{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T18:40:10Z","timestamp":1790275210258,"version":"4.1.0"},"reference-count":56,"publisher":"The Royal Society","issue":"1959","license":[{"start":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T00:00:00Z","timestamp":1790208000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,24]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>All animals exhibit complex behaviours that share features with those that in humans we call \u2018emotional expressions\u2019. Sidestepping contentious issues about the conscious experience of emotion, their expression across species provides a wealth of detail about their emergence in evolution, the shared features that distinguish them from reflexes, and principles for their neural control. This paper provides a high-level survey of this vast field.<\/jats:p>\n                  <jats:p>This article is part of the theme issue \u2018Mechanisms, development, phylogeny and functions of emotional expressions\u2019.<\/jats:p>","DOI":"10.1098\/rstb.2025.0135","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T07:08:09Z","timestamp":1790233689000},"source":"Crossref","is-referenced-by-count":1,"title":["Comparative neuroanatomy of emotion expression: from flies to humans"],"prefix":"10.1098","volume":"381","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8053-9692","authenticated-orcid":false,"given":"Ralph","family":"Adolphs","sequence":"first","affiliation":[{"name":"Department of Biology and Biological Engineering, California Institute of Technology , Pasadena, CA 91125, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,9,24]]},"reference":[{"key":"2026092403080094300_RSTB20250135C1","doi-asserted-by":"crossref","DOI":"10.1037\/10001-000","volume-title":"The expression of the emotions in man and animals,","author":"Darwin","year":"1872","edition":"3rd edn"},{"key":"2026092403080094300_RSTB20250135C2","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780195096736.001.0001","volume-title":"Affective neuroscience: the foundations of human and animal emotions","author":"Panksepp","year":"1998"},{"key":"2026092403080094300_RSTB20250135C3","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/9780262019569.001.0001","volume-title":"The cognitive-emotional brain: from interactions to integration","author":"Pessoa","year":"2013"},{"key":"2026092403080094300_RSTB20250135C4","doi-asserted-by":"crossref","first-page":"1070","DOI":"10.1016\/j.cub.2011.05.017","article-title":"Agitated honeybees exhibit pessimistic cognitive biases","volume":"21","author":"Bateson","year":"2011","journal-title":"Curr. Biol."},{"key":"2026092403080094300_RSTB20250135C5","doi-asserted-by":"crossref","first-page":"105475","DOI":"10.1016\/j.neubiorev.2023.105475","article-title":"Emotion in action: when emotions meet motor circuits","volume":"155","author":"Braine","year":"2023","journal-title":"Neurosci. Biobehav. Rev."},{"key":"2026092403080094300_RSTB20250135C6","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1093\/scan\/nsw153","article-title":"How should neuroscience study emotions? By distinguishing emotion states, concepts and experiences","volume":"12","author":"Adolphs","year":"2017","journal-title":"Soc. Cognit. Affect. Neurosci."},{"key":"2026092403080094300_RSTB20250135C7","volume-title":"The neuroscience of emotion: a new synthesis","author":"Adolphs","year":"2018"},{"key":"2026092403080094300_RSTB20250135C8","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.conb.2021.01.003","article-title":"Expressions of emotions across species","volume":"68","author":"Zych","year":"2021","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026092403080094300_RSTB20250135C9","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1016\/j.cell.2014.03.003","article-title":"A framework for studying emotions across species","volume":"157","author":"Anderson","year":"2014","journal-title":"Cell"},{"key":"2026092403080094300_RSTB20250135C10","volume-title":"The new science of emotion: mind, brain and behavior","author":"Adolphs","year":"2027"},{"key":"2026092403080094300_RSTB20250135C11","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1038\/nrn3301","article-title":"The many paths to fear","volume":"13","author":"Gross","year":"2012","journal-title":"Nat. Rev. Neurosci."