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A brain motivated to play: insights into the neurobiology of playfulness
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Play is an important part of normal childhood development and is seen in varied forms among many mammals. While not indispensable to normal development, playful social experiences as juveniles may provide an opportunity to develop flexible behavioural strategies when novel and uncertain situations arise as an adult. To understand the neurobiological mechanisms responsible for play and how the functions of play may relate to these neural substrates, the rat has become the model of choice. Play in the rat is easily quantified, tightly regulated, and can be modulated by genetic factors and postnatal experiences. Brain areas most likely to be involved in the modulation of play include regions within the prefrontal cortex, dorsal and ventral striatum, some regions of the amygdala, and habenula. This paper discusses what we currently know about the neurobiological substrates of play and how this can help illuminate functional questions about the putative benefits of play.
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Play is itself a social behavior and involves the activity of many regions in the social circuitry, such as the medial amygdala, lateral septum, and hypothalamus.
“given play is itself a social behavior and involves the activity of many regions in the social circuitry, such as the medial amygdala, lateral septum, and hypothalamus (summarized in Siviy, 2016 and VanRyzin et al., 2020a).”
From Social play experience in juvenile rats is indispensable for appropriate socio-sexual behavior in adulthood in males but not females (2023) · cites this paper as Siviy, 2016
The Introduction describes social rough-and-tumble play in rats, while noting that human play can also be solitary. The Subcortical involvement section links play to activity in the medial amygdala and hypothalamus, and reports that a lateral septal infusion changes play in rats; it does not establish play-related neural activity in the lateral septum. The claim therefore overgeneralizes both the social nature of play and the evidence for activity in every listed region (page numbers unavailable). (LLM, conf=0.85)
Experimental — accuracy not yet measuredLaboratory rats have been an important model species for studying the neurobiology of rough-and-tumble play.
“Laboratory rats have been an important model species with which to study the neurobiology of RTP (Siviy and Panksepp, 2011; Siviy, 2016; Vanderschuren et al., 2016).”
From The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain (2022) · cites this paper as Siviy, 2016
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“Dendritic complexity of pyramidal cells within the medial prefrontal cortex has also been shown to be sensitive to play experience as a juvenile (Bell et al., 2010).”
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“Vasopressin originating from cells within the paraventricular nucleus may also be a relevant modulator of play as a positive correlation has been reported in rats between levels of play and mRNA levels for vasopressin (Paul et al., 2014).”
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Marquardt, A. E., VanRyzin, J. W., Fuquen, R. W., & McCarthy, M. M. (2023) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2022.1076765 · Cited by 31
“This specificity to socio-sexual behavior makes sense, given play is itself a social behavior and involves the activity of many regions in the social circuitry, such as the medial amygdala, lateral septum, and hypothalamus (summarized in Siviy, 2016 and VanRyzin et al., 2020a).”
BackgroundThe rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Pellis, S. M., Pellis, V. C., Ham, J. R., & Achterberg, E. J. M. (2022) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2022.1033999 · Cited by 29
“Laboratory rats have been an important model species with which to study the neurobiology of RTP (Siviy and Panksepp, 2011; Siviy, 2016; Vanderschuren et al., 2016).”
What do you hear what do you say? Ultrasonic calls as signals during play fighting in rats
Burke, C. J., Euston, D. R., & Pellis, S. M. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1720126 · Cited by 23
Achterberg, E. J. M., Burke, C. J., & Pellis, S. M. (2023) · Behavioural Processes · doi:10.1016/j.beproc.2023.104933 · Cited by 20
Social behavior in prepubertal neurexin 1α deficient rats: A model of neurodevelopmental disorders.
Kight, K. E., Argue, K. J., Bumgardner, J. G., Bardhi, K., Waddell, J., & McCarthy, M. M. (2021) · Behavioral Neuroscience · doi:10.1037/bne0000482 · Cited by 17
Chapter 1: Problem-based learning: A 21st-century teaching and learning strategy
Research Unit Self-Directed Learning, F. O. E., Golightly, A., Sebatana, M. J., & School of Mathematics, S. A. T. E. (2023) · NWU Self-Directed Learning Series · doi:10.4102/aosis.2023.bk409.01 · Cited by 1
Ham, J. R., Iwaniuk, A. N., & Pellis, S. M. (2026) · Behavioural Brain Research · doi:10.1016/j.bbr.2026.116198 · Cited by 1
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
Metadata
- Authors
- Siviy, S. M.
- Affiliations
- Gettysburg College (US)
- Corresponding author
- Siviy, Stephen M.
- ORCID
- Siviy, Stephen M.
- Venue
- Behaviour
- Publisher
- Walter de Gruyter GmbH
- Published
- 2016
- Volume
- 153
- Issue
- 6-7
- Pages
- 819-844
- ISSN
- 0005-7959 (ISSN-L)
- 1568-539X
- PMC
- PMC5646690
- OpenAlex
- W2334239925
- Cited by
- 113
- References
- 20
- Citations by year
- 2026: 11 · 2025: 9 · 2024: 13 · 2023: 26 · 2022: 12 · 2021: 9 · 2020: 9 · 2019: 7 · 2018: 10 · 2017: 6 · 2016: 1
- FWCI
- 12.58
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Behaviour (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- The Cupola: Scholarship at Gettysburg College (Gettysburg College) (submittedVersion)
- Funders
- National Institute of Mental Health (R15MH100585)
- Topics
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Neuroendocrine regulation and behavior
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- playfulness, play behavior, prefrontal cortex, striatum, amygdala, habenula, behavioral flexibility, postnatal experience, genetic factors, child development
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