{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,20]],"date-time":"2026-06-20T17:41:41Z","timestamp":1781977301284,"version":"3.54.5"},"reference-count":139,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2024,8,27]],"date-time":"2024-08-27T00:00:00Z","timestamp":1724716800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Behav. Neurosci."],"abstract":"<jats:p>Social interactions are often analyzed by scoring segments of predefined behavior and then statistically assessing numerical and sequential patterns to identify the structure of the encounters. However, this approach can miss the dynamics of the animals\u2019 relationship over the course of the encounter, one that often involves invariant bonds, say a nose-to-nose orientation, with many different movements performed by both partners acting to counteract each other\u2019s attempts to break or maintain the relationship. Moreover, these invariant bonds can switch from one configuration to another during an interaction, leading from one stable configuration to another. It is this stepwise sequence of configurational stabilities that lead to functional outcomes, such as mating, aggression, or predation. By focusing on the sequence of invariant relational configurations, the deep structure of interactions can be discerned. This deep structure can then be used to differentiate between compensatory movements, no matter how seemingly stereotyped they may appear, from movement patterns which are restricted to a particular form when more than one option is available. A dynamic perspective requires suitable tools for analysis, and such tools are highlighted as needed in describing particular interactions.<\/jats:p>","DOI":"10.3389\/fnbeh.2024.1451283","type":"journal-article","created":{"date-parts":[[2024,8,27]],"date-time":"2024-08-27T13:05:18Z","timestamp":1724763918000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":6,"title":["Oppositions, joints, and targets: the attractors that are the glue of social interactions"],"prefix":"10.3389","volume":"18","author":[{"given":"Jackson R.","family":"Ham","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sergio M.","family":"Pellis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vivien C.","family":"Pellis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2024,8,27]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"105319","DOI":"10.1016\/j.neubiorev.2023.105319","article-title":"The neurobiology of social play behavior: past, present and future","volume":"152","author":"Achterberg","year":"2023","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref2","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1037\/a0016437","article-title":"Effects of neonatal handling on playfulness by means of reversal of the desire to play in rats (Rattus norvegicus)","volume":"123","author":"Aguilar","year":"2009","journal-title":"J. Comp. Psychol."},{"key":"ref3","volume-title":"Play fighting","author":"Aldis","year":"1975"},{"key":"ref4","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1007\/s10329-024-01123-x","article-title":"Automatic identification of stone-handling behaviour in Japanese macaques using LabGym artificial intelligence","volume":"65","author":"Ardoin","year":"2024","journal-title":"Primates"},{"key":"ref5","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1002\/dev.10069","article-title":"Effects of neonatal handling and maternal separation on rough-and-tumble play in the rat","volume":"41","author":"Arnold","year":"2002","journal-title":"Dev. Psychobiol."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1016\/0003-3472(67)90018-8","article-title":"Comparison of behavioural development in socially and isolated grouped rats","volume":"15","author":"Baenninger","year":"1967","journal-title":"Anim. Behav."},{"key":"ref7","doi-asserted-by":"publisher","first-page":"105868","DOI":"10.1016\/j.applanim.2023.105868","article-title":"The interconnection of hierarchy, affiliative behaviours, and social play shapes social dynamics in Maremmana beef cattle","volume":"260","author":"Bagnato","year":"2023","journal-title":"App. Anim. Behav. Sci."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"177","DOI":"10.2307\/1381018","article-title":"Fighting behavior of wild Sus scrofa","volume":"67","author":"Barrette","year":"1986","journal-title":"J. Mammal."},{"key":"ref9","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1155\/1968\/96242","article-title":"The mating behavior of Gromphadorhina portentosa (Schaum) (Blattaria, Blaberoidea, Oxyhaloinae): an anomalous pattern for a cockroach","volume":"75","author":"Barth","year":"1968","journal-title":"Psyche"},{"key":"ref10","volume-title":"Control in living systems: an exploration of the cybernetic properties of interactive behaviour","author":"Bell","year":"2013"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"196","DOI":"10.46867\/ijcp.2014.27.02.02","article-title":"Behavioral variability in the service of constancy","volume":"27","author":"Bell","year":"2014","journal-title":"In. J. Comp. Psychol."},{"key":"ref12","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1016\/j.anbehav.2011.06.016","article-title":"A cybernetic perspective on food protection in rats: simple rules can generate complex and adaptable behaviour","volume":"82","author":"Bell","year":"2011","journal-title":"Anim. Behav."