The neurobiology of social play behaviour: Past, present and future
Open accessAbstract
Social play behaviour is a highly energetic and rewarding activity that is of great importance for the development of brain and behaviour. Social play is abundant during the juvenile and early adolescent phases of life, and it occurs in most mammalian species, as well as in certain birds and reptiles. To date, the majority of research into the neural mechanisms of social play behaviour has been performed in male rats. In the present review we summarize studies on the neurobiology of social play behaviour in rats, including work on pharmacological and genetic models for autism spectrum disorders, early life manipulations and environmental factors that influence play in rats. We describe several recent developments that expand the field, and highlight outstanding questions that may guide future studies.
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Treatment with psychostimulant drugs in doses that evoke hyperactivity typically results in a reduction in social play.
“For example, treatment with psychostimulant drugs in doses that evoke hyperactivity, typically results in a reduction in social play (for reviews see Vanderschuren et al. 2016; Achterberg and Vanderschuren 2023)”
From Neurexin1α knockout in rats causes aberrant social behaviour: relevance for autism and schizophrenia (2025) · cites this paper as Achterberg and Vanderschuren 2023
Locomotor hyperactivity and social play behaviour can be pharmacologically dissociated.
“Furthermore, locomotor hyperactivity and social play behaviour can be pharmacologically dissociated.”
From Neurexin1α knockout in rats causes aberrant social behaviour: relevance for autism and schizophrenia (2025) · cites this paper as Achterberg and Vanderschuren 2023
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Animal play and evolution: Seven timely research issues about enigmatic phenomena
Burghardt, G. M., Pellis, S. M., Schank, J. C., Smaldino, P. E., Vanderschuren, L. J. M. J., & Palagi, E. (2024) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2024.105617 · Cited by 32 · see citing works
Self-citationPellis, S. M., Pellis, V. C., & Ham, J. R. (2024) · Frontiers in Ethology · doi:10.3389/fetho.2024.1362052 · Cited by 15 · see citing works
Playful brains: a possible neurobiological pathway to cognitive health in aging
Golland, Y., Ben‐David, B. M., Mather, M., & Keisari, S. (2025) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2025.1490864 · Cited by 16 · see citing works
Kaplan, G. (2024) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2024.105621 · Cited by 15 · see citing works
Steinerová, K., Parker, S., Brown, J., & Seddon, Y. M. (2024) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2024.106263 · Cited by 16 · see citing works
Play partner preferences among groups of unfamiliar juvenile male rats
Ham, J. R., & Pellis, S. M. (2024) · Scientific Reports · doi:10.1038/s41598-024-66988-w · Cited by 13 · see citing works
Social play fosters cooperation in wild adult chimpanzees
Samuni, L., Mielke, A., Crockford, C., & Wittig, R. M. (2024) · Current Biology · doi:10.1016/j.cub.2024.10.058 · Cited by 15 · see citing works
Neurexin1α knockout in rats causes aberrant social behaviour: relevance for autism and schizophrenia
Achterberg, E. J. M., Biemans, B., & Vanderschuren, L. J. M. J. (2025) · Psychopharmacology · doi:10.1007/s00213-024-06559-z · Cited by 9 · see citing works
“…psychostimulant drugs in doses that evoke hyperactivity, typically results in a reduction in social play (for reviews see Vanderschuren et al. 2016; Achterberg and Vanderschuren 2023), and we have previously shown that treatment with methylphenidate in a social setting does not alter locomotion,…”
Self-citationSocial play behavior is driven by glycine-dependent mechanisms
Dvorzhak, A., Brecht, M., & Schmitz, D. (2024) · Current Biology · doi:10.1016/j.cub.2024.06.073 · Cited by 9 · see citing works
Skateboarding and the surplus value of city play
Glenney, B., Bjorke, I., & Buchetti, A. (2024) · Frontiers in Sports and Active Living · doi:10.3389/fspor.2024.1454274 · Cited by 9 · see citing works
Nyberg, H., Bogen, I. L., Nygaard, E., Achterberg, E. J. M., & Andersen, J. M. (2024) · Psychopharmacology · doi:10.1007/s00213-024-06718-2 · Cited by 11 · see citing works
Self-citationSteinerová, K., Harding, J. C. S., Parker, S., Wilson, H. L., Finatto, A. N., & Seddon, Y. M. (2024) · Frontiers in Veterinary Science · doi:10.3389/fvets.2024.1460993 · Cited by 8 · see citing works
Nuancing ‘Emotional’ Social Play: Does Play Behaviour Always Underlie a Positive Emotional State?
