The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Open accessAbstract
social play, rats, social dynamics, social skills, individual differences Play fighting, the most commonly reported form of social play, involves competition to gain an advantage (Aldis, 1975), but there are two features that make it different from serious fighting.First, it is highly pleasurable and associated with a positive affective state (Vanderschuren et al., 2016).Second, the competition is moderated by cooperation, ensuring that the interactions have a degree of reciprocity or turn taking between partners (Palagi et al., 2016a).That is, the player with the advantage may voluntarily relinquish it, thus allowing a role reversal to occur, with the original defender gaining the opportunity to become the attacker (Pellis and Pellis, 2017).Thus, although actions performed during play fighting can be accurately described as involving attack, defense, and counterattack, the context of their use should not be confused with that of aggression.For this reason, we will refer to this kind of play as rough-and-tumble play (RTP), to highlight the cooperative aspect of these interactions.Nonetheless, competition during RTP can create ambiguity as to whether a partner may be taking unfair advantage of the situation, with effective communication being important to avoid the risk of escalating to aggression or to partners being ostracized if they play too roughly (Palagi et al., 2016b).Creating and resolving ambiguity, which requires balancing competition and cooperation, also provides a vehicle by which juveniles and adolescents can train socio-cognitive skills (Pellis and Pellis, 2009, 2017).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).As shown in Figure 1, RTP in rats involves competition to gain access to the partner's nape of the neck, which is nuzzled with the snout if contacted (Pellis and Pellis, 1987;Siviy and Panksepp, 2011).A variety of tactics are used to attack and defend the nape, including launching counterattacks following a successful defense (Himmler et al., 2013;Pellis et al., 2022).Rats are a particularly good model species for studying RTP, as this behavior not only differs from serious fighting in the ways described above, but also because serious fighting involves attacking other body targets, namely the flanks and rump, which are bitten if contacted (Blanchard et al., 1977;Pellis and Pellis, 1987).Consequently, it can be readily discerned when a playful encounter escalates to serious fighting, as the aggressor switches from attacking the nape to biting the partner's posterior (Stark and Pellis, 2020).
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Written from the paper’s full text.
TL;DR
This review argues that rough-and-tumble play (RTP) in rats is a natural, cooperative-competitive behavior ideal for studying the social brain, with the mPFC and mesolimbic dopamine/opioid systems implicated in its regulation and reward. It highlights individual differences, partner choice/group dynamics, and the value of combining naturalistic observation with dyadic and operant experimental paradigms.
Key findings
- RTP in rats involves competition to gain access to the partner’s nape, which is nuzzled if contacted; serious fighting instead targets the flanks and rump and bites the posterior, so escalation can be discerned when an aggressor switches from nape attack to biting the partner’s posterior.
- RTP begins to emerge in the third week after birth, peaks between the fourth and fifth week, declines with puberty, and continues into adulthood at a lower level; once play begins around 17 days of age, the repertoire of tactics takes about 10 days to mature fully, with little influence of rearing experiences.
- Cortical systems, especially the medial prefrontal cortex (mPFC), are critical for modulating playful responses depending on partner actions and identity; juvenile RTP with peers (about 28–40 days post birth) influences mPFC development and socio-cognitive skills.
- Individual differences exist: high players initiate more nape attacks and preferentially use different defensive tactics than low players; mesolimbic dopamine activity is linked to launching nape attacks and to motivation to work for access to a playmate, while opioid systems are implicated in play reward. Group testing shows rats prefer some partners over others, and the standard dyadic paradigm removes partner choice.
Methods
- Design: conceptual/review article, not a primary empirical study; it synthesizes existing literature and proposes combining naturalistic observations with experimental manipulations.
- Data or sample: no new empirical sample or dataset is reported; the paper discusses laboratory rat studies and common behavioral paradigms, including home-enclosure/litter observations and dyadic testing after social isolation, with fixed test durations of 5–20 min being most common.
- Main analysis: narrative synthesis and conceptual argument, with illustrative examples and references; no statistical analysis is reported.