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1163\/156853991X00247","article-title":"Feeding habits and predatory behaviour of marbled polecats (Vormella peregusna syriaca): I. Killing methods with relation to prey size and prey behaviour","volume":"118","author":"Ben-David","year":"1991","journal-title":"Behaviour"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1037\/11525-018","article-title":"The colony model of aggression and defense","volume-title":"Contemporary issues in comparative psychology","author":"Blanchard","year":"1990"},{"key":"ref15","first-page":"133","article-title":"Environmental targets and sensorimotor systems in aggression and defence","volume-title":"Ethology and psychopharmacology","author":"Blanchard","year":"1994"},{"key":"ref16","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1007\/BF03395855","article-title":"Conspecific wounding in free ranging Rattus norvegicus","volume":"35","author":"Blanchard","year":"1985","journal-title":"Psychol. Rec."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1016\/0003-3472(77)90113-0","article-title":"Attack and defensive behaviour in the albino rat","volume":"25","author":"Blanchard","year":"1977","journal-title":"Anim. Behav."},{"key":"ref18","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1002\/1098-2337(1978)4:1<31::AID-AB2480040104>3.0.CO;2-Y","article-title":"Aggressive behaviors of the Japanese brown bear","volume":"4","author":"Blanchard","year":"1979","journal-title":"Aggress. Behav."},{"key":"ref19","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1016\/0003-3472(64)90062-4","article-title":"The ontogeny of behavior in the albino rat","volume":"12","author":"Bolles","year":"1964","journal-title":"Anim. Behav."},{"key":"ref20","first-page":"197","article-title":"Agonistic interactions of two cockroach species, Gromphadorhina portentosa and Supella longipalpa (orthoptera (dictyoptera): Blaberidae, blattellidae)","volume":"54","author":"Breed","year":"1981","journal-title":"J. Kans. Entomol. Soc."},{"key":"ref21","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1016\/j.jneumeth.2018.06.013","article-title":"T-pattern detection and analysis for the discovery of hidden features of behaviour","volume":"310","author":"Casarrubea","year":"2018","journal-title":"J. Neurosci. Methods"},{"key":"ref22","doi-asserted-by":"publisher","first-page":"112938","DOI":"10.1016\/j.physbeh.2020.112938","article-title":"Inferring tool use behaviour patterns from the temporal structure of object play sequences in a non-human primate species","volume":"222","author":"Cenni","year":"2020","journal-title":"Physiol. Behav."},{"key":"ref23","doi-asserted-by":"publisher","first-page":"1111908","DOI":"10.3389\/fnbeh.2023.1111908","article-title":"Alpha tracker: a multi-animal tracking and behavioral analysis tool","volume":"17","author":"Chen","year":"2023","journal-title":"Front. Behav. Neurosci."},{"key":"ref24","doi-asserted-by":"publisher","first-page":"1313","DOI":"10.1016\/j.anbehav.2012.09.002","article-title":"Social tolerance and adult play in macaque societies: a comparison with different human cultures","volume":"84","author":"Ciani","year":"2012","journal-title":"Anim. Behav."},{"key":"ref25","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1007\/BF01990081","article-title":"Social interactions and aggression among male Madagascar hissing cockroaches (Gromphadorhina portentosa) in groups (Dictyoptera: Blaberidae)","volume":"7","author":"Clark","year":"1994","journal-title":"J. Insect Behav."},{"key":"ref26","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/0003-3472(79)90141-6","article-title":"The logical stag: adaptive aspects of fighting in red deer (Cervus elaphus L.)","volume":"27","author":"Clutton-Brock","year":"1979","journal-title":"Anim. Behav."},{"key":"ref27","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1016\/S0003-3472(73)80007-7","article-title":"Agonistic behaviour in males of five species of voles","volume":"21","author":"Colvin","year":"1973","journal-title":"Anim. Behav."},{"key":"ref28","doi-asserted-by":"publisher","first-page":"104778","DOI":"10.1016\/j.beproc.2022.104778","article-title":"Play fighting versus real fighting in piglets: similar patterns, different structure","volume":"203","author":"Cordoni","year":"2022","journal-title":"Behav. Process."},{"key":"ref29","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1016\/j.anbehav.2021.02.016","article-title":"The dark side of play: play fighting as a substitute for real fighting in domestic pigs, Sus scrofa","volume":"175","author":"Cordoni","year":"2021","journal-title":"Anim. Behav."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"278","DOI":"10.1037\/com0000035","article-title":"Unveiling the \u201csecret\u201d of play in dogs (Canis lupus familiaris): asymmetry and signals","volume":"130","author":"Cordoni","year":"2016","journal-title":"J. Comp. Psychol."