Differential rates of attack, defense, and counterattack during the developmental decrease in play fighting by male and female rats
Abstract
During postweaning development, rats exhibit several well documented trends in their play fighting: (1) It peaks between 30–40 days and then declines with the approach of sexual maturity; (2) males initiate more play fights than females; and (3) the overall complexity of play fights, as expressed by such measures as duration of bouts, also decreases with increasing age. Such trends could arise from changes in attack or defense, or some combination of both. In this article it is shown that (a) the decline in play fighting with the onset of sexual maturity in rats results from a decline in attack, not in defense; (b) the differences in play fighting by male and female rats are due to sex‐specific rates of both attack and defense; (c) the developmental decrease in the complexity of play fighting arises from a decrease in the frequency of counterattacks (i.e., after an animal defends itself, it is less likely to launch an attack). In this way, age and sex differences in play fighting can be traced to differences in its subcomponents.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Males are more likely to counterattack, engage in boxing, and pin more frequently than females.
“Males are also more likely to counterattack, engage in boxing, and pin more frequently, all of which may reflect the initial difference in the rate of play initiation (Pellis & Pellis, 1990).”
From Developmental origins of sex differences in the neural circuitry of play (2020) · cites this paper as Pellis & Pellis, 1990
The "turn and face" play behavior characteristic of LE rats may provide an opportunity to quantify more nuanced social interactions.
“though the “turn and face” play behavior characteristic of LE rats may provide an opportunity to quantify more nuanced social interactions [26–33].”
From Behavioral Phenotyping of Juvenile Long-Evans and Sprague-Dawley Rats: Implications for Preclinical Models of Autism Spectrum Disorders (2016) · cites this paper as [27]
Paradigms optimized for eliciting play behavior in LE rats utilize 24 hours of social isolation prior to testing, repeated habituation to the test cage, testing in complete darkness and the presence of bedding.
“In contrast, paradigms optimized for eliciting play behavior in LE rats utilize 24 hours of social isolation prior to testing, repeated habituation to the test cage, testing in complete darkness and the presence of bedding [27,41,57,82].”
From Behavioral Phenotyping of Juvenile Long-Evans and Sprague-Dawley Rats: Implications for Preclinical Models of Autism Spectrum Disorders (2016) · cites this paper as [27]
The animals from one cage were not used in the final analysis because this cage contained animals of two different ages and social play changes with age.
“The animals from one cage were not used in the final analysis because this cage contained animals of two different ages and social play changes with age [22, 23].”
From Sex-specific effects of neonatal progestin receptor antagonism on juvenile social play behavior in rats (2021) · cites this paper as [22]
Chronic exposure to males in the home cage can decrease how much females play even when the females are tested in same-sex pairs.
“although chronic exposure to males in the home cage can decrease how much females play even when the females are tested in same‐sex pairs (e.g., Pellis and Pellis 1990)”
From Partner Choice and Context‐Dependent Sex Differences in Rat Rough‐and‐Tumble Play (2026) · cites this paper as Pellis and Pellis 1990
In male rats, the probability of defense against a playful attack does not change with age.
“With regard to playful defense it has been shown that while the probability of defense against a playful attack does not change with age in males (Pellis & Pellis, 1990;Pellis et al., 1997), the use of complete rotations, in response to playful attack, decreases with the onset of puberty (Pellis & Pellis, 1990, 1991).”
From Play fighting in androgen-insensitivetfm rats: Evidence that androgen receptors are necessary for the development of adult playful attack and defense (2006) · cites this paper as Pellis & Pellis, 1990
It is unlikely that the procedures used in the present experiment account for the reversed sex difference.
“Given that the testing paradigm we used is the same as that which has been used to consistently show sex differences in the frequency of play fighting in Long Evans hooded rats (e.g., Pellis & Pellis, 1990;Pellis et al., 1994), it is unlikely that the procedures used in the present experiment account for the reversed sex difference.”
From Play fighting in androgen-insensitivetfm rats: Evidence that androgen receptors are necessary for the development of adult playful attack and defense (2006) · cites this paper as Pellis & Pellis, 1990
Pellis and Pellis described in Long Evans hooded male and female rats a peak of play fighting around 41-45 days of age in same-sex pairs.
“Pellis and Pellis [55] described in Long Evans hooded male and female rats a peak of play fighting around 41–45 days of age in same-sex pairs.”