Limitations
- Stated: the dyadic paradigm gives experimental control but removes the rats’ ability to choose their play partner; experimental manipulations may alter partner discrimination and go undetected in dyadic tests, potentially leading to the false conclusion that a manipulation has no effect.
- Stated: open questions remain about exactly which neural circuits are associated with what types of playful actions and styles of play, and how cortical and subcortical circuits regulating affective and aversive ultrasonic vocalizations are modulated to accommodate different play styles.
- Evident from the text: as a review/perspective, the paper presents no new empirical data, sample, or statistical results; its conclusions are synthesized from cited studies.
Model-written summary. Check it against the paper before relying on it.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
21 of 69 cited sources checked · 8 supported · 0 flagged · accuracy not yet measured
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
In play fighting, the player with the advantage may voluntarily relinquish it, allowing a role reversal in which the original defender can become the attacker.
“That is, the player with the advantage may voluntarily relinquish it, thus allowing a role reversal to occur, with the original defender gaining the opportunity to become the attacker (Pellis and Pellis, 2017).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Pellis and Pellis, 2017 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Laboratory 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).”
Quote in the paperSubstantial claim1 of 3 sources checked - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
The medial prefrontal cortex has a crucial role in coordinating actions with partners as juveniles and as adults in both playful and non-playful social interactions.
“and the mPFC has a crucial role in coordinating actions with partners as juveniles and as adults in both playful and non-playful social interactions (Bell et al., 2009; Van Kerkhof et al., 2013; Himmler et al., 2014; Stark and Pellis, 2020).”
Quote in the paperSubstantial claim1 of 3 sources checked- cites Bell et al., 2009 source
- cites Van Kerkhof et al., 2013 source
The Concluding Remarks say that medial PFC activity helps adolescent rats initiate play and respond appropriately to a partner’s play initiation. The paper’s discussion of non-playful social exploration does not establish a role in coordinating those interactions, and its mention of adult sexual behavior concerns neuronal activity rather than coordinated actions. The claim therefore extends beyond the paper’s findings. (LLM, conf=0.87)
Experimental — accuracy not yet measured - cites Himmler et al., 2014 source
- Premisecitation
More sophisticated methods are becoming available for continuous recording and scoring of behavior in a home cage.
“Indeed, more sophisticated methods are becoming available for continuous recording and scoring of behavior in a home cage (Greico et al., 2021).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Greico et al., 2021 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Rats have a degree of sophistication in their rough-and-tumble play comparable to that of many primates and other social mammals.
“rats have a degree of sophistication in their RTP that is comparable to that of many primates and other social mammals (Pellis and Pellis, 2017)”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Pellis and Pellis, 2017 source
In “The many routes to restraint during play fighting” (page unavailable), the paper describes rats’ tactical role reversals and reports a similar pattern in spider monkeys and macaques. This supports a comparison with some primates, but it does not establish comparable overall sophistication across many primates and other social mammals. (LLM, conf=0.82)
Experimental — accuracy not yet measured
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Under some rearing and testing conditions, male rats initiate more nape attacks.
“under some rearing and testing conditions, males initiate more nape attacks (Thor and Holloway, 1984)”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Thor and Holloway, 1984 source
In “WHEN DOES PLAY FIGHTING BEGIN?” (page unavailable), the paper reports that males initiated play by pouncing more often than females at ages 26–55 days, but not at ages 21–25 days. That supports a condition-dependent difference in play initiation; our copy does not specify that the pounces were nape attacks or link this difference to particular rearing and testing conditions. (LLM, conf=0.80)
Experimental — accuracy not yet measured
- Premisecitation
There is strong evidence that the difference in launching nape attacks is linked to differences in mesolimbic dopamine activity.
“There is strong evidence that the difference in launching nape attacks is linked to differences in mesolimbic dopamine activity (Vanderschuren et al., 2016; Siviy, 2020).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Vanderschuren et al., 2016 source
- cites Siviy, 2020 source
In “The use of Fischer 344 rats in play research” (page unavailable), the paper reports strain differences in nape contacts. In “Using strain differences to study the underlying neurobiological substrates of play” (page unavailable), it reports dopamine-related differences and evidence that nucleus accumbens dopamine modulates play, but says it remains unclear whether impaired striatal dopamine function accounts for the F344 rats’ lower playfulness. It therefore does not support the claim of strong evidence linking the difference in launching nape attacks specifically to mesolimbic dopamine activity. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
The development of the medial prefrontal cortex and associated socio-cognitive skills is influenced by the experience of rough-and-tumble play with peers in the juvenile period (about 28-40 days post birth).