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"e0193096","DOI":"10.1371\/journal.pone.0193096","article-title":"Differences in play can illuminate differences in affiliation: a comparative study on chimpanzees and gorillas","volume":"13","author":"Cordoni","year":"2018","journal-title":"PLoS One"},{"key":"ref32","doi-asserted-by":"publisher","first-page":"749","DOI":"10.1163\/1568539X-00003326","article-title":"Aggression and hierarchical steepness inhibit social play in adult wolves","volume":"153","author":"Cordoni","year":"2016","journal-title":"Behaviour"},{"key":"ref33","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1002\/zoo.21824","article-title":"Agonistic interactions and social behaviors in the Saharan Dorcas gazelle (Gazella Dorcas neglecta): using social network analysis to evaluate relationships and social structure in captive male groups","volume":"43","author":"Cort\u00e9s","year":"2024","journal-title":"Zoo Biol."},{"key":"ref34","first-page":"500","article-title":"Wild pigs and boars","volume-title":"The encyclopaedia of mammals","author":"Cumming","year":"1984"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1007\/978-3-031-35651-3_8","article-title":"Play, sexual display, or just boredom relief?","volume-title":"Sex in cetaceans","author":"Da Silva","year":"2023"},{"key":"ref36","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1111\/j.1439-0310.1989.tb00518.x","article-title":"A comparative study of agonistic behaviour in hairy-footed gerbils (genus Gerbillurus)","volume":"83","author":"Dempster","year":"1989","journal-title":"Ethology"},{"key":"ref37","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.lmot.2017.03.003","article-title":"Social order: using the sequential structure of social interaction to discriminate abnormal social behavior in the rat","volume":"61","author":"Donaldson","year":"2018","journal-title":"Learn. Motiv."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1111\/j.1439-0396.2012.01331.x","article-title":"Physiological and hormonal aspects of female domestic pigeons (Columba livia) associated with breeding and growth","volume":"97","author":"Dong","year":"2013","journal-title":"J. Anim. Physiol. Anim. Nutr."},{"key":"ref39","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1002\/dev.420210707","article-title":"The ontogeny of exploratory behavior in the house rat (Rattus rattus): the mobility gradient","volume":"21","author":"Eilam","year":"1989","journal-title":"Dev. Psychobiol."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"872634","DOI":"10.3389\/fnmol.2022.872634","article-title":"Chemogenetic modulation of sensory afferents induces locomotor changes and plasticity after spinal cord injury","volume":"15","author":"Eisdorfer","year":"2022","journal-title":"Front. Mol. Neurosci."},{"key":"ref41","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1016\/0003-3472(69)90139-0","article-title":"Hormonal control of courtship behaviour in the pigeon (Columbia livia)","volume":"17","author":"Erpino","year":"1969","journal-title":"Anim. Behav."},{"key":"ref42","volume-title":"Movement notation","author":"Eshkol","year":"1958"},{"key":"ref43","volume-title":"The safari companion. A guide to watching African mammals","author":"Estes","year":"1993"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"534","DOI":"10.1016\/0003-3472(63)90276-8","article-title":"Laboratory observations on the reproductive behaviour of the pigeon (Columbia livia) during the pre-incubation phase of the breeding cycle","volume":"11","author":"Fabricius","year":"1963","journal-title":"Anim. Behav."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1002\/1098-2337(1990)16:1<41::AID-AB2480160106>3.0.CO;2-V","article-title":"Ethological analysis of the male rat's socioagonistic behaviour in a resident-intruder paradigm","volume":"16","author":"Fern\u00e1ndez-Espejo","year":"1990","journal-title":"Aggress. Behav."},{"key":"ref46","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1037\/0735-7036.110.3.298","article-title":"An analysis of sex differences in the movement patterns used during the food wrenching and dodging paradigm","volume":"110","author":"Field","year":"1996","journal-title":"J. Comp. Psychol."},{"key":"ref47","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1002\/(SICI)1098-2337(1997)23:3<197::AID-AB5>3.0.CO;2-J","article-title":"The organization of sex-typical patterns of defense during food protection in the rat: the role of the opponent\u2019s sex","volume":"23","author":"Field","year":"1997","journal-title":"Aggress. Behav."},{"key":"ref48","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/B978-0-12-821410-7.00008-X","article-title":"Movement analysis: expanding the resolution of analysis in animal behavior","volume-title":"Exploring animal behavior in laboratory and field","author":"Foroud","year":"2021"},{"key":"ref49","first-page":"133","article-title":"A comparison of behaviour in the Suidae","volume-title":"The behaviour of ungulates and its relation to management","author":"Fr\u00e4drich","year":"1974"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"194","DOI":"10.1016\/S0003-3472(84)80337-1","article-title":"Communication in the courtship of a Madagascan hissing cockroach. I. Normal courtship","volume":"32","author":"Fraser","year":"1984","journal-title":"Anim. Behav."