Cordoni, G., & Norscia, I. (2024) · Animals · doi:10.3390/ani14192769 · Cited by 9 · see citing works
Oppositions, joints, and targets: the attractors that are the glue of social interactions
Ham, J. R., Pellis, S. M., & Pellis, V. C. (2024) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2024.1451283 · Cited by 5 · see citing works
Steinerová, K., Krause, A., Parker, S., & Seddon, Y. M. (2025) · Frontiers in Veterinary Science · doi:10.3389/fvets.2025.1518153 · Cited by 5 · see citing works
Ham, J. R., & Pellis, S. M. (2025) · iScience · doi:10.1016/j.isci.2025.112562 · Cited by 5 · see citing works
Juvenile male rats form preferences based on strain when playing in groups but not in pairs
Ham, J. R., Jaiswal, D., Waner-Mariquito, R., Pellis, S. M., & Achterberg, E. J. M. (2025) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2025.1617178 · Cited by 5 · see citing works
Self-citationPlay behavior in ectothermic vertebrates
Леонидович, Д. В. (2023) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2023.105428 · Cited by 5 · see citing works
Steinerová, K., Parker, S., & Seddon, Y. M. (2025) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2024.106502 · Cited by 4 · see citing works
Shad, K. B. (2025) · Journal of Sociocybernetics · doi:10.26754/ojs_jos/jos.2025111599 · Cited by 4 · see citing works
Adriaense, J. E. C., Ringen, E. J., Ohashi, A., & Burkart, J. M. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.08.14.607991 · Cited by 4 · see citing works
Wilson, S. C. (2024) · Animals · doi:10.3390/ani14142086 · Cited by 3 · see citing works
Polzin, B. J., Zhao, C., Stevenson, S. A., Gammie, S. C., & Riters, L. V. (2024) · Genes Brain & Behavior · doi:10.1111/gbb.12908 · Cited by 2 · see citing works
Ruiz‐Sobremazas, D., Coca, M. R., Morales‐Navas, M., Rodulfo-Cárdenas, R., López-Granero, C., Colomina, M. T., Pérez-Fernández, C., & Sánchez-Santed, F. (2025) · NeuroToxicology · doi:10.1016/j.neuro.2025.04.007 · Cited by 3 · see citing works
Achterberg, E. J. M., Klooster, J. G. L. T., Pimentel, T. C., Aleksić, A., Hendriks, J., Nijhof, S. L., van der Ent, C. K., & Lesscher, H. M. B. (2024) · Frontiers in Ethology · doi:10.3389/fetho.2024.1458136 · Cited by 2 · see citing works
Self-citationWallis, A. S. J., Mendl, M., Lecorps, B., & Held, S. D. E. (2025) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2025.1962 · Cited by 3 · see citing works
Alhama-Riba, J., de Theije, C. G. M., Salvi, V., Viveen, D., Corstjens, S. W. T., Achterberg, E. J. M., van de Looij, Y., Hoebeek, F. E., & Nijboer, C. H. A. (2026) · The Journal of Physiology · doi:10.1113/jp290257 · Cited by 2 · see citing works
Self-citationAdding a layer of playfulness?