“the development of the medial prefrontal cortex (mPFC) and associated socio-cognitive skills is influenced by the experience of RTP with peers in the juvenile period (about 28–40 days post birth; Bell et al., 2010; Baarendse et al., 2013; Schneider et al., 2016)”
Quote in the paperSubstantial claim2 of 3 sources checked- cites Bell et al., 2010 source
- cites Baarendse et al., 2013 source
The Introduction and Discussion link juvenile social experience to prefrontal development and cognitive control, and report altered mPFC dopamine sensitivity after social isolation in rats. But the study isolated rats during postnatal days 21–42; it did not isolate the effects of rough-and-tumble play or establish the narrower 28–40-day window. No page numbers are provided in our copy. (LLM, conf=0.86)
Experimental — accuracy not yet measured - cites Schneider et al., 2016 source
The Introduction says peer play is crucial for social competence and reports that play deprivation interferes with fronto-cortical development. The Methods examine play on postnatal days 30 and 35, but the copy does not establish the full 28–40-day window or report a medial prefrontal cortex finding from this study. (LLM, conf=0.82)
Experimental — accuracy not yet measured
- Premisecitation
Even within members of the same sex and same strain, not all individual rats play to the same degree; some rats consistently play more than others.
“Even within members of the same sex and same strain, not all individuals play to the same degree—some rats consistently play more than others (Lampe et al., 2017; Lesscher et al., 2021).”
Quote in the paperSubstantial claim1 of 2 sources checked - Interpretationcitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Sex and strain differences in play have provided valuable tools for identifying the roles of specific neural systems and networks in regulating particular aspects of play.
“These differences have provided valuable tools for using sex and strain differences to identify the roles of specific neural systems and neural networks in regulating particular aspects of play (Siviy, 2020; VanRyzin et al., 2020).”
Quote in the paperSubstantial claim2 of 2 sources checked- cites Siviy, 2020 source
The section “Using strain differences to study the underlying neurobiological substrates of play” describes using behavioral differences between rat strains to investigate neural systems involved in play. The passages from our copy do not establish the same use of sex differences, so Siviy (2020) supports the strain component but not the full claim. (LLM, conf=0.84)
Experimental — accuracy not yet measured - cites VanRyzin et al., 2020 source
The Introduction says sex differences in play provide an experimental means of identifying how specific brain regions contribute to behavior. The section on sexually differentiated modulatory nodes also describes sex differences as a way to distinguish play-specific features of neural circuitry. Our copy does not support the claim about strain differences serving that purpose. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Premisecitation
Dopamine manipulations affect how hard rats will work for the reward of access to a playmate.
“in which dopamine manipulations affect how hard rats will work for the reward of access to a playmate (Achterberg et al., 2016).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Achterberg et al., 2016 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
The motivation to play can be modified by early experiences mostly derived from the mother.
“While the motivation to play can be modified by early experiences mostly derived from the mother (Parent and Meaney, 2008; Van Hasselt et al., 2012)”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Parent and Meaney, 2008 source
- cites Van Hasselt et al., 2012 source
The Discussion (page not provided) reports that maternal care predicts adolescent social play in male rats, and describes the play test as probing social motivation. But it found no such correlation in females and explicitly cautions that the correlation does not establish that maternal care causes play to develop. The claim omits those limits and presents modification by early experience more broadly. (LLM, conf=0.84)
Experimental — accuracy not yet measured
- Interpretationcitation
Identifying that turn taking, as shown by role reversals, is a key feature of rough-and-tumble play that promotes the development of socio-cognitive skills has been important for engineering therapeutic play contexts that promote those skills in children.