},{"key":"ref51","doi-asserted-by":"publisher","first-page":"104338","DOI":"10.1016\/j.beproc.2021.104338","article-title":"Let\u2019s unite in play! Play modality and group membership in wild geladas","volume":"184","author":"Gallo","year":"2021","journal-title":"Behav. Process."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"551","DOI":"10.2307\/1377327","article-title":"On the rutting behavior of the mountain goal","volume":"45","author":"Geist","year":"1965","journal-title":"J. Mammal."},{"key":"ref53","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1163\/156853966X00155","article-title":"The evolution of hornlike organs","volume":"27","author":"Geist","year":"1966","journal-title":"Behaviour"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"192","DOI":"10.2307\/3798378","article-title":"On fighting injuries and dermal shields of mountain goals","volume":"31","author":"Geist","year":"1967","journal-title":"J. Wildl. Manag."},{"key":"ref55","first-page":"1","article-title":"On weapons, combat and ecology","volume-title":"Advances in the study of communication and affect Vol. 4 aggression, dominance and individual spacing","author":"Geist","year":"1978"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/978-1-4615-7572-6_2","article-title":"Homeostatic motor processes in mammalian interactions: a choreography of display","volume-title":"Perspectives in ethology","author":"Golani","year":"1976"},{"key":"ref57","first-page":"372","article-title":"The search for invariants in behavior","volume-title":"Behavioral development: the Bielefeld interdisciplinary project","author":"Golani","year":"1981"},{"key":"ref58","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1002\/dev.420180609","article-title":"Early ontogeny of face grooming in mice","volume":"18","author":"Golani","year":"1985","journal-title":"Dev. Psychobiol."},{"key":"ref59","first-page":"65","article-title":"A motility-immobility gradient in the behaviour of the \u2018inferior\u2019 wolf during \u2018ritualized fighting\u2019","volume-title":"Advances in the study of mammalian behavior","author":"Golani","year":"1983"},{"key":"ref60","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1016\/0006-8993(79)90019-2","article-title":"A proposed natural geometry of recovery from akinesia in the lateral hypothalamic rat","volume":"164","author":"Golani","year":"1979","journal-title":"Brain Res."},{"key":"ref61","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1016\/S0065-3454(08)60409-X","article-title":"Development of social displays: form, development, form fixation and change in context","volume":"22","author":"Groothuis","year":"1993","journal-title":"Adv. Stud. Behav."},{"key":"ref62","doi-asserted-by":"publisher","first-page":"104695","DOI":"10.1016\/j.beproc.2022.104695","article-title":"The emergence and early development of socio-sexual behavior in beluga calves (Delphinapterus leucas)","volume":"200","author":"Ham","year":"2022","journal-title":"Behav. Process."},{"key":"ref63","first-page":"129","article-title":"Non-conceptive sexual behavior in cetaceans: comparison of form and function","volume-title":"Sex in cetaceans","author":"Ham","year":""},{"key":"ref64","doi-asserted-by":"publisher","first-page":"730","DOI":"10.1002\/zoo.21788","article-title":"Playful mouth-to-mouth interactions of belugas (Delphinapterus leucas) in managed care","volume":"42","author":"Ham","year":"","journal-title":"Zoo Biol."},{"key":"ref65","doi-asserted-by":"publisher","first-page":"e52407","DOI":"10.3791\/52407-v","article-title":"Moderate prenatal alcohol exposure and quantification of social behavior in adult rats","volume":"94","author":"Hamilton","year":"2014","journal-title":"JoVE"},{"key":"ref66","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1111\/j.1439-0310.1985.tb00123.x","article-title":"The form of combative strategy in interactions among wolf pups (Canis lupus)","volume":"68","author":"Havkin","year":"1985","journal-title":"Ethology"},{"key":"ref67","doi-asserted-by":"publisher","first-page":"105","DOI":"10.12966\/abc.05.01.2015","article-title":"Developing a catalog of socio-sexual behaviors of beluga whales (Delphinapterus leucas)","volume":"2","author":"Hill","year":"2015","journal-title":"Anim. Behav. Cogn."},{"key":"ref68","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1578\/AM.48.6.2022.541","article-title":"Observations of mating practice by non-sexually mature male belugas (Delphinapterus lecuas)","volume":"48","author":"Hill","year":"2022","journal-title":"Aquat. Mamm."},{"key":"ref69","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1037\/bne0000010","article-title":"The role of the medial prefrontal cortex in regulating inter-animal coordination of movements","volume":"128","author":"Himmler","year":"2014","journal-title":"Behav. Neurosci."},{"key":"ref70","doi-asserted-by":"publisher","first-page":"1","DOI":"10.46867\/ijcp.2015.28.00.10","article-title":"The development of juvenile-typical patterns of play fighting in juvenile rats does not depend on peer\u2013peer play experience in the peri-weaning period","volume":"28","author":"Himmler","year":"2015","journal-title":"Int. J. Comp. Psychol."},{"key":"ref71","doi-asserted-by":"publisher","first-page":"385","DOI":"10.46867\/ijcp.2014.27.03.09","article-title":"The development of strain typical defensive patterns in the play fighting of laboratory rats","volume":"27","author":"Himmler","year":"2014","journal-title":"Int. J. Comp. Psychol."