Siviy, S. M. (2023) · Neuropsychoanalysis · doi:10.1080/15294145.2023.2263002 · Cited by 2 · see citing works
Byun, Y., & Noh, J. (2023) · Physiology & Behavior · doi:10.1016/j.physbeh.2023.114379 · Cited by 2 · see citing works
Disruption of social play influences development of caution in juvenile ground squirrels
Ryan, W. J., Kuan, M. B., & Nunes, S. (2024) · Frontiers in Ethology · doi:10.3389/fetho.2024.1410334 · Cited by 2 · see citing works
Play behavior: Why do adults play less than juveniles?
Pellis, S. M., & Ham, J. R. (2024) · Current Biology · doi:10.1016/j.cub.2024.06.054 · Cited by 2 · see citing works
Sex and Age-at-Injury as Determinants of Social Behavior Outcomes After TBI
Semple, B. D., & Mychasiuk, R. (2024) · Advances in neurobiology · doi:10.1007/978-3-031-69832-3_10 · Cited by 2 · see citing works
Partner Choice and Context‐Dependent Sex Differences in Rat Rough‐and‐Tumble Play
Ham, J. R., Pellis, S. M., & McDonald, R. J. (2026) · European Journal of Neuroscience · doi:10.1111/ejn.70426 · Cited by 1 · see citing works
“Rats less motivated to play are more inclined to evade an attack than roll over to supine (Pellis et al. 1997; Varlinskaya et al. 1999) as rolling to supine is thought to be an active solicitation of the partner, so facilitating the continuation of play (Achterberg and Vanderschuren 2023).”
Ham, J. R., Iwaniuk, A. N., & Pellis, S. M. (2026) · Behavioural Brain Research · doi:10.1016/j.bbr.2026.116198 · Cited by 1 · see citing works
Piechal, A., Blecharz‐Klin, K., Jakimiuk, A. A., Pyrzanowska, J., Joniec‐Maciejak, I., Mirowska-Guzel, D. M., & Widy‐Tyszkiewicz, E. (2024) · Neuroscience · doi:10.1016/j.neuroscience.2024.12.017 · Cited by 1 · see citing works
Understanding play from neurobiological, evolutionary, and comparative perspectives
Schank, J. C., Burghardt, G. M., Palagi, E., & Pellis, S. M. (2025) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2025.106100 · Cited by 1 · see citing works
Endocannabinoids and social behavior
Trezza, V., Rava, A., & Feo, A. (2025) · Handbook of behavioral neuroscience · doi:10.1016/b978-0-443-13908-6.00019-0 · Cited by 1 · see citing works
Puccio, G., Franceschini, S., Carbone, S., Filotto, C., Gori, S., Pievani, T., Trezza, V., Mascheretti, S., Facoetti, A., & Bertoni, S. (2026) · Computers in Human Behavior Reports · doi:10.1016/j.chbr.2026.101183 · Cited by 2 · see citing works
The role of the dopamine D1 receptor in anticipatory pleasure and social play
Witt, K. M., Harper, D. N., & Ellenbroek, B. A. (2024) · Neuropharmacology · doi:10.1016/j.neuropharm.2024.110225 · Cited by 1 · see citing works
A rat model of volitional mutual social interactions
Lis, C. A., Casile, A., Feulner, B., Garcia, J., Madangopal, R., Papastrat, K. M., Huang, Z., Pacheco-Spiewak, A., Ramsey, L. A., & Vènniro, M. (2025) · Neuropsychopharmacology · doi:10.1038/s41386-025-02113-3 · Cited by 1 · see citing works
Rat Anterior Insula Symmetrically Represents Tickling-Induced Playful EmotionsPreprint
Dagher, S., & Ishiyama, S. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.04.02.587725 · Cited by 1 · see citing works
Tivey, E. K. L., Martin, J., Brown, S. M., Bishop, V. R., Bombail, V., Lawrence, A. B., & Meddle, S. (2024) · Research Square · doi:10.21203/rs.3.rs-4472769/v1 · Cited by 1 · see citing works
Nuancing 'Emotional' Social Play: Does Play Behaviour Always Underlie a Positive Emotion?Preprint