“Indeed, identifying that turn taking, as shown by the occurrence of role reversals, is a key feature of RTP that promotes the development of socio-cognitive skills (Pellis et al., 2019; Stark et al., 2021), has been important for engineering therapeutic play contexts that similarly promote the development of those skills in children (e.g., Diamond et al., 2007; Nijhoff et al., 2018).”
Quote in the paperNot a substantial claim3 of 3 sources checked- cites Pellis et al., 2019 source
Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measured - cites Stark et al., 2021 source
- cites Diamond et al., 2007 source
Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measured
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
It can be readily discerned when a playful encounter escalates to serious fighting, as the aggressor switches from attacking the nape to biting the partner's posterior.
“Consequently, it can be readily discerned when a playful encounter escalates to serious fighting, as the aggressor switches from attacking the nape to biting the partner's posterior (Stark and Pellis, 2020).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Stark and Pellis, 2020 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Serious fighting in rats involves attacking other body targets, namely the flanks and rump, which are bitten if contacted.
“serious fighting involves attacking other body targets, namely the flanks and rump, which are bitten if contacted (Blanchard et al., 1977; Pellis and Pellis, 1987).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Play fighting involves competition to gain an advantage.
“Play fighting, the most commonly reported form of social play, involves competition to gain an advantage (Aldis, 1975), but there are two features that make it different from serious fighting.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Aldis, 1975 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
In play fighting, competition is moderated by cooperation, ensuring a degree of reciprocity or turn taking between partners.
“Second, the competition is moderated by cooperation, ensuring that the interactions have a degree of reciprocity or turn taking between partners (Palagi et al., 2016a).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Palagi et al., 2016a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Although actions during play fighting involve attack, defense, and counterattack, their context should not be confused with aggression.
“Thus, although actions performed during play fighting can be accurately described as involving attack, defense, and counterattack, the context of their use should not be confused with that of aggression.”
Quote in the paperSubstantial claim - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Competition during rough-and-tumble play can create ambiguity as to whether a partner may be taking unfair advantage of the situation.
“Nonetheless, competition during RTP can create ambiguity as to whether a partner may be taking unfair advantage of the situation”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Palagi et al., 2016b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Rats are a particularly good model species for studying rough-and-tumble play.
“Rats are a particularly good model species for studying RTP, as this behavior not only differs from serious fighting in the ways described above, but also because serious fighting involves attacking other body targets, namely the flanks and rump, which are bitten if contacted (Blanchard et al., 1977; Pellis and Pellis, 1987).”
Quote in the paperSubstantial claim - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Effective communication during rough-and-tumble play is important to avoid the risk of escalating to aggression or of partners being ostracized if they play too roughly.
“with effective communication being important to avoid the risk of escalating to aggression or to partners being ostracized if they play too roughly (Palagi et al., 2016b).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Palagi et al., 2016b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Initial studies of play in rats involved observing them in their home enclosures with the whole litter present to assess the occurrence of rough-and-tumble play.
“Initial studies of play in rats involved observing them in their home enclosures with the whole litter present to assess the occurrence of RTP (Baeninger, 1967; Meaney and Stewart, 1981; Pellis and Pellis, 1983).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation
Attack and defense during rough-and-tumble play tend to involve independent mechanisms.
“For instance, attack and defense during RTP tend to involve independent mechanisms (Himmler et al., 2016).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Himmler et al., 2016 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
There are differences across rat strains in the relative frequency of nape attacks and in the use of defensive tactics that lead to evading contact or promoting close-quarter wrestling.
“Similarly, there are differences across strains in the relative frequency of nape attacks and in the use of defensive tactics that lead to evading contact or promoting close-quarter wrestling (Himmler et al., 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Himmler et al., 2016 source
- Interpretationreasoning
A collective paradigm, although naturalistic, makes testing the effects of specific treatments on play nearly impossible.
“Such a collective paradigm, although naturalistic, makes testing the effects of specific treatments on play nearly impossible.”
Quote in the paperSubstantial claim - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
In early adulthood, male rats are more likely to use defensive tactics that make their rough-and-tumble play seem rougher.