},{"key":"ref72","doi-asserted-by":"publisher","first-page":"e54540","DOI":"10.7554\/eLife.54540","article-title":"A Matlab-based toolbox for characterizing behavior of rodents engaged in string pulling","volume":"9","author":"Inayat","year":"2020","journal-title":"eLife"},{"key":"ref73","doi-asserted-by":"publisher","first-page":"1005","DOI":"10.1006\/anbe.2003.2124","article-title":"Is the parallel walk between competing male fallow deer, Dama dama, a lateral display of individual quality?","volume":"65","author":"Jennings","year":"2003","journal-title":"Anim. Behav."},{"key":"ref74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.46867\/ijcp.2019.32.00.10","article-title":"Targets, tactics and cooperation in the play fighting of two genera of Old World Monkeys (Mandrillus and Papio): accounting for similarities and differences","volume":"32","author":"Kraus","year":"2019","journal-title":"Int. J. Comp. Psychol."},{"key":"ref75","first-page":"1","article-title":"Ontogeny of social behaviour in Burmese red junglefowl (Gallus gallus spadiceus) Bonnaterre","volume":"12","author":"Kruijt","year":"1964","journal-title":"Behaviour. Supp."},{"key":"ref76","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1038\/s41592-022-01443-0","article-title":"Multi-animal pose estimation, identification and tracking with DeepLabCut","volume":"19","author":"Lauer","year":"2022","journal-title":"Nat. Methods"},{"key":"ref77","doi-asserted-by":"publisher","first-page":"714","DOI":"10.1016\/S0003-3472(71)80175-6","article-title":"The agonistic of behaviour of the golden hamster Mesocricetus auratus (Waterhouse)","volume":"19","author":"Lerwill","year":"1971","journal-title":"Anim. Behav."},{"key":"ref78","doi-asserted-by":"publisher","first-page":"104025","DOI":"10.1016\/j.beproc.2019.104025","article-title":"The development of socio-sexual behavior in belugas (Delphinapterus leucas) under human care","volume":"171","author":"Lilley","year":"2020","journal-title":"Behav. Process."},{"key":"ref79","doi-asserted-by":"publisher","first-page":"104718","DOI":"10.1016\/j.beproc.2022.104718","article-title":"Investigation of lateralization of socio-sexual behavior in belugas (Delphinapterus leucas)","volume":"201","author":"Lilley","year":"2022","journal-title":"Behav. Process."},{"key":"ref80","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1111\/eth.12594","article-title":"Strong competition does not always predict play asymmetry: the case of south American sea lions (Otaria flavescens)","volume":"123","author":"Llamazares-Mart\u00edn","year":"2017","journal-title":"Ethology"},{"key":"ref81","first-page":"173","article-title":"Sexual behaviors of Odontocetes in managed care","volume-title":"Sex in cetaceans","author":"Manitzas Hill","year":""},{"key":"ref82","doi-asserted-by":"publisher","first-page":"100071","DOI":"10.1016\/j.therwi.2024.100071","article-title":"A review of beluga (Delphinapterus leucas) sexual behavior and reproductive physiology leading to conception","volume":"4","author":"Manitzas Hill","year":"2024","journal-title":"Theriogenol. Wild"},{"key":"ref83","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1080\/21594937.2022.2152536","article-title":"Social games that belugas (Delphinapterus leucas) play","volume":"12","author":"Manitzas Hill","year":"","journal-title":"Int. J. Play"},{"key":"ref84","doi-asserted-by":"publisher","first-page":"1281","DOI":"10.1038\/s41593-018-0209-y","article-title":"DeepLabCut: markerless pose estimation of user-defined body parts with deep learning","volume":"21","author":"Mathis","year":"2018","journal-title":"Nat. Neurosci."},{"key":"ref85","doi-asserted-by":"publisher","first-page":"1146","DOI":"10.1016\/S0003-3472(81)80067-X","article-title":"A description of relational patterns of movement during \u2018ritualized fighting\u2019 in wolves","volume":"29","author":"Moran","year":"1981","journal-title":"Anim. Behav."},{"key":"ref86","doi-asserted-by":"publisher","first-page":"286","DOI":"10.1016\/0003-3472(69)90014-1","article-title":"The endocrine basis of breeding behaviour in the feral pigeon (Columba livia): I. Effects of exogenous hormones on the pre-incubation behaviour of intact males","volume":"17","author":"Murton","year":"1969","journal-title":"Anim. Behav."},{"key":"ref87","doi-asserted-by":"publisher","first-page":"2152","DOI":"10.1038\/s41596-019-0176-0","article-title":"Using DeepLabCut for 3D markerless pose estimation across species and behaviors","volume":"14","author":"Nath","year":"2019","journal-title":"Nat. Protoc."},{"key":"ref88","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1007\/BF00292773","article-title":"Sound production in the cockroach, Gromphadorhina portentosa: evidence for communication by hissing","volume":"6","author":"Nelson","year":"1980","journal-title":"Behav. Ecol. Sociobiol."},{"key":"ref89","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1016\/0376-6357(88)90004-6","article-title":"Playful behaviour of piglets","volume":"17","author":"Newberry","year":"1988","journal-title":"Behav. Process."