From Developmental exposure to low levels of ethinylestradiol affects social play in juvenile male rats (2020) · cites this paper as [55]
In laboratory rats, rough-and-tumble play begins to emerge in the third week after birth, peaks between the fourth and fifth week, declines with the onset of puberty, and continues into adulthood at a lower level.
“In laboratory rats, RTP begins to emerge in the third week after birth, peaks in occurrence between the fourth and fifth week and then declines with the onset of puberty, but continues into adulthood, albeit at a lower level (Thor and Holloway, 1984; Pellis and Pellis, 1990, 1997).”
From The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain (2022) · cites this paper as Pellis and Pellis, 1990
Studies of social play in juvenile rats have revealed sex differences in playful defence behaviours, with males more likely to adopt a supine posture and females more likely to evade through lateral rotation.
“Studies of social play in juvenile rats have revealed sex differences in playful defence behaviours, with males more likely to adopt a supine posture and females more likely to evade through lateral rotation ( Pellis & McKenna, 1992; Pellis & Pellis, 1990).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Pellis & Pellis, 1990
Even when playing with males and being exposed to more nape attacks, females were more likely to evade with lateral rotation.
“Even when playing with males and being exposed to more nape attacks, females were more likely to evade with lateral rotation ( Pellis & Pellis, 1990);”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Pellis & Pellis, 1990
Juvenile male rats exhibit more playful initiation behaviour than females.
“Juvenile male rats exhibit more playful initiation behaviour than females ( Meaney & Stewart, 1981; Pellis & Pellis, 1990; Thor & Holloway, 1984) and produce more USVs than females during social play including during pinning ( Kisko et al., 2021).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Pellis & Pellis, 1990
Cites
The psychobiology of play: Theoretical and methodological perspectives
Panksepp, J., Siviy, S., & Normansell, L. (1984) · Neuroscience & Biobehavioral Reviews · doi:10.1016/0149-7634(84)90005-8 · Cited by 468 · see citing works
Developmental analyses of social play behavior in juvenile rats
Thor, D. H., & Holloway, W. R. (1984) · Bulletin of the Psychonomic Society · doi:10.3758/bf03333916 · Cited by 105 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
The adolescent brain and age-related behavioral manifestations
Spear, L. P. (2000) · Neuroscience & Biobehavioral Reviews · doi:10.1016/s0149-7634(00)00014-2 · Cited by 5,433 · see citing works
Douglas, L. A., Varlinskaya, E. I., & Spear, L. P. (2004) · Developmental Psychobiology · doi:10.1002/dev.20025 · Cited by 389 · see citing works
The neurobiology of social play and its rewarding value in rats
Vanderschuren, L. J. M. J., Achterberg, E. J. M., & Trezza, V. (2016) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2016.07.025 · Cited by 329 · see citing works
Burke, A. R., McCormick, C. M., Pellis, S. M., & Lukkes, J. L. (2017) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2017.01.018 · Cited by 225 · see citing works
Self-citationStress in adolescence and drugs of abuse in rodent models: role of dopamine, CRF, and HPA axis
Burke, A. R., & Miczek, K. A. (2014) · Psychopharmacology · doi:10.1007/s00213-013-3369-1 · Cited by 207 · see citing works
Vanderschuren, L. J. M. J., & Trezza, V. (2013) · Current Topics in Behavioral Neurosciences · doi:10.1007/978-3-662-45758-0_268 · Cited by 200 · see citing works
Sex-specific modulation of juvenile social play by vasopressin
Veenema, A. H., Bredewold, R., & De Vries, G. J. (2013) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2013.06.002 · Cited by 166 · see citing works
“Social play was assessed in 35-day-old juvenile rats because social play is highest at this age (Panksepp, 1981; Pellis & Pellis, 1990), following a procedure described in Veenema & Neumann (2009).”
MethodThiel, K. J., Okun, A. C., & Neisewander, J. L. (2008) · Drug and Alcohol Dependence · doi:10.1016/j.drugalcdep.2008.02.013 · Cited by 164 · see citing works
Smith, C. J. W., Poehlmann, M. L., Li, S., Ratnaseelan, A. M., Bredewold, R., & Veenema, A. H. (2017) · Brain Structure and Function · doi:10.1007/s00429-016-1260-7 · Cited by 143 · see citing works
Von Frijtag, J. C., Schot, M., van den Bos, R., & Spruijt, B. M. (2002) · Developmental Psychobiology · doi:10.1002/dev.10057 · Cited by 116 · see citing works
The role of the medial prefrontal cortex in the play fighting of rats.