“in early adulthood are more likely to use defensive tactics that makes their RTP seem rougher (Pellis, 2002).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Pellis, 2002 source
- Interpretationcitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
There are subtle differences between sexes and strains of rats in play that can be informative.
“Even though both sexes and all strains of rats thus far studied share the same basic play repertoire, there are subtle differences that can be informative.”
Quote in the paperSubstantial claim - Premisecitation
The uneven distribution of play in groups is not simply a by-product of proximity; rats will travel to initiate play with a particular animal.
“and this is not simply a by-product of which of the animals are in closest proximity—rats will leave the company of animals in one part of the enclosure and travel to the other side to initiate play with a particular animal (Pellis et al., 2022).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
- Interpretationreasoning§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Social play is a valuable window into the social brain.
“That is, social play is a valuable window into the social brain.”
Quote in the paperNot a substantial claim - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Once play begins at around 17 days of age, it takes about 10 days for the repertoire of tactics used during play to mature fully.
“once play begins at around 17 days of age, it takes about 10 days for the repertoire of tactics used during play to mature fully”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Himmler et al., 2015 source
- Premisecitation
A dyadic paradigm was developed in which rats are tested in pairs in an enclosure to which they have been habituated following a period of social isolation to increase their motivation to engage in play.
“Consequently, a dyadic paradigm was developed—rats are tested in pairs in an enclosure to which they have been habituated following a period of social isolation to increase their motivation to engage in play (Panksepp and Beatty, 1980; Panksepp, 1981).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
When rats are tested in groups with multiple partners available, play is not distributed evenly; some partners are favored over others.
“When rats are tested in groups, in which multiple partners are available, play is not distributed evenly. Rather, some partners are favored over others”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
- Interpretationreasoning
Individual differences in play provide an opportunity to refine the search for the neural mechanisms that regulate play.
“Such individual differences provide an opportunity to refine the search for the neural mechanisms that regulate play.”
Quote in the paperSubstantial claim - Interpretationreasoning
The downside to the level of control achieved by the dyadic paradigm is that rats lose the ability to choose their play partner, as the experimenter selects the partner.
“However, the downside to the level of control achieved by the dyadic paradigm is that the rats lose the ability to choose their play partner, as it is the experimenter who selects the partner.”
Quote in the paperNot a substantial claim - Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
The maturation of the play tactic repertoire appears to be little influenced by rearing experiences.
“this maturation appears to be little influenced by rearing experiences (Himmler et al., 2015).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Himmler et al., 2015 source
- Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brain
Cortical systems, especially those of the prefrontal cortex, are critical to allow rats to modulate aspects of their playful responses depending on their partner's actions and identity.
“but cortical systems, especially those of the prefrontal cortex, are critical to allow rats to modulate aspects of their playful responses, depending on their partner's actions and their identity (Pellis and Pellis, 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis and Pellis, 2016
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
Outliers in naturally occurring behavior like rough-and-tumble play can be highly informative about underlying biological processes, rather than being viewed as noise that interferes with interpretation and replication.
“As indicated by the two examples above, so-called outliers in naturally occurring behavior like RTP can be highly informative about underlying biological processes, rather than, as is typically the case in many contrived experiments, being viewed as noise that interferes with the interpretation of results and future replication.”
Quote in the paperSubstantial claim - Premisecitation
In a group setting, rats preferentially play with particular partners.
“In a group setting, rats preferentially play with particular partners (Pellis et al., 2022)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
- May combine two claims. 12
- Classified as probably not a substantial claim. 6
May combine two claims.
Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainA variety of tactics are used to attack and defend the nape, including launching counterattacks following a successful defense.
“A variety of tactics are used to attack and defend the nape, including launching counterattacks following a successful defense (Himmler et al., 2013; Pellis et al., 2022).”
Quote in the paperNot a substantial claim1 of 2 sources checkedMay combine two claims.
PremisecitationMesolimbic dopamine activity affects not just the launching of nape attacks but also the motivation to engage in play.