},{"key":"ref90","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1016\/j.anbehav.2021.07.012","article-title":"Play fighting in wild spotted hyenas: like a bridge over troubled water of a hierarchical society","volume":"180","author":"Nolfo","year":"2021","journal-title":"Anim. Behav."},{"key":"ref91","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1016\/j.beproc.2014.09.001","article-title":"Down but not out: supine postures as facilitators of play in domestic dogs","volume":"110","author":"Norman","year":"2015","journal-title":"Behav. Process."},{"key":"ref92","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.beproc.2015.07.014","article-title":"A structural comparison of female-male and female-female mounting in Japanese macaques (Macaca fuscata)","volume":"119","author":"Ottenheimer Carrier","year":"2015","journal-title":"Behav. Process."},{"key":"ref93","doi-asserted-by":"publisher","first-page":"e52767","DOI":"10.1371\/journal.pone.0052767","article-title":"The right time to happen: play developmental divergence in the two pan species","volume":"7","author":"Palagi","year":"2012","journal-title":"PLoS One"},{"key":"ref94","doi-asserted-by":"publisher","first-page":"1195","DOI":"10.1163\/1568539X-00003336","article-title":"Fair play and its connection with social tolerance, reciprocity and the ethology of peace","volume":"153","author":"Palagi","year":"2016","journal-title":"Behaviour"},{"key":"ref95","doi-asserted-by":"publisher","first-page":"767","DOI":"10.1002\/dev.20342","article-title":"The influence of natural variations in maternal care on play fighting in the rat","volume":"50","author":"Parent","year":"2008","journal-title":"Dev. Psychobiol."},{"key":"ref96","doi-asserted-by":"publisher","first-page":"61","DOI":"10.3727\/015613881791560685","article-title":"A description of social play by the Australian magpie Gymnorhina tibicen based on Eshkol\u2013Wachman Notation","volume":"3","author":"Pellis","year":"1981","journal-title":"Bird Behav."},{"key":"ref97","doi-asserted-by":"publisher","first-page":"30","DOI":"10.3727\/015613882791560620","article-title":"An analysis of courtship and mating in the cape barren goose Cereopsis novaehollandiae Latham based on Eshkol\u2013Wachmann movement notation","volume":"4","author":"Pellis","year":"1982","journal-title":"Bird Behav."},{"key":"ref98","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1002\/1098-2337(1988)14:2<85::AID-AB2480140203>3.0.CO;2-5","article-title":"Agonistic versus amicable targets of attack and defense: consequences for the origin, function and descriptive classification of play-fighting","volume":"14","author":"Pellis","year":"1988","journal-title":"Aggress. Behav."},{"key":"ref99","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1002\/(SICI)1098-2337(1997)23:2<107::AID-AB3>3.0.CO;2-K","article-title":"Targets and tactics: the analysis of moment-to-moment decision making in animal combat","volume":"23","author":"Pellis","year":"1997","journal-title":"Aggress. Behav."},{"key":"ref100","doi-asserted-by":"publisher","first-page":"255","DOI":"10.46867\/ijcp.2011.24.03.04","article-title":"Head and foot coordination in head scratching and food manipulation by purple swamp hens (Porphyrio porphyrio): rules for minimizing the computational costs of combining movements from multiple parts of the body","volume":"24","author":"Pellis","year":"2011","journal-title":"Int. J. Comp. Psychol."},{"key":"ref101","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/B978-0-12-818948-1.00005-8","article-title":"Unraveling the dynamics of dyadic interactions: perceptual control in animal contests","volume-title":"The interdisciplinary handbook of perceptual control theory: living control systems IV","author":"Pellis","year":"2020"},{"key":"ref102","doi-asserted-by":"publisher","first-page":"1567","DOI":"10.1163\/1568539X-00003110","article-title":"Drawn into the vortex: the facing-past encounter and combat in lekking male greater sage-grouse (Centrocercus urophasianus)","volume":"150","author":"Pellis","year":"2013","journal-title":"Behaviour"},{"key":"ref103","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1002\/(SICI)1098-2302(199909)35:2<156::AID-DEV8>3.0.CO;2-C","article-title":"The development of a sex-differentiated defensive motor-pattern in rats: a possible role for juvenile experience","volume":"35","author":"Pellis","year":"1999","journal-title":"Dev. Psychobiol."},{"key":"ref104","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1002\/dev.420160403","article-title":"Locomotor-rotational movements in the ontogeny and play of the laboratory rat Rattus norvegicus","volume":"16","author":"Pellis","year":"1983","journal-title":"Dev. Psychobiol."},{"key":"ref105","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1002\/1098-2337(1987)13:4<227::AID-AB2480130406>3.0.CO;2-C","article-title":"Play-fighting differs from serious fighting in both target of attack and tactics of fighting in the laboratory rat Rattus norvegicus","volume":"13","author":"Pellis","year":"1987","journal-title":"Aggress. Behav."},{"key":"ref106","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1002\/1098-2337(1989)15:3<217::AID-AB2480150304>3.0.CO;2-B","article-title":"Targets of attack and defense in the play fighting by the Djungarian hamster Phodopus campbelli: links to fighting and sex","volume":"15","author":"Pellis","year":"1989","journal-title":"Aggress. Behav."