Cordoni, G., & Norscia, I. (2024) · Preprints.org · doi:10.20944/preprints202407.1348.v1 · Cited by 1 · see citing works
Evaluating the play fighting of rats: A sociological perspective
Ham, J. R., Pellis, S. M., Pellis, V. C., & Achterberg, E. J. M. (2026) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2026.110836
Self-citationŠpinka, M. (2025) · Elsevier eBooks · doi:10.1016/b978-0-443-29068-8.00135-5
Ferland, J. N. (2025) · Springer eBooks · doi:10.1007/978-3-031-87990-6_8
The role of endocannabinoids in nonpharmacological addiction
Mostallino, R., & Fattore, L. (2025) · Handbook of behavioral neuroscience · doi:10.1016/b978-0-443-13908-6.00025-6
Roles of eCB in adolescence and sensitive periods
Zamberletti, E., Gabaglio, M., & Rubino, T. (2025) · Handbook of behavioral neuroscience · doi:10.1016/b978-0-443-13908-6.00017-7
Chen, X., Tao, X., Zhong, Z., Zhang, Y., Li, Y., Ouyang, Y., Ding, Z., An, M., Wang, M., & Li, Y. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.11.12.687985
Heightened Play following Play Observation in the Absence of 22KHz Calls in Juvenile RatsPreprint
Michon, F., Achterberg, E. J. M., Kaufmann, L. V., Packheiser, J., Bos, S., van Reenen, K., Gazzola, V., & Keysers, C. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2025.12.31.697143
Self-citationMethylation and merriment: an exploration of epigenetic mechanisms underlying play behaviour
Malka, R. J., Arranz, M., & Baselga-Carretero, I. (2026) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1002/brv.70142
Watanasriyakul, W. T. (2026) · Developmental Psychobiology · doi:10.1002/dev.70158
The spectre and promise of species diversity
Pellis, S. M. (2026) · International Journal of Play · doi:10.1080/21594937.2026.2650984
Heightened play following play observation in the absence of 22-kHz calls in juvenile rats
Michon, F., Achterberg, E. J. M., Kaufmann, L. V., Packheiser, J., Bos, S., van Reenen, K., Gazzola, V., & Keysers, C. (2026) · Behavioural Brain Research · doi:10.1016/j.bbr.2026.116382 · Cited by 1 · see citing works
Self-citationHeld, S. D. E. (2026) · Wiley eBooks · doi:10.1002/9781394182237.ch8
Primate Play and the Classificatory Making of Human Nature
Bădescu, I. (2026) · Evolutionary Anthropology Issues News and Reviews · doi:10.1002/evan.70047
- Authors
- Achterberg, E. J. M., & Vanderschuren, L. J. M. J.
- Affiliations
- Utrecht University (NL)
- Corresponding author
- Achterberg, E.J. Marijke
- Vanderschuren, Louk J.M.J.
- Venue
- Neuroscience & Biobehavioral Reviews
- Publisher
- Elsevier BV
- Published
- 2023-09
- Volume
- 152
- Pages
- 105319
- ISSN
- 0149-7634 (ISSN-L)
- OpenAlex
- W4384283590
- Cited by
- 56
- References
- 239
- Citations by year
- 2026: 11 · 2025: 17 · 2024: 25 · 2023: 3
- FWCI
- 2.73
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Neuroscience & Biobehavioral Reviews (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Utrecht University Repository (Utrecht University) (submittedVersion, other-oa)
- Funders
- Nederlandse Organisatie voor Wetenschappelijk Onderzoek
- Topics
- Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Cognitive Neuroscience › Autism Spectrum Disorder Research
- Keywords
- rats, pharmacological models, genetic models, early life experience, environmental enrichment, juvenile development, brain development, social behavior
- MeSH terms
- Animals, Brain, Male, Mammals, Social Behavior, Rats
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