“That mesolimbic dopamine activity does not just affect the launching of nape attacks, but also the motivation to engage in play, has been shown by operant conditioning methods”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Achterberg et al., 2016 source
The ‘Dissociable Roles of Dopamine and Noradrenaline in Social Play Motivation and Expression’ section reports that dopamine-enhancing drugs increased operant responding for social play, supporting an effect on motivation. The ‘Operant Conditioning’ section identifies pouncing as an attempt to contact a partner’s nape, and the study measures it separately. But the drugs were not targeted to the mesolimbic pathway, so the study does not directly establish the claim’s regional specificity. (LLM, conf=0.82)
Experimental — accuracy not yet measured
May combine two claims.
Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainIn 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).”
Quote in the paperNot a substantial claim1 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationAttempts to explain the neural mechanisms associated with individual differences in rough-and-tumble play have started to emerge in the literature.
“However, attempts to explain the neural mechanisms associated with individual differences in RTP have started to emerge in the literature (e.g., Reppucci et al., 2020).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Reppucci et al., 2020 source
The Discussion (no page available) says neural mechanisms underlying motivated social behavior can differ between males and females and cites studies of juvenile social play. But this paper tests social-versus-food preference, not the neural mechanisms behind individual differences in rough-and-tumble play; its methods say playful behaviors were not observed during the test. (LLM, conf=0.78)
Experimental — accuracy not yet measured
May combine two claims.
Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainRough-and-tumble play in rats involves competition to gain access to the partner's nape of the neck, which is nuzzled with the snout if contacted.
“As shown in Figure 1, RTP in rats involves competition to gain access to the partner's nape of the neck, which is nuzzled with the snout if contacted (Pellis and Pellis, 1987; Siviy and Panksepp, 2011).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainPlay fighting is highly pleasurable and associated with a positive affective state.
“First, it is highly pleasurable and associated with a positive affective state (Vanderschuren et al., 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Vanderschuren et al., 2016 source
May combine two claims.
InterpretationcitationExperimental manipulations may alter the ability of rats to discriminate between partners, and this may go undetected in the standard dyadic play paradigm, potentially leading to the false conclusion that the manipulation has no effect.
“Experimental manipulations may alter the ability of rats to discriminate between partners (Pellis et al., 2006), and this may go undetected in the standard, dyadic play paradigm, in which interaction with only one partner is possible, potentially leading to the false conclusion that the experimental manipulation has no effect as the amount of play is the same as that of the control.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis et al., 2006 source
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationOpioid systems have been implicated in the rewards derived from engaging in play.
“While opioid systems have been implicated in the rewards derived from engaging in play (Vanderschuren et al., 2016; Achterberg et al., 2019)”
Quote in the paperNot a substantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationThe dyadic paradigm has become the most widely used experimental paradigm for testing rough-and-tumble play in rats.
“This dyadic paradigm has now become the most widely used experimental paradigm for testing RTP in rats (Pellis et al., 2022).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
May combine two claims.
Premisecitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainNeither the age-typical changes in the frequency of rough-and-tumble play nor the availability of the full behavioral repertoire used during rough-and-tumble play depend on the cortex.
“Neither the age-typical changes in the frequency of RTP nor the availability of the full behavioral repertoire used during RTP depend on the cortex”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis and Pellis, 2016
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
PremisecitationWithin a rat strain, high players initiate more nape attacks and preferentially use a different suite of rat-typical defensive tactics than low players.
“Within a strain, high players not only initiate more nape attacks, but also preferentially use a different suite of the rat-typical defensive tactics than do low players (Pellis et al., 2022).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
May combine two claims.
Interpretationcitation§ The rough-and-tumble play of rats as a natural behavior suitable for studying the social brainRats are an ideal laboratory species to study play and, by extension, aspects of the social brain that make play possible.
“they are an ideal laboratory species to study not only play, but also, by extension, aspects of the social brain that make play possible.”
Quote in the paperSubstantial claim- Rests on an assumption: Rough-and-tumble play is naturally occurring.
May combine two claims.
InterpretationcitationRats with different styles of rough-and-tumble play may need to modify their use of ultrasonic vocalizations to play together effectively, making rough-and-tumble play in rats a useful window into subtle social communication processes.