},{"key":"ref107","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1002\/1098-2337(1992)18:4<301::AID-AB2480180406>3.0.CO;2-1","article-title":"An analysis of the targets and tactics of conspecific attack and predatory attack in northern grasshopper mice Onychomys leucogaster","volume":"18","author":"Pellis","year":"1992","journal-title":"Aggress. Behav."},{"key":"ref108","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1002\/(SICI)1098-2337(1997)23:1<41::AID-AB5>3.0.CO;2-W","article-title":"Targets, tactics and the open mouth face during play fighting in three species of primates","volume":"23","author":"Pellis","year":"","journal-title":"Aggress. Behav."},{"key":"ref109","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1002\/(SICI)1098-2302(199711)31:3<193::AID-DEV4>3.0.CO;2-N","article-title":"The pre-juvenile onset of play fighting in rats (Rattus norvegicus)","volume":"31","author":"Pellis","year":"","journal-title":"Dev. Psychobiol."},{"key":"ref110","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.physbeh.2015.03.038","article-title":"Are agonistic behavior patterns signals or combat tactics \u2013 or does it matter? Targets as organizing principles of fighting","volume":"146","author":"Pellis","year":"2015","journal-title":"Physiol. Behav."},{"key":"ref111","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1163\/1568539X-00003346","article-title":"Play fighting in Visayan warty pigs (Sus cebifrons): insights on restraint and reciprocity in the maintenance of play","volume":"153","author":"Pellis","year":"2016","journal-title":"Behaviour"},{"key":"ref112","doi-asserted-by":"publisher","first-page":"355","DOI":"10.3758\/s13420-017-0264-3","article-title":"What is play fighting and what is it good for?","volume":"45","author":"Pellis","year":"2017","journal-title":"Learn. Behav."},{"key":"ref113","doi-asserted-by":"crossref","DOI":"10.1017\/9781108650151","volume-title":"Understanding animal behaviour: what to measure and why","author":"Pellis","year":"2021"},{"key":"ref114","doi-asserted-by":"publisher","first-page":"128","DOI":"10.12966\/abc.05.04.2014","article-title":"One good turn deserves another: combat versus other functions of acrobatic maneuvers in the play fighting of vervet monkeys (Chlorocebus aethiops)","volume":"2","author":"Pellis","year":"2014","journal-title":"Anim. Behav. Cogn."},{"key":"ref115","doi-asserted-by":"publisher","first-page":"e337","DOI":"10.1002\/cpz1.337","article-title":"Measuring play fighting in rats: a multilayered approach","volume":"2","author":"Pellis","year":"2022","journal-title":"Curr. Protoc."},{"key":"ref116","doi-asserted-by":"publisher","first-page":"1362052","DOI":"10.3389\/fetho.2024.1362052","article-title":"Play fighting revisited: its design features and how they shape our understanding of its mechanisms and functions","volume":"3","author":"Pellis","year":"2024","journal-title":"Front. Ethol."},{"key":"ref117","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1038\/s41592-022-01426-1","article-title":"SLEAP: a deep learning system for multi-animal pose tracking","volume":"19","author":"Pereira","year":"2022","journal-title":"Nat. Methods"},{"key":"ref118","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1002\/dev.20305","article-title":"Social play in crested and Japanese macaques: testing the covariation hypothesis","volume":"50","author":"Petit","year":"2008","journal-title":"Dev. Psychobiol."},{"key":"ref119","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1093\/sxmrev\/qead034","article-title":"From distal to proximal to interactive: behavioral and brain synchrony during attraction, courtship, and sexual interaction\u2014implications for clinical assessments of relationship style and quality","volume":"11","author":"Pfaus","year":"2023","journal-title":"Sex. Med. Rev."},{"key":"ref120","doi-asserted-by":"publisher","first-page":"105110","DOI":"10.1016\/j.neubiorev.2023.105110","article-title":"Non-angry aggressive arousal and angriffsberietschaft: a narrative review of the phenomenology and physiology of proactive\/offensive aggression motivation and escalation in people and other animals","volume":"147","author":"Potegal","year":"2023","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref121","volume-title":"Behavior: the control of perception","author":"Powers","year":"2005"},{"key":"ref122","doi-asserted-by":"publisher","first-page":"166","DOI":"10.46867\/IJCP.2010.23.02.05","article-title":"Targets and tactics of play fighting: competitive versus cooperative styles of play in Japanese and Tonkean macaques","volume":"23","author":"Reinhart","year":"2010","journal-title":"Int. J. Comp. Psychol."},{"key":"ref123","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1002\/1098-2337(1987)13:6<329::AID-AB2480130602>3.0.CO;2-3","article-title":"Offence and defence in fighting between young pigs (Sus scrofa)","volume":"13","author":"Rushen","year":"1987","journal-title":"Aggress. Behav."},{"key":"ref124","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2173\/tbna.425.p","article-title":"Sage grouse: Centrocerus urophasianus","volume":"425","author":"Schroeder","year":"1999","journal-title":"Bird. N. Am."