"Cell Metab"},{"key":"1688_CR12","doi-asserted-by":"publisher","first-page":"829","DOI":"10.1016\/j.cell.2015.01.033","volume":"160","author":"Y Chen","year":"2015","unstructured":"Chen Y, Lin Y-C, Kuo T-W, Knight ZA (2015) Sensory detection of food rapidly modulates arcuate feeding circuits. Cell 160:829\u2013841. https:\/\/doi.org\/10.1016\/j.cell.2015.01.033","journal-title":"Cell"},{"key":"1688_CR13","doi-asserted-by":"publisher","first-page":"500","DOI":"10.1172\/JCI88622","volume":"127","author":"M Chen","year":"2017","unstructured":"Chen M, Shrestha YB, Podyma B et al (2017) Gs\u03b1 deficiency in the dorsomedial hypothalamus underlies obesity associated with Gs\u03b1 mutations. J Clin Invest 127:500\u2013510. https:\/\/doi.org\/10.1172\/JCI88622","journal-title":"J Clin Invest"},{"key":"1688_CR14","doi-asserted-by":"publisher","first-page":"33059","DOI":"10.1074\/jbc.M006328200","volume":"275","author":"SP Commins","year":"2000","unstructured":"Commins SP, Watson PM, Levin N et al (2000) Central leptin regulates the UCP1 and ob genes in brown and white adipose tissue via different beta-adrenoceptor subtypes. J Biol Chem 275:33059\u201333067. https:\/\/doi.org\/10.1074\/jbc.M006328200","journal-title":"J Biol Chem"},{"key":"1688_CR15","doi-asserted-by":"publisher","first-page":"e58764","DOI":"10.7554\/eLife.58764","volume":"9","author":"JD Deem","year":"2020","unstructured":"Deem JD, Faber CL, Pedersen C et al (2020) Cold-induced hyperphagia requires AgRP neuron activation in mice. Elife 9:e58764. https:\/\/doi.org\/10.7554\/eLife.58764","journal-title":"Elife"},{"key":"1688_CR16","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1146\/annurev-genet-121415-122024","volume":"50","author":"IA Droujinine","year":"2016","unstructured":"Droujinine IA, Perrimon N (2016) Interorgan communication pathways in physiology: focus on drosophila. Annu Rev Genet 50:539\u2013570. https:\/\/doi.org\/10.1146\/annurev-genet-121415-122024","journal-title":"Annu Rev Genet"},{"key":"1688_CR17","doi-asserted-by":"publisher","first-page":"1375","DOI":"10.1016\/s0896-6273(00)80656-x","volume":"21","author":"CF Elias","year":"1998","unstructured":"Elias CF, Lee C, Kelly J et al (1998) Leptin activates hypothalamic CART neurons projecting to the spinal cord. Neuron 21:1375\u20131385. https:\/\/doi.org\/10.1016\/s0896-6273(00)80656-x","journal-title":"Neuron"},{"key":"1688_CR18","doi-asserted-by":"publisher","first-page":"e63671","DOI":"10.7554\/eLife.63671","volume":"10","author":"CL Faber","year":"2021","unstructured":"Faber CL, Deem JD, Phan BA et al (2021) Leptin receptor neurons in the dorsomedial hypothalamus regulate diurnal patterns of feeding, locomotion, and metabolism. Elife 10:e63671. https:\/\/doi.org\/10.7554\/eLife.63671","journal-title":"Elife"},{"key":"1688_CR19","doi-asserted-by":"publisher","first-page":"5414","DOI":"10.1210\/en.2007-0655","volume":"148","author":"M Faouzi","year":"2007","unstructured":"Faouzi M, Leshan R, Bj\u00f6rnholm M et al (2007) Differential accessibility of circulating leptin to individual hypothalamic sites. Endocrinol 148:5414\u20135423. https:\/\/doi.org\/10.1210\/en.2007-0655","journal-title":"Endocrinol"},{"key":"1688_CR20","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1210\/er.2013-1087","volume":"35","author":"C Fekete","year":"2014","unstructured":"Fekete C, Lechan RM (2014) Central regulation of hypothalamic-pituitary-thyroid axis under physiological and pathophysiological conditions. Endocr Rev 35:159\u2013194. https:\/\/doi.org\/10.1210\/er.2013-1087","journal-title":"Endocr Rev"},{"key":"1688_CR21","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.jtherbio.2005.12.001","volume":"31","author":"CR Fetsch","year":"2006","unstructured":"Fetsch CR, Heideman PD, Griffin JD (2006) Effects of melatonin on thermally classified anterior hypothalamic neurons in the white-footed mouse (Peromyscus leucopus). J Therm Biol 31:40\u201349. https:\/\/doi.org\/10.1016\/j.jtherbio.2005.12.001","journal-title":"J