“rats with different styles of RTP may need to modify their use of such calls (Pellis et al., 2022) to play together effectively, making RTP in rats a useful window into subtle social communication processes.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pellis et al., 2022 source
Probably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningMulti-animal paradigms are needed to offset the disadvantages of the dyadic paradigm.
“Thus, multi-animal paradigms are needed to offset the disadvantages that have come with the advantages gained from the dyadic paradigm.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationThe use of ultrasonic vocalizations can be critical for communication during play, facilitating role reversals and avoiding escalation to aggression.
“As the use of ultrasonic vocalizations (USV) can be critical for communication during play, facilitating role reversals and avoiding escalation to aggression (Kisko et al., 2017; Burke et al., 2020)”
Quote in the paperSubstantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S2 (checkable), S4 (based).
InterpretationreasoningSome degree of experimental control over behavior may be needed to identify what is most relevant to the animals.
“However, while outliers can focus our attention on phenomena that may be missed when comparing group means, some degree of experimental control over the behavior may be needed to identify what is most relevant to the animals.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationStudying rough-and-tumble play can be a valuable tool both for basic research on communication and other processes involved in regulating social behavior and for translational research on neurodevelopmental disorders.
“Studying RTP can be a valuable tool both for basic research on communication and other processes that are involved in regulating social behavior (Palagi et al., 2016b), and for translational research on neurodevelopmental disorders (Burke et al., 2017).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Palagi et al., 2016b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Burke et al., 2017 source
Probably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningNaturalistic observations and experimental manipulations can be used together in an iterative manner.
“That is, naturalistic observations and experimental manipulations can be used together in an iterative manner.”
Quote in the paperNot a substantial claim
The paper relies on this without support.
Rough-and-tumble play is naturally occurring.
“Given that RTP is naturally occurring, no experimental training is needed to teach the animals to play.”
Cites
Preschool Program Improves Cognitive Control
Diamond, A., Barnett, W. S., Thomas, J., & Munro, S. (2007) · Science · doi:10.1126/science.1151148 · Cited by 2,025 · see citing works
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“Consequently, a dyadic paradigm was developed—rats are tested in pairs in an enclosure to which they have been habituated following a period of social isolation to increase their motivation to engage in play (Panksepp and Beatty, 1980; Panksepp, 1981).”
BackgroundEthotransmission: communication of emotional states through ultrasonic vocalization in rats.
Brudzynski, S. M. (2013) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2013.01.014 · Cited by 416 · cited 2× · see citing works
“As the use of ultrasonic vocalizations (USV) can be critical for communication during play, facilitating role reversals and avoiding escalation to aggression (Kisko et al., 2017; Burke et al., 2020), rats with different styles of RTP may need to modify their use of such calls (Pellis et al., 2022) to play together eff…”
BackgroundSocial deprivation and play in rats
Panksepp, J., & Beatty, W. W. (1980) · Behavioral and Neural Biology · doi:10.1016/s0163-1047(80)91077-8 · Cited by 409 · cited 1× · see citing works
“Consequently, a dyadic paradigm was developed—rats are tested in pairs in an enclosure to which they have been habituated following a period of social isolation to increase their motivation to engage in play (Panksepp and Beatty, 1980; Panksepp, 1981).”
BackgroundA descriptive study of social development in the rat (Rattus norvegicus)
Meaney, M. J., & Stewart, J. (1981) · Animal Behaviour · doi:10.1016/s0003-3472(81)80149-2 · Cited by 403 · cited 1× · see citing works
“Initial studies of play in rats involved observing them in their home enclosures with the whole litter present to assess the occurrence of RTP (Baeninger, 1967; Meaney and Stewart, 1981; Pellis and Pellis, 1983).”
BackgroundAttack and defensive behaviour in the albino rat
Blanchard, R. J., Blanchard, D. C., Takahashi, T., & Kelley, M. J. (1977) · Animal Behaviour · doi:10.1016/0003-3472(77)90113-0 · Cited by 371 · cited 1× · see citing works
“Rats are a particularly good model species for studying RTP, as this behavior not only differs from serious fighting in the ways described above, but also because serious fighting involves attacking other body targets, namely the flanks and rump, which are bitten if contacted (Blanchard et al., 1977; Pellis and Pellis…”
BackgroundNijhof, S. L., Vinkers, C. H., van Geelen, S. M., Duijff, S. N., Achterberg, E. J. M., van der Net, J., Veltkamp, R. C., Grootenhuis, M. A., van de Putte, E. M., Hillegers, M. H. J., et al. (2018) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2018.09.024 · Cited by 363 · cited 1× · see citing works
“, 2021), has been important for engineering therapeutic play contexts that similarly promote the development of those skills in children (e.g., Diamond et al., 2007; Nijhoff et al., 2018).”
BackgroundSelf-citationThe 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
Self-citationPellis, S. M., & Pellis, V. C. (1990) · Developmental Psychobiology · doi:10.1002/dev.420230303 · Cited by 255 · cited 1× · 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-citationBurke, 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 · cited 1× · see citing works
“, 2016b), and for translational research on neurodevelopmental disorders (Burke et al., 2017).”
BackgroundSelf-citationBell, H. C., Pellis, S. M., & Kolb, B. (2010) · Behavioural Brain Research · doi:10.1016/j.bbr.2009.09.029 · Cited by 217 · cited 1× · see citing works
“For example, the development of the medial prefrontal cortex (mPFC) and associated socio-cognitive skills is influenced by the experience of RTP with peers in the juvenile period (about 28–40 days post birth; Bell et al., 2010; Baarendse et al., 2013; Schneider et al., 2016), and the mPFC has a crucial role in coordin…”
BackgroundSelf-citationRough‐and‐tumble play as a window on animal communication
Palagi, E., Burghardt, G. M., Smuts, B., Cordoni, G., Dall'Olio, S., Fouts, H. N., Řeháková‐Petrů, M., Siviy, S. M., & Pellis, S. M. (2016) · Biological Reviews · doi:10.1111/brv.12172 · Cited by 209 · see citing works
Self-citationBaarendse, P. J. J., Counotte, D. S., O'Donnell, P., & Vanderschuren, L. J. M. J. (2013) · Neuropsychopharmacology · doi:10.1038/npp.2013.47 · Cited by 206 · cited 2× · see citing works
“…socio-cognitive skills is influenced by the experience of RTP with peers in the juvenile period (about 28–40 days post birth; Bell et al., 2010; Baarendse et al., 2013; Schneider et al., 2016), and the mPFC has a crucial role in coordinating actions with partners as juveniles and as adults in…”
BackgroundThe influence of natural variations in maternal care on play fighting in the rat
Parent, C. I., & Meaney, M. J. (2008) · Developmental Psychobiology · doi:10.1002/dev.20342 · Cited by 175 · cited 1× · see citing works
“While themotivation to play can be modified by early experiences mostly derived from the mother (Parent and Meaney, 2008; Van Hasselt et al., 2012), once play begins at around 17 days of age, it takes about 10 days for the repertoire of tactics used during play to mature fully and this maturation appears to be little…”
BackgroundIn search of the neurobiological substrates for social playfulness in mammalian brains
Siviy, S. M., & Panksepp, J. (2011) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2011.03.006 · Cited by 162 · cited 2× · see citing works
“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).”
Backgroundvan Kerkhof, L. W., Damsteegt, R., Trezza, V., Voorn, P., & Vanderschuren, L. J. (2013) · Neuropsychopharmacology · doi:10.1038/npp.2013.83 · Cited by 161 · see citing works
What is play fighting and what is it good for?
Pellis, S. M., & Pellis, V. C. (2017) · Learning & Behavior · doi:10.3758/s13420-017-0264-3 · Cited by 160 · cited 3× · see citing works
“Moreover, as rats have a degree of sophistication in their RTP that is comparable to that of many primates and other social mammals (Pellis and Pellis, 2017), they are an ideal laboratory species to study not only play, but also, by extension, aspects of the social brain that make play possible.”
BackgroundSelf-citationComparison of behavioural development in socially isolated and grouped rats
Baenninger, L. P. (1967) · Animal Behaviour · doi:10.1016/0003-3472(67)90018