Bell, H. C., McCaffrey, D. R., Forgie, M. L., Kolb, B., & Pellis, S. M. (2009) · Behavioral Neuroscience · doi:10.1037/a0017617 · Cited by 114 · see citing works
Self-citationHedonic sensitivity in adolescent and adult rats: Taste reactivity and voluntary sucrose consumption
Wilmouth, C. E., & Spear, L. P. (2009) · Pharmacology Biochemistry and Behavior · doi:10.1016/j.pbb.2009.02.009 · Cited by 96 · see citing works
Schneider, P., Bindila, L., Schmahl, C., Bohus, M., Meyer-Lindenberg, A., Lutz, B., Spanagel, R., & Schneider, M. (2016) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2016.00203 · Cited by 89 · see citing works
Himmler, B. T., Kisko, T. M., Euston, D. R., Kolb, B., & Pellis, S. M. (2014) · Behavioural Processes · doi:10.1016/j.beproc.2014.04.014 · Cited by 87 · see citing works
Self-citationRat tickling: A systematic review of applications, outcomes, and moderators.
LaFollette, M. R., O’Haire, M. E., Cloutier, S., Blankenberger, W. B., & Gaskill, B. N. (2017) · PLOS ONE · doi:10.1371/journal.pone.0175320 · Cited by 84 · see citing works
Himmler, B. T., Stryjek, R., Modlinska, K., Derksen, S. M., Pisula, W., & Pellis, S. M. (2013) · Journal of Comparative Psychology · doi:10.1037/a0032187 · Cited by 82 · see citing works
Self-citationPeering into the Dynamics of Social Interactions: Measuring Play Fighting in Rats
Himmler, B. T., Pellis, V. C., & Pellis, S. M. (2013) · Journal of Visualized Experiments · doi:10.3791/4288 · Cited by 81 · see citing works
Self-citationBurke, C. J., Kisko, T. M., Swiftwolfe, H., Pellis, S. M., & Euston, D. R. (2017) · PLOS ONE · doi:10.1371/journal.pone.0175841 · Cited by 78 · see citing works
Self-citationPlay fighting and the development of the social brain: The rat’s tale
Pellis, S. M., Pellis, V. C., Ham, J. R., & Stark, R. A. (2023) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2023.105037 · Cited by 76 · see citing works
Self-citationKu, K. M., Weir, R. K., Silverman, J. L., Berman, R. F., & Bauman, M. D. (2016) · PLOS ONE · doi:10.1371/journal.pone.0158150 · Cited by 74 · see citing works
Burleson, C. A., Pedersen, R. W., Seddighi, S., DeBusk, L. E., Burghardt, G. M., & Cooper, M. A. (2016) · Behavioral Neuroscience · doi:10.1037/bne0000148 · Cited by 69 · see citing works
The neurobiology of social play behaviour: Past, present and future
Achterberg, E. J. M., & Vanderschuren, L. J. M. J. (2023) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2023.105319 · Cited by 56 · see citing works
Raza, S., Himmler, B. T., Himmler, S. M., Harker, A., Kolb, B., Pellis, S. M., & Gibb, R. (2015) · Behavioural Pharmacology · doi:10.1097/fbp.0000000000000169 · Cited by 37 · see citing works
Self-citationDevelopmental origins of sex differences in the neural circuitry of play
VanRyzin, J. W., Marquardt, A. E., & McCarthy, M. M. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1723370 · Cited by 34 · see citing works
“Males are also more likely to counterattack, engage in boxing, and pin more frequently, all of which may reflect the initial difference in the rate of play initiation (Pellis & Pellis, 1990).”
BackgroundStark, R., & Pellis, S. M. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1720142 · Cited by 32 · see citing works
Self-citationMarquardt, 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 · see citing works
“Additionally, there are known sex differences in the qualitative characteristics of social play interactions (Pellis and Pellis, 1990, 1997).”