},{"key":"2026092403080094300_RSTB20250135C12","doi-asserted-by":"crossref","first-page":"111222","DOI":"10.1016\/j.celrep.2022.111222","article-title":"A central alarm system that gates multi-sensory innate threat cues to the amygdala","volume":"40","author":"Kang","year":"2022","journal-title":"Cell Rep."},{"key":"2026092403080094300_RSTB20250135C13","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1038\/s41586-018-0244-6","article-title":"A synaptic threshold mechanism for computing escape decisions","volume":"558","author":"Evans","year":"2018","journal-title":"Nature"},{"key":"2026092403080094300_RSTB20250135C14","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1093\/cercor\/bhac109","article-title":"Rapid processing of threatening faces in the amygdala of nonhuman primates: subcortical inputs and dual roles","volume":"33","author":"Inagaki","year":"2023","journal-title":"Cereb. Cortex"},{"key":"2026092403080094300_RSTB20250135C15","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1038\/s41583-020-0350-y","article-title":"Multidimensional processing in the amygdala","volume":"21","author":"Gothard","year":"2020","journal-title":"Nat. Rev. Neurosci."},{"key":"2026092403080094300_RSTB20250135C16","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1016\/S0079-6123(08)61871-3","article-title":"Columnar organization in the midbrain periaqueductal gray and the integration of emotional expression","volume":"107","author":"Bandler","year":"1996","journal-title":"Progress in Brain Research"},{"key":"2026092403080094300_RSTB20250135C17","first-page":"447","article-title":"Integrated cardio-behavioral responses to threat define defensive states","volume":"26","author":"Signoret-Genest","year":"2023","journal-title":"Nat. Neurosci."},{"key":"2026092403080094300_RSTB20250135C18","doi-asserted-by":"crossref","first-page":"e74860","DOI":"10.7554\/eLife.74860","article-title":"Proposing a neural framework for the evolution of elaborate courtship displays","volume":"11","author":"Schwark","year":"2022","journal-title":"eLife"},{"key":"2026092403080094300_RSTB20250135C19","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1016\/j.conb.2013.01.017","article-title":"Neuroscience of affect: brain mechanisms of pleasure and displeasure","volume":"23","author":"Berridge","year":"2013","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026092403080094300_RSTB20250135C20","doi-asserted-by":"crossref","first-page":"903857","DOI":"10.3389\/fpsyg.2022.903857","article-title":"Birdsong and the neural regulation of positive emotion","volume":"13","author":"Riters","year":"2022","journal-title":"Front. Psychol."},{"key":"2026092403080094300_RSTB20250135C21","doi-asserted-by":"crossref","first-page":"3856","DOI":"10.1242\/jeb.151308","article-title":"Studying emotion in invertebrates: what has been done, what can be measured, and what they can provide","volume":"220","author":"Perry","year":"2017","journal-title":"J. Exp. Biol."},{"key":"2026092403080094300_RSTB20250135C22","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1017\/S0140525X00059057","article-title":"The command neuron concept","volume":"1","author":"Kupfermann","year":"1978","journal-title":"Behav. Brain Sci."},{"key":"2026092403080094300_RSTB20250135C23","doi-asserted-by":"crossref","first-page":"2237","DOI":"10.1111\/brv.12891","article-title":"A synthesis of deimatic behavior","volume":"97","author":"Drinkwater","year":"2022","journal-title":"Biol. Rev."},{"key":"2026092403080094300_RSTB20250135C24","doi-asserted-by":"crossref","first-page":"R1119","DOI":"10.1016\/j.cub.2023.09.008","article-title":"Embodied mechanisms of motor control in the octopus","volume":"33","author":"Hochner","year":"2023","journal-title":"Curr. Biol."},{"key":"2026092403080094300_RSTB20250135C25","doi-asserted-by":"crossref","first-page":"R1095","DOI":"10.1016\/j.cub.2023.08.095","article-title":"Neural control of cephalopod camouflage","volume":"33","author":"Montague","year":"2023","journal-title":"Curr. Biol."