Therm Biol"},{"key":"1688_CR22","doi-asserted-by":"publisher","first-page":"156100","DOI":"10.1016\/j.metabol.2024.156100","volume":"163","author":"M Francois","year":"2025","unstructured":"Francois M, Kaiser L, He Y et al (2025) Leptin receptor neurons in the dorsomedial hypothalamus require distinct neuronal subsets for thermogenesis and weight loss. Metabolism 163:156100. https:\/\/doi.org\/10.1016\/j.metabol.2024.156100","journal-title":"Metabolism"},{"key":"1688_CR23","doi-asserted-by":"publisher","first-page":"111380","DOI":"10.1016\/j.celrep.2022.111380","volume":"40","author":"A Fukushima","year":"2022","unstructured":"Fukushima A, Kataoka N, Nakamura K (2022) An oxytocinergic neural pathway that stimulates thermogenic and cardiac sympathetic outflow. Cell Rep 40:111380. https:\/\/doi.org\/10.1016\/j.celrep.2022.111380","journal-title":"Cell Rep"},{"key":"1688_CR24","doi-asserted-by":"publisher","first-page":"863","DOI":"10.1038\/nn.4011","volume":"18","author":"AS Garfield","year":"2015","unstructured":"Garfield AS, Li C, Madara JC et al (2015) A neural basis for melanocortin-4 receptor regulated appetite. Nat Neurosci 18:863\u2013871. https:\/\/doi.org\/10.1038\/nn.4011","journal-title":"Nat Neurosci"},{"key":"1688_CR25","doi-asserted-by":"publisher","first-page":"1628","DOI":"10.1038\/nn.4392","volume":"19","author":"AS Garfield","year":"2016","unstructured":"Garfield AS, Shah BP, Burgess CR et al (2016) Dynamic GABAergic afferent modulation of AgRP neurons. Nat Neurosci 19:1628\u20131635. https:\/\/doi.org\/10.1038\/nn.4392","journal-title":"Nat Neurosci"},{"key":"1688_CR26","doi-asserted-by":"publisher","first-page":"1460","DOI":"10.1002\/cne.22283","volume":"518","author":"JC Geerling","year":"2010","unstructured":"Geerling JC, Shin J-W, Chimenti PC, Loewy AD (2010) Paraventricular hypothalamic nucleus: axonal projections to the brainstem. J Comp Neurol 518:1460\u20131499. https:\/\/doi.org\/10.1002\/cne.22283","journal-title":"J Comp Neurol"},{"key":"1688_CR85","doi-asserted-by":"publisher","unstructured":"Nakamura K, Morrison SF (2010) A thermosensory pathway mediating heat-defense responses. Proc Natl Acad Sci U S A 107:8848\u20138853 https:\/\/doi.org\/10.1073\/pnas.0913358107","DOI":"10.1073\/pnas.0913358107"},{"key":"1688_CR27","doi-asserted-by":"publisher","first-page":"17686","DOI":"10.1074\/jbc.272.28.17686","volume":"272","author":"D Grujic","year":"1997","unstructured":"Grujic D, Susulic VS, Harper M-E et al (1997) \u03b23-Adrenergic receptors on white and brown adipocytes mediate \u03b23-selective agonist-induced effects on energy expenditure, insulin secretion, and food intake. J Biol Chem 272:17686\u201317693. https:\/\/doi.org\/10.1074\/jbc.272.28.17686","journal-title":"J Biol Chem"},{"key":"1688_CR28","doi-asserted-by":"publisher","first-page":"1433470","DOI":"10.3389\/fpubh.2024.1433470","volume":"12","author":"E Gutierrez","year":"2024","unstructured":"Gutierrez E, Garc\u00eda N, Carrera O (2024) Disordered eating in anorexia nervosa: give me heat, not just food. Front Public Health 12:1433470. https:\/\/doi.org\/10.3389\/fpubh.2024.1433470","journal-title":"Front Public Health"},{"key":"1688_CR29","doi-asserted-by":"publisher","first-page":"e2017218118","DOI":"10.1073\/pnas.2017218118","volume":"118","author":"J Houtz","year":"2021","unstructured":"Houtz J, Liao G-Y, An JJ, Xu B (2021) Discrete TrkB-expressing neurons of the dorsomedial hypothalamus regulate feeding and thermogenesis. Proc Natl Acad Sci U S A 118:e2017218118. https:\/\/doi.org\/10.1073\/pnas.2017218118","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1688_CR30","doi-asserted-by":"publisher","first-page":"zsab093","DOI":"10.1093\/sleep\/zsab093","volume":"44","author":"Y-G Huang","year":"2021","unstructured":"Huang Y-G, Flaherty SJ, Pothecary CA et al (2021) The relationship between fasting-induced torpor, sleep, and wakefulness in laboratory mice. Sleep 