BackgroundField, E. F., Whishaw, I. Q., Pellis, S. M., & Watson, N. V. (2006) · Developmental Psychobiology · doi:10.1002/dev.20121 · Cited by 30 · see citing works
Self-citationThe 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 · see citing works
“In laboratory rats, RTP begins to emerge in the third week after birth, peaks in occurrence between the fourth and fifth week and then declines with the onset of puberty, but continues into adulthood, albeit at a lower level (Thor and Holloway, 1984; Pellis and Pellis, 1990, 1997).”
BackgroundSelf-citationWhat 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 · see citing works
Self-citationBlake, B. E., & McCoy, K. A. (2015) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2015.04.021 · Cited by 20 · see citing works
Sex differences in juvenile play behavior differ among rat strains
Northcutt, K. V., & Nwankwo, V. C. (2018) · Developmental Psychobiology · doi:10.1002/dev.21760 · Cited by 20 · see citing works
Pinning in the Play Fighting of Rats: A Comparative Perspective With Methodological Recommendations
Himmler, S. M., Himmler, B. T., Stryjek, R., Modlińska, K., Pisula, W., & Pellis, S. M. (2016) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2016.29.00.06 · Cited by 19 · see citing works
Self-citationField, E. F., Martens, D. J., Watson, N. V., & Pellis, S. M. (2005) · Journal of Comparative Psychology · doi:10.1037/0735-7036.119.2.238 · Cited by 7 · see citing works
Self-citationDevelopmental exposure to low levels of ethinylestradiol affects social play in juvenile male rats
Zaccaroni, M., Massolo, A., Beani, L., Seta, D. D., Farabollini, F., Giannelli, G., Fusani, L., & Dessì-Fulgheri, F. (2020) · Toxicological Research · doi:10.1007/s43188-019-00035-z · Cited by 5 · see citing works
Bombail, V., Brown, S. M., Beyts, C., Martin, J. E., Li, T., Meddle, S. L., Mendl, M., Robinson, E. S., Hammond, T. J., Nielsen, B. L., et al. (2025) · F1000Research · doi:10.12688/f1000research.166417.2 · Cited by 4 · see citing works
“Studies of social play in juvenile rats have revealedsexdifferences inplayfuldefencebehaviours,withmalesmorelikely toadoptasupinepostureandfemalesmore likely to evade through lateral rotation (Pellis & McKenna, 1992; Pellis & Pellis, 1990).”
Orsucci, I. C., Becker, K. D., Ham, J. R., Lee, J. D. A., Bowden, S. M., & Veenema, A. H. (2025) · Physiology & Behavior · doi:10.1016/j.physbeh.2025.115080 · Cited by 3 · 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
“…is, although chronic exposure to males in the home cage can decrease how much females play even when the females are tested in same-sex pairs (e.g., Pellis and Pellis 1990), our present data show that males can also influence the play of females acutely, even for females housed in same-sex pairs.”
Self-citationForbes-Lorman, R. M. (2021) · Behavioral and Brain Functions · doi:10.1186/s12993-021-00183-z · Cited by 1 · see citing works
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
Self-citationEvaluating 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-citation
- Authors
- Pellis, S. M., & Pellis, V. C.
- Affiliations
- University of Florida (US)
- Corresponding author
- Pellis, Sergio M.
- Venue
- Developmental Psychobiology
- Publisher
- Wiley
- Published
- 1990-04
- Volume
- 23
- Issue
- 3
- Pages
- 215-231
- ISSN
- 0012-1630 (ISSN-L)
- 1098-2302
- OpenAlex
- W2061863994
- Cited by
- 255
- References
- 2
- Citations by year
- 2026: 9 · 2025: 12 · 2024: 8 · 2023: 10 · 2022: 7 · 2021: 14 · 2020: 11 · 2019: 4 · 2018: 7 · 2017: 13 · 2016: 13 · 2015: 12 · 2014: 9 · 2013: 12 · 2012: 6
- FWCI
- 9.32
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Developmental Psychobiology (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
- Keywords
- play fighting, rats, sexual maturity, sex differences, attack behavior, defensive behavior, counterattack, post-weaning development, behavioral development, social behavior
- MeSH terms
- Aggression, Aging, Agonistic Behavior, Animals, Body Weight, Female, Male, Play and Playthings, Rats, Inbred Strains, Sex Factors, Rats
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review