},{"key":"2026092403080094300_RSTB20250135C26","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1038\/s41586-018-0591-3","article-title":"Elucidating the control and development of skin patterning in cuttlefish","volume":"562","author":"Reiter","year":"2018","journal-title":"Nature"},{"key":"2026092403080094300_RSTB20250135C27","doi-asserted-by":"crossref","first-page":"e58942","DOI":"10.7554\/eLife.58942","article-title":"Cell types and neuronal circuitry underlying female aggression in Drosophila","volume":"9","author":"Schretter","year":"2020","journal-title":"eLife"},{"key":"2026092403080094300_RSTB20250135C28","doi-asserted-by":"crossref","first-page":"e59502","DOI":"10.7554\/eLife.59502","article-title":"The neural basis for a persistent internal state in Drosophila females","volume":"9","author":"Deutsch","year":"2020","journal-title":"eLife"},{"key":"2026092403080094300_RSTB20250135C29","doi-asserted-by":"crossref","first-page":"5998","DOI":"10.1016\/j.cell.2024.08.015","article-title":"A line attractor encoding a persistent internal state requires neuropeptide signaling","volume":"187","author":"Mountoufaris","year":"2024","journal-title":"Cell"},{"key":"2026092403080094300_RSTB20250135C30","volume-title":"Facial action coding system: a technique for the measurement of facial movement.","author":"Ekman","year":"1978"},{"key":"2026092403080094300_RSTB20250135C31","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1177\/1529100619832930","article-title":"Emotional expressions reconsidered: challenges to inferring emotions from human facial movements","volume":"20","author":"Barrett","year":"2019","journal-title":"Psychol. Sci. Public Interest"},{"key":"2026092403080094300_RSTB20250135C32","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.neubiorev.2020.02.031","article-title":"Measuring the evolution of facial \u2018expression\u2019 using multi-species FACS","volume":"113","author":"Waller","year":"2020","journal-title":"Neurosci. Biobehav. Rev."},{"key":"2026092403080094300_RSTB20250135C33","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1126\/science.aaz9468","article-title":"Facial expressions of emotion states and their neuronal correlates in mice","volume":"368","author":"Dolensek","year":"2020","journal-title":"Science"},{"key":"2026092403080094300_RSTB20250135C34","doi-asserted-by":"crossref","first-page":"43","DOI":"10.3389\/fnins.2014.00043","article-title":"The amygdala-motor pathways and the control of facial expressions","volume":"8","author":"Gothard","year":"2014","journal-title":"Front. Neurosci."},{"key":"2026092403080094300_RSTB20250135C35","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.jhevol.2004.10.003","article-title":"Evolution of the brainstem orofacial motor system in primates: a comparative study of trigeminal, facial, and hypoglossal nuclei","volume":"48","author":"Sherwood","year":"2005","journal-title":"J. Hum. Evol."},{"key":"2026092403080094300_RSTB20250135C36","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.neubiorev.2013.11.002","article-title":"The facial motor system","volume":"38","author":"Cattaneo","year":"2014","journal-title":"Neurosci. Biobehav. Rev."},{"key":"2026092403080094300_RSTB20250135C37","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1002\/cne.21165","article-title":"Amygdala interconnections with the cingulate motor cortex in the rhesus monkey","volume":"500","author":"Morecraft","year":"2007","journal-title":"J. Comp. Neurol."},{"key":"2026092403080094300_RSTB20250135C38","doi-asserted-by":"crossref","first-page":"3035","DOI":"10.1093\/brain\/awy219","article-title":"Motor and emotional behaviours elicited by electrical stimulation of the human cingulate cortex","volume":"141","author":"Caruana","year":"2018","journal-title":"Brain"},{"key":"2026092403080094300_RSTB20250135C39","doi-asserted-by":"crossref","first-page":"1115","DOI":"10.1126\/science.7652558","article-title":"Double dissociation of conditioning and declarative knowledge relative to the amygdala and hippocampus in humans","volume":"269","author":"Bechara","year":"1995","journal-title":"Science"},{"key":"2026092403080094300_RSTB20250135C40","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.cub.2010.11.042","article-title":"The human amygdala and the induction and experience of fear","volume":"21","author":"Feinstein","year":"2011","journal-title":"Curr. Biol."},{"key":"2026092403080094300_RSTB20250135C41","first-page":"1","volume-title":"Living without an amygdala","author":"Feinstein","year":"2016"},{"key":"2026092403080094300_RSTB20250135C42","doi-asserted-by":"crossref","first-page":"imag_a_00517","DOI":"10.1162\/imag_a_00517","article-title":"Cerebral topographies of perceived and felt emotion","volume":"3","author":"Saarim\u00e4ki","year":"2025","journal-title":"Imaging Neurosci."