44:zsab093. https:\/\/doi.org\/10.1093\/sleep\/zsab093","journal-title":"Sleep"},{"key":"1688_CR31","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1016\/j.molmet.2015.03.006","volume":"4","author":"JH Jeong","year":"2015","unstructured":"Jeong JH, Lee DK, Blouet C et al (2015) Cholinergic neurons in the dorsomedial hypothalamus regulate mouse brown adipose tissue metabolism. Mol Metab 4:483\u2013492. https:\/\/doi.org\/10.1016\/j.molmet.2015.03.006","journal-title":"Mol Metab"},{"key":"1688_CR32","doi-asserted-by":"publisher","first-page":"e2004399","DOI":"10.1371\/journal.pbio.2004399","volume":"16","author":"JH Jeong","year":"2018","unstructured":"Jeong JH, Lee DK, Liu S-M et al (2018) Activation of temperature-sensitive TRPV1-like receptors in ARC POMC neurons reduces food intake. PLoS Biol 16:e2004399. https:\/\/doi.org\/10.1371\/journal.pbio.2004399","journal-title":"PLoS Biol"},{"key":"1688_CR33","doi-asserted-by":"publisher","first-page":"rs-7032725","DOI":"10.21203\/rs.3.rs-7032725\/v1","volume":"3","author":"L Kaiser","year":"2025","unstructured":"Kaiser L, Lee N, Williams J et al (2025) Obesogenic effects of warm temperature involve feeding adaptation by preoptic area leptin receptor neurons. Res Sq Rs. 3:rs-7032725. https:\/\/doi.org\/10.21203\/rs.3.rs-7032725\/v1","journal-title":"Res Sq Rs."},{"key":"1688_CR34","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1081\/CBI-100101036","volume":"17","author":"G K\u00f6rtner","year":"2000","unstructured":"K\u00f6rtner G, Geiser F (2000) The temporal organization of daily torpor and hibernation: circadian and circannual rhythms. Chronobiol Int 17:103\u2013128. https:\/\/doi.org\/10.1081\/CBI-100101036","journal-title":"Chronobiol Int"},{"key":"1688_CR35","doi-asserted-by":"publisher","first-page":"1424","DOI":"10.1172\/JCI46229","volume":"121","author":"MJ Krashes","year":"2011","unstructured":"Krashes MJ, Koda S, Ye C et al (2011) Rapid, reversible activation of AgRP neurons drives feeding behavior in mice. J Clin Invest 121:1424\u20131428. https:\/\/doi.org\/10.1172\/JCI46229","journal-title":"J Clin Invest"},{"key":"1688_CR36","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1016\/j.cmet.2013.03.004","volume":"17","author":"F Langlet","year":"2013","unstructured":"Langlet F, Levin BE, Luquet S et al (2013) Tanycytic VEGF-A boosts blood-hypothalamus barrier plasticity and access of metabolic signals to the arcuate nucleus in response to fasting. Cell Metab 17:607\u2013617. https:\/\/doi.org\/10.1016\/j.cmet.2013.03.004","journal-title":"Cell Metab"},{"key":"1688_CR37","doi-asserted-by":"publisher","first-page":"1131","DOI":"10.1038\/nn1949","volume":"10","author":"M Lazarus","year":"2007","unstructured":"Lazarus M, Yoshida K, Coppari R et al (2007) EP3 prostaglandin receptors in the median preoptic nucleus are critical for fever responses. Nat Neurosci 10:1131\u20131133. https:\/\/doi.org\/10.1038\/nn1949","journal-title":"Nat Neurosci"},{"key":"1688_CR38","doi-asserted-by":"publisher","first-page":"1891","DOI":"10.1002\/cne.23265","volume":"521","author":"SJ Lee","year":"2013","unstructured":"Lee SJ, Kirigiti M, Lindsley SR et al (2013) Efferent projections of neuropeptide Y-expressing neurons of the dorsomedial hypothalamus in chronic hyperphagic models. J Comp Neurol 521:1891\u20131914. https:\/\/doi.org\/10.1002\/cne.23265","journal-title":"J Comp Neurol"},{"key":"1688_CR39","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1080\/23328940.2021.1993734","volume":"9","author":"NLS Machado","year":"2022","unstructured":"Machado NLS, Saper CB (2022) Genetic identification of preoptic neurons that regulate body temperature in mice. Temp Austin Tex 9:14\u201322. https:\/\/doi.org\/10.1080\/23328940.2021.1993734","journal-title":"Temp Austin Tex"},{"key":"1688_CR40","doi-asserted-by":"publisher","first-page":"2291","DOI":"10.1016\/j.cub.2018.05.064","volume":"28","author":"NLS Machado","year":"2018","unstructured":"Machado NLS, Abbott SBG, Resch J et al (2018) A glutamatergic hypothalamomedullary circuit mediates thermogenesis but not heat