},{"key":"ref125","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1002\/dev.20202","article-title":"Early weaning decreases play-fighting behavior during the postweaning developmental period of Wistar rats","volume":"49","author":"Shimozuru","year":"2007","journal-title":"Dev. Psychobiol."},{"key":"ref126","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.applanim.2010.02.003","article-title":"Playing and fighting by piglets around weaning on farms, employing individual or group housing of lactating sows","volume":"124","author":"\u0160ilerov\u00e1","year":"2010","journal-title":"App. Anim. Behav. Sci."},{"key":"ref127","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/j.physbeh.2016.11.035","article-title":"Effects of cross-fostering on play and anxiety in juvenile Fischer 344 and Lewis rats","volume":"169","author":"Siviy","year":"2017","journal-title":"Physiol. Behav."},{"key":"ref128","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1037\/0735-7036.122.1.1","article-title":"Effects of neonatal handling on play behavior and fear towards predator odor in juvenile rats (Rattus norvegicus)","volume":"122","author":"Siviy","year":"2008","journal-title":"J. Comp. Psychol."},{"key":"ref129","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1111\/j.1467-9280.1992.tb00249.x","article-title":"Towards a synthetic physiological psychology","volume":"3","author":"Teitelbaum","year":"1992","journal-title":"Psychol. Sci."},{"key":"ref130","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1002\/dev.420230605","article-title":"Weaning in rats: II. Pup behavior patterns","volume":"23","author":"Thiels","year":"1990","journal-title":"Dev. Psychobiol."},{"key":"ref131","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1016\/j.physbeh.2011.12.007","article-title":"Within-litter variation in maternal care received by individual pups correlates with adolescent social play behavior in male rats","volume":"106","author":"Van Hasselt","year":"2012","journal-title":"Physiol. Behav."},{"key":"ref132","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1016\/j.psyneuen.2008.10.017","article-title":"Maternal separation enhances offensive play-fighting, basal corticosterone and hypothalamic vasopressin mRNA expression in juvenile male rats","volume":"34","author":"Veenema","year":"2008","journal-title":"Psychoneuroendocrinology"},{"key":"ref133","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1016\/0165-0270(88)90061-1","article-title":"Food wrenching and dodging: use of action patterns for the analysis of sensorimotor and social behavior in the rat","volume":"24","author":"Whishaw","year":"1988","journal-title":"J. Neurosci. Methods"},{"key":"ref134","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/0166-4328(90)90053-H","article-title":"The structure of skilled forelimb reaching in the rat: a proximally driven stereotyped movement with a single rotatory component","volume":"41","author":"Whishaw","year":"1990","journal-title":"Behav. Brain Res."},{"key":"ref135","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1016\/S0166-4328(05)80042-7","article-title":"The impairments in reaching and the movements of compensation in rats with motor cortex lesions: a videorecording and movement notation analysis","volume":"42","author":"Whishaw","year":"1991","journal-title":"Behav. Brain Res."},{"key":"ref136","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1163\/156853973X00319","article-title":"The strut display of male sage grouse \u2013 a \u2018fixed\u2019 action pattern","volume":"47","author":"Wiley","year":"","journal-title":"Behaviour"},{"key":"ref137","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/0003-3472(73)90004-3","article-title":"Territoriality and non-random mating in sage grouse, Centrocerus urophasianus","volume":"6","author":"Wiley","year":"","journal-title":"Anim. Behav. Monogr."},{"key":"ref138","doi-asserted-by":"publisher","first-page":"1560","DOI":"10.1111\/1365-2656.13932","article-title":"DeepWild: application of the pose estimation tool DeepLabCut for behaviour tracking in wild chimpanzees and bonobos","volume":"92","author":"Wiltshire","year":"2023","journal-title":"J. Anim. Ecol."},{"key":"ref139","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1111\/j.1439-0310.1987.tb00924.x","article-title":"Superiority and inferiority: a morphological analysis of free and stimulus bound behaviour in honey badger (Mellivora capensis) interactions","volume":"74","author":"Yaniv","year":"1987","journal-title":"Ethology"}],"container-title":["Frontiers in Behavioral Neuroscience"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnbeh.2024.1451283\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,8,27]],"date-time":"2024-08-27T13:05:35Z","timestamp":1724763935000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnbeh.2024.1451283\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,8,27]]},"references-count":139,"alternative-id":["10.3389\/fnbeh.2024.1451283"],"URL":"https:\/\/doi.org\/10.3389\/fnbeh.2024.1451283","relation":{},"ISSN":["1662-5153"],"issn-type":[{"value":"1662-5153","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,8,27]]},"article-number":"1451283"}}