},{"key":"2026092403080094300_RSTB20250135C43","volume-title":"How emotions are made: the secret life of the brain","author":"Barrett","year":"2017"},{"key":"2026092403080094300_RSTB20250135C44","doi-asserted-by":"crossref","first-page":"3887","DOI":"10.1016\/j.neuron.2025.11.003","article-title":"The neural computation of affective internal states in the hypothalamus: a dynamical systems perspective","volume":"113","author":"Nair","year":"2025","journal-title":"Neuron"},{"key":"2026092403080094300_RSTB20250135C45","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1037\/h0024127","article-title":"Motivational and emotional controls of cognition","volume":"74","author":"Simon","year":"1967","journal-title":"Psychol. Rev."},{"key":"2026092403080094300_RSTB20250135C46","first-page":"185","volume-title":"Evolution and learning","author":"Fanselow","year":"1988"},{"key":"2026092403080094300_RSTB20250135C47","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1038\/nrn3945","article-title":"Neuronal circuits for fear and anxiety","volume":"16","author":"Tovote","year":"2015","journal-title":"Nat. Rev. Neurosci."},{"key":"2026092403080094300_RSTB20250135C48","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1146\/annurev-psych-122216-011854","article-title":"The emotion process: event appraisal and component differentiation","volume":"70","author":"Scherer","year":"2019","journal-title":"Annu. Rev. Psychol."},{"key":"2026092403080094300_RSTB20250135C49","author":"Fodor","year":"1983"},{"key":"2026092403080094300_RSTB20250135C50","doi-asserted-by":"crossref","first-page":"974","DOI":"10.1038\/s41593-019-0392-5","article-title":"Evidence that recurrent circuits are critical to the ventral stream's execution of core object recognition behavior","volume":"22","author":"Kar","year":"2019","journal-title":"Nat. Neurosci."},{"key":"2026092403080094300_RSTB20250135C51","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1016\/j.cell.2022.11.027","article-title":"An approximate line attractor in the hypothalamus encodes an aggressive state","volume":"186","author":"Nair","year":"2023","journal-title":"Cell"},{"key":"2026092403080094300_RSTB20250135C52","doi-asserted-by":"crossref","first-page":"110378","DOI":"10.1016\/j.isci.2024.110378","article-title":"A shared structure for emotion experiences from narratives, videos, and everyday life","volume":"27","author":"Han","year":"2024","journal-title":"iScience"},{"key":"2026092403080094300_RSTB20250135C53","doi-asserted-by":"crossref","first-page":"726","DOI":"10.1007\/s42761-025-00337-6","article-title":"Investigating the structure of emotion: tools, pitfalls, and recommendations","volume":"6","author":"Lin","year":"2025","journal-title":"Affect. Sci."},{"key":"2026092403080094300_RSTB20250135C54","article-title":"Unsupervised mapping of causal relations between brain lesions and behavior","author":"Wahle","year":"2023"},{"key":"2026092403080094300_RSTB20250135C55","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1038\/s41586-019-1858-z","article-title":"Internal state dynamics shape brainwide activity and foraging behaviour","volume":"577","author":"Marques","year":"2020","journal-title":"Nature"},{"key":"2026092403080094300_RSTB20250135C56","volume-title":"The ecological approach to visual perception","author":"Gibson","year":"1979"}],"container-title":["Philosophical Transactions of the Royal Society B: Biological Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article-pdf\/doi\/10.1098\/rstb.2025.0135\/6167197\/rstb.2025.0135.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article-pdf\/doi\/10.1098\/rstb.2025.0135\/6167197\/rstb.2025.0135.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T07:08:17Z","timestamp":1790233697000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article\/381\/1959\/20250135\/483379\/Comparative-neuroanatomy-of-emotion-expression"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,24]]},"references-count":56,"journal-issue":{"issue":"1959","published-online":{"date-parts":[[2026,9,24]]},"published-print":{"date-parts":[[2026,9,24]]}},"URL":"https:\/\/doi.org\/10.1098\/rstb.2025.0135","relation":{},"ISSN":["0962-8436","1471-2970"],"issn-type":[{"value":"0962-8436","type":"print"},{"value":"1471-2970","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,24]]},"article-number":"20250135"}}