conservation during stress-induced hyperthermia. Curr Biol CB 28:2291-2301.e5. https:\/\/doi.org\/10.1016\/j.cub.2018.05.064","journal-title":"Curr Biol CB"},{"key":"1688_CR41","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-025-09056-1","author":"N Machado","year":"2025","unstructured":"Machado N, Lynch N, Costa L et al (2025) Preoptic EP3R neurons constitute a two-way switch for fever and torpor. Nature. https:\/\/doi.org\/10.1038\/s41586-025-09056-1","journal-title":"Nature"},{"key":"1688_CR42","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1016\/s0306-4522(03)00527-x","volume":"122","author":"CJ Madden","year":"2003","unstructured":"Madden CJ, Morrison SF (2003) Excitatory amino acid receptor activation in the raphe pallidus area mediates prostaglandin-evoked thermogenesis. Neuroscience 122:5\u201315. https:\/\/doi.org\/10.1016\/s0306-4522(03)00527-x","journal-title":"Neuroscience"},{"key":"1688_CR43","doi-asserted-by":"publisher","first-page":"R831","DOI":"10.1152\/ajpregu.91007.2008","volume":"296","author":"CJ Madden","year":"2009","unstructured":"Madden CJ, Morrison SF (2009) Neurons in the paraventricular nucleus of the hypothalamus inhibit sympathetic outflow to brown adipose tissue. Am J Physiol - Regul Integr Comp Physiol 296:R831\u2013R843. https:\/\/doi.org\/10.1152\/ajpregu.91007.2008","journal-title":"Am J Physiol - Regul Integr Comp Physiol"},{"key":"1688_CR44","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/j.neulet.2018.11.027","volume":"696","author":"CJ Madden","year":"2019","unstructured":"Madden CJ, Morrison SF (2019) Central nervous system circuits that control body temperature. Neurosci Lett 696:225\u2013232. https:\/\/doi.org\/10.1016\/j.neulet.2018.11.027","journal-title":"Neurosci Lett"},{"key":"1688_CR45","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1016\/s0196-9781(00)00359-4","volume":"22","author":"V Mart\u00ednez","year":"2001","unstructured":"Mart\u00ednez V, Wang L, Tach\u00e9 Y (2001) Central TRH receptor 1 antisense blocks cold-induced gastric emptying but not brain c-Fos induction. Peptides 22:81\u201390. https:\/\/doi.org\/10.1016\/s0196-9781(00)00359-4","journal-title":"Peptides"},{"key":"1688_CR46","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau5324","volume":"362","author":"JR Moffitt","year":"2018","unstructured":"Moffitt JR, Bambah-Mukku D, Eichhorn SW et al (2018) Molecular, spatial, and functional single-cell profiling of the hypothalamic preoptic region. Sci 362:eaau5324. https:\/\/doi.org\/10.1126\/science.aau5324","journal-title":"Sci"},{"key":"1688_CR47","doi-asserted-by":"publisher","first-page":"3448","DOI":"10.1210\/en.2013-2092","volume":"155","author":"B Monge-Roffarello","year":"2014","unstructured":"Monge-Roffarello B, Labbe SM, Roy M-C et al (2014) The PVH as a site of CB1-mediated stimulation of thermogenesis by MC4R agonism in male rats. Endocrinol 155:3448\u20133458. https:\/\/doi.org\/10.1210\/en.2013-2092","journal-title":"Endocrinol"},{"key":"1688_CR48","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1146\/annurev-physiol-020518-114546","volume":"81","author":"SF Morrison","year":"2019","unstructured":"Morrison SF, Nakamura K (2019) Central mechanisms for thermoregulation. Annu Rev Physiol 81:285\u2013308. https:\/\/doi.org\/10.1146\/annurev-physiol-020518-114546","journal-title":"Annu Rev Physiol"},{"key":"1688_CR49","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1152\/physrev.00030.2013","volume":"94","author":"R Mullur","year":"2014","unstructured":"Mullur R, Liu Y-Y, Brent GA (2014) Thyroid hormone regulation of metabolism. Physiol Rev 94:355\u2013382. https:\/\/doi.org\/10.1152\/physrev.00030.2013","journal-title":"Physiol Rev"},{"key":"1688_CR50","doi-asserted-by":"publisher","first-page":"451","DOI":"10.12688\/f1000research.24260.1","volume":"9","author":"H M\u00fcnzberg","year":"2020","unstructured":"M\u00fcnzberg H, Singh P, Heymsfield SB et al (2020) Recent advances in understanding the role of leptin in energy homeostasis. F1000Res 9:451. https:\/\/doi.org\/10.12688\/f1000research.24260.1","journal-title":"F1000Res"},{"key":"1688_CR51","doi-asserted-by":"publisher","first-page":"R127","DOI":"10.1152\/ajpregu.00427.2006","volume":"292","author":"K Nakamura","year":"2007","unstructured":"Nakamura K, Morrison SF (2007) Central efferent pathways mediating skin cooling-evoked sympathetic thermogenesis in brown adipose tissue. Am J Physiol Regul Integr Comp Physiol 292:R127-136. https:\/\/doi.org\/10.1152\/ajpregu.00427.2006","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"key":"1688_CR52","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.neures.2004.09.007","volume":"51","author":"K Nakamura","year":"2005","unstructured":"Nakamura K, Matsumura K, Kobayashi S, Kaneko T (2005) Sympathetic premotor neurons mediating thermoregulatory functions. Neurosci Res 51:1\u20138. https:\/\/doi.org\/10.1016\/j.neures.2004.09.007","journal-title":"Neurosci Res"},{"key":"1688_CR53","doi-asserted-by":"publisher","first-page":"322","DOI":"10.1016\/j.cmet.2016.12.002","volume":"25","author":"Y Nakamura","year":"2017","unstructured":"Nakamura Y, Yanagawa Y, Morrison SF, Nakamura K (2017) Medullary reticular neurons mediate neuropeptide y-induced metabolic inhibition and mastication. Cell Metab 25:322\u2013334. https:\/\/doi.org\/10.1016\/j.cmet.2016.12.002","journal-title":"Cell Metab"},{"key":"1688_CR54","doi-asserted-by":"publisher","first-page":"eadd5463","DOI":"10.1126\/sciadv.add5463","volume":"8","author":"Y Nakamura","year":"2022","unstructured":"Nakamura Y, Yahiro T, Fukushima A et al (2022) Prostaglandin EP3 receptor-expressing preoptic neurons bidirectionally control body temperature via tonic GABAergic signaling. Sci Adv 8:eadd5463. https:\/\/doi.org\/10.1126\/sciadv.add5463","journal-title":"Sci Adv"},{"key":"1688_CR55","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1007\/s00360-010-0503-9","volume":"181","author":"R Oelkrug","year":"2011","unstructured":"Oelkrug R, Heldmaier G, Meyer CW (2011) Torpor patterns, arousal rates, and temporal organization of torpor entry in wildtype and UCP1-ablated mice. J Comp Physiol [b] 181:137\u2013145. https:\/\/doi.org\/10.1007\/s00360-010-0503-9","journal-title":"J Comp Physiol [b]"},{"key":"1688_CR56","doi-asserted-by":"publisher","first-page":"e202400190","DOI":"10.1002\/bies.202400190","volume":"47","author":"A Ohba","year":"2025","unstructured":"Ohba A, Yamaguchi H (2025) The art of chilling out: how neurons regulate torpor. BioEssays News Rev Mol Cell Dev Biol 47:e202400190. https:\/\/doi.org\/10.1002\/bies.202400190","journal-title":"BioEssays News Rev Mol Cell Dev Biol"},{"key":"1688_CR57","doi-asserted-by":"publisher","first-page":"515","DOI":"10.1016\/s0306-4522(01)00555-3","volume":"110","author":"BJ Oldfield","year":"2002","unstructured":"Oldfield BJ, Giles ME, Watson A et al (2002) The neurochemical characterisation of hypothalamic pathways projecting polysynaptically to brown adipose tissue in the rat. Neuroscience 110:515\u2013526. https:\/\/doi.org\/10.1016\/s0306-4522(01)00555-3","journal-title":"Neuroscience"},{"key":"1688_CR58","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1016\/0006-8993(86)90338-0","volume":"373","author":"M Palkovits","year":"1986","unstructured":"Palkovits M, Mezey E, Eskay RL, Brownstein MJ (1986) Innervation of the nucleus of the solitary tract and the dorsal vagal nucleus by thyrotropin-releasing hormone-containing raphe neurons. Brain Res 373:246\u2013251. https:\/\/doi.org\/10.1016\/0006-8993(86)90338-0","journal-title":"Brain Res"},{"key":"1688_CR59","doi-asserted-by":"publisher","first-page":"1530","DOI":"10.1038\/s41593-018-0249-3","volume":"21","author":"RA Pi\u00f1ol","year":"2018","unstructured":"Pi\u00f1ol RA, Zahler SH, Li C et al (2018) Brs3 neurons in the mouse dorsomedial hypothalamus regulate body temperature, energy expenditure and heart rate, but not food intake. Nat Neurosci 21:1530\u20131540. https:\/\/doi.org\/10.1038\/s41593-018-0249-3","journal-title":"Nat Neurosci"},{"key":"1688_CR60","doi-asserted-by":"publisher","first-page":"1389","DOI":"10.1016\/j.cmet.2021.05.001","volume":"33","author":"RA Pi\u00f1ol","year":"2021","unstructured":"Pi\u00f1ol RA, Mogul AS, Hadley CK et al (2021) Preoptic BRS3 neurons increase body temperature and heart rate via multiple pathways. Cell Metab 33:1389-1403.e6. https:\/\/doi.org\/10.1016\/j.cmet.2021.05.001","journal-title":"Cell Metab"},{"key":"1688_CR61","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1016\/j.molmet.2014.07.008","volume":"3","author":"K Rezai-Zadeh","year":"2014","unstructured":"Rezai-Zadeh K, Yu S, Jiang Y et al (2014) Leptin receptor neurons in the dorsomedial hypothalamus are key regulators of energy expenditure and body weight, but not food intake. Molecular Metabolism 3:681\u2013693. https:\/\/doi.org\/10.1016\/j.molmet.2014.07.008","journal-title":"Molecular Metabolism"},{"key":"1688_CR62","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1016\/j.cmet.2017.09.004","volume":"26","author":"R Schreiber","year":"2017","unstructured":"Schreiber R, Diwoky C, Schoiswohl G et al (2017) Cold-induced thermogenesis depends on ATGL-mediated lipolysis in cardiac muscle, but not brown adipose tissue. Cell Metab 26:753-763.e7. https:\/\/doi.org\/10.1016\/j.cmet.2017.09.004","journal-title":"Cell Metab"},{"key":"1688_CR63","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1038\/35007534","volume":"404","author":"MW Schwartz","year":"2000","unstructured":"Schwartz MW, Woods SC, Porte D et al (2000) Central nervous system control of food intake. Nature 404:661\u2013671. https:\/\/doi.org\/10.1038\/35007534","journal-title":"Nature"},{"key":"1688_CR64","doi-asserted-by":"publisher","first-page":"13193","DOI":"10.1073\/pnas.1407843111","volume":"111","author":"BP Shah","year":"2014","unstructured":"Shah BP, Vong L, Olson DP et al (2014) MC4R-expressing glutamatergic neurons in the paraventricular hypothalamus regulate feeding and are synaptically connected to the parabrachial nucleus. Proc Natl Acad Sci U S A 111:13193\u201313198. https:\/\/doi.org\/10.1073\/pnas.1407843111","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1688_CR65","doi-asserted-by":"publisher","first-page":"764","DOI":"10.1016\/j.cmet.2017.09.002","volume":"26","author":"H Shin","year":"2017","unstructured":"Shin H, Ma Y, Chanturiya T et al (2017) Lipolysis in brown adipocytes is not essential for cold-induced thermogenesis in mice. Cell Metab 26:764-777.e5. https:\/\/doi.org\/10.1016\/j.cmet.2017.09.002","journal-title":"Cell Metab"},{"issue":"2","key":"1688_CR66","doi-asserted-by":"publisher","first-page":"R417","DOI":"10.1152\/ajpregu.00174.2008","volume":"295","author":"CK Song","year":"2008","unstructured":"Song CK, Vaughan CH, Keen-Rhinehart E et al (2008) Melanocortin-4 receptor mRNA expressed in sympathetic outflow neurons to brown adipose tissue: neuroanatomical and functional evidence. Am J Physiol Regul Integr Comp Physiol 295(2):R417. https:\/\/doi.org\/10.1152\/ajpregu.00174.2008","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"key":"1688_CR67","doi-asserted-by":"publisher","first-page":"1393","DOI":"10.1126\/science.aaf7537","volume":"353","author":"K Song","year":"2016","unstructured":"Song K, Wang H, Kamm GB et al (2016) The TRPM2 channel is a hypothalamic heat sensor that limits fever and can drive hypothermia. Sci 353:1393\u20131398. https:\/\/doi.org\/10.1126\/science.aaf7537","journal-title":"Sci"},{"key":"1688_CR68","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1002\/cne.20742","volume":"492","author":"RL Stornetta","year":"2005","unstructured":"Stornetta RL, Rosin DL, Simmons JR et al (2005) Coexpression of vesicular glutamate transporter-3 and \u03b3-aminobutyric acidergic markers in rat rostral medullary raphe and intermediolateral cell column. J Comp Neurol 492:477\u2013494. https:\/\/doi.org\/10.1002\/cne.20742","journal-title":"J Comp Neurol"},{"key":"1688_CR69","doi-asserted-by":"publisher","first-page":"15306","DOI":"10.1523\/JNEUROSCI.0226-14.2014","volume":"34","author":"AK Sutton","year":"2014","unstructured":"Sutton AK, Pei H, Burnett KH et al (2014) Control of food intake and energy expenditure by Nos1 neurons of the paraventricular hypothalamus. J Neurosci 34:15306\u201315318. https:\/\/doi.org\/10.1523\/JNEUROSCI.0226-14.2014","journal-title":"J Neurosci"},{"key":"1688_CR70","doi-asserted-by":"publisher","first-page":"410","DOI":"10.1159\/000123111","volume":"31","author":"LW Swanson","year":"1980","unstructured":"Swanson LW, Sawchenko PE (1980) Paraventricular nucleus: a site for the integration of neuroendocrine and autonomic mechanisms. Neuroendocrinol 31:410\u2013417. https:\/\/doi.org\/10.1159\/000123111","journal-title":"Neuroendocrinol"},{"key":"1688_CR71","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1523\/JNEUROSCI.3721-05.2006","volume":"26","author":"SJ Swoap","year":"2006","unstructured":"Swoap SJ, Gutilla MJ, Liles LC et al (2006) The full expression of fasting-induced torpor requires beta 3-adrenergic receptor signaling. J Neurosci off J Soc Neurosci 26:241\u2013245. https:\/\/doi.org\/10.1523\/JNEUROSCI.3721-05.2006","journal-title":"J Neurosci off J Soc Neurosci"},{"key":"1688_CR72","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1038\/s41586-020-2163-6","volume":"583","author":"TM Takahashi","year":"2020","unstructured":"Takahashi TM, Sunagawa GA, Soya S et al (2020) A discrete neuronal circuit induces a hibernation-like state in rodents. Nature 583:109\u2013114. https:\/\/doi.org\/10.1038\/s41586-020-2163-6","journal-title":"Nature"},{"key":"1688_CR73","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.cell.2016.08.028","volume":"167","author":"CL Tan","year":"2016","unstructured":"Tan CL, Cooke EK, Leib DE et al (2016) Warm-Sensitive Neurons that Control Body Temperature. Cell 167:47-59.e15. https:\/\/doi.org\/10.1016\/j.cell.2016.08.028","journal-title":"Cell"},{"key":"1688_CR74","doi-asserted-by":"publisher","first-page":"1550","DOI":"10.1210\/en.2006-1389","volume":"148","author":"A Voss-Andreae","year":"2007","unstructured":"Voss-Andreae A, Murphy JG, Ellacott KLJ et al (2007) Role of the central melanocortin circuitry in adaptive thermogenesis of brown adipose tissue. Endocrinol 148:1550\u20131560. https:\/\/doi.org\/10.1210\/en.2006-1389","journal-title":"Endocrinol"},{"key":"1688_CR86","doi-asserted-by":"publisher","unstructured":"Yahiro T, Kataoka N, Nakamura K (2023) Two Ascending Thermosensory Pathways from the Lateral Parabrachial Nucleus That Mediate Behavioral and Autonomous Thermoregulation. J Neurosci Off J Soc Neurosci 43:5221\u20135240 https:\/\/doi.org\/10.1523\/JNEUROSCI.0643-23.2023","DOI":"10.1523\/JNEUROSCI.0643-23.2023"},{"key":"1688_CR75","doi-asserted-by":"publisher","first-page":"5381","DOI":"10.1016\/j.cub.2023.10.076","volume":"33","author":"H Yamaguchi","year":"2023","unstructured":"Yamaguchi H, Murphy KR, Fukatsu N et al (2023) Dorsomedial and preoptic hypothalamic circuits control torpor. Curr Biol CB 33:5381-5389.e4. https:\/\/doi.org\/10.1016\/j.cub.2023.10.076","journal-title":"Curr Biol CB"},{"key":"1688_CR76","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2021.109502","volume":"36","author":"S Yang","year":"2021","unstructured":"Yang S, Tan YL, Wu X et al (2021) An mPOA-ARCAgRP pathway modulates cold-evoked eating behavior. Cell Rep 36:109502. https:\/\/doi.org\/10.1016\/j.celrep.2021.109502","journal-title":"Cell Rep"},{"key":"1688_CR77","doi-asserted-by":"publisher","first-page":"5034","DOI":"10.1523\/JNEUROSCI.0213-16.2016","volume":"36","author":"S Yu","year":"2016","unstructured":"Yu S, Qualls-Creekmore E, Rezai-Zadeh K et al (2016) Glutamatergic Preoptic Area Neurons That Express Leptin Receptors Drive Temperature-Dependent Body Weight Homeostasis. J Neurosci off J Soc Neurosci 36:5034\u20135046. https:\/\/doi.org\/10.1523\/JNEUROSCI.0213-16.2016","journal-title":"J Neurosci off J Soc Neurosci"},{"key":"1688_CR78","doi-asserted-by":"publisher","first-page":"e33505","DOI":"10.7554\/eLife.33505","volume":"7","author":"S Yu","year":"2018","unstructured":"Yu S, Cheng H, Fran\u00e7ois M et al (2018) Preoptic leptin signaling modulates energy balance independent of body temperature regulation. Elife 7:e33505. https:\/\/doi.org\/10.7554\/eLife.33505","journal-title":"Elife"},{"key":"1688_CR79","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/s0006-8993(01)03369-8","volume":"928","author":"MV Zaretskaia","year":"2002","unstructured":"Zaretskaia MV, Zaretsky DV, Shekhar A, DiMicco JA (2002) Chemical stimulation of the dorsomedial hypothalamus evokes non-shivering thermogenesis in anesthetized rats. Brain Res 928:113\u2013125. https:\/\/doi.org\/10.1016\/s0006-8993(01)03369-8","journal-title":"Brain Res"},{"key":"1688_CR80","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/s0304-3940(03)00047-8","volume":"340","author":"MV Zaretskaia","year":"2003","unstructured":"Zaretskaia MV, Zaretsky DV, DiMicco JA (2003) Role of the dorsomedial hypothalamus in thermogenesis and tachycardia caused by microinjection of prostaglandin E2 into the preoptic area in anesthetized rats. Neurosci Lett 340:1\u20134. https:\/\/doi.org\/10.1016\/s0304-3940(03)00047-8","journal-title":"Neurosci Lett"},{"key":"1688_CR81","doi-asserted-by":"publisher","first-page":"1873","DOI":"10.1523\/JNEUROSCI.3223-10.2011","volume":"31","author":"Y Zhang","year":"2011","unstructured":"Zhang Y, Kerman IA, Laque A et al (2011) Leptin-receptor-expressing neurons in the dorsomedial hypothalamus and median preoptic area regulate sympathetic brown adipose tissue circuits. J Neurosci off J Soc Neurosci 31:1873\u20131884. https:\/\/doi.org\/10.1523\/JNEUROSCI.3223-10.2011","journal-title":"J Neurosci off J Soc Neurosci"},{"key":"1688_CR82","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1038\/s41586-020-2683-0","volume":"585","author":"KX Zhang","year":"2020","unstructured":"Zhang KX, D\u2019Souza S, Upton BA et al (2020a) Violet-light suppression of thermogenesis by opsin 5 hypothalamic neurons. Nature 585:420\u2013425. https:\/\/doi.org\/10.1038\/s41586-020-2683-0","journal-title":"Nature"},{"key":"1688_CR83","doi-asserted-by":"publisher","first-page":"6378","DOI":"10.1038\/s41467-020-20050-1","volume":"11","author":"Z Zhang","year":"2020","unstructured":"Zhang Z, Reis FMCV, He Y et al (2020b) Estrogen-sensitive medial preoptic area neurons coordinate torpor in mice. Nat Commun 11:6378. https:\/\/doi.org\/10.1038\/s41467-020-20050-1","journal-title":"Nat Commun"},{"key":"1688_CR84","doi-asserted-by":"publisher","unstructured":"Boulant JA, Dean JB (1986) Temperature receptors in the central nervous system. Annu Rev Physiol 48:639\u2013654 https:\/\/doi.org\/10.1146\/annurev.ph.48.030186.003231","DOI":"10.1146\/annurev.ph.48.030186.003231"}],"container-title":["Journal of Comparative Physiology B"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00360-026-01688-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00360-026-01688-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00360-026-01688-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T16:13:33Z","timestamp":1785514413000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00360-026-01688-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,27]]},"references-count":86,"alternative-id":["1688"],"URL":"https:\/\/doi.org\/10.1007\/s00360-026-01688-7","relation":{},"ISSN":["0174-1578","1432-136X"],"issn-type":[{"value":"0174-1578","type":"print"},{"value":"1432-136X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,27]]},"assertion":[{"value":"9 January 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"4 June 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Since submission of this manuscript, H.M. has provided consulting services to Novo Nordisk. The other authors declare no conflicts of interest relevant to the content of this paper.","order":1,"name":"Ethics","label":"Conflict of Interest","group":{"name":"EthicsHeading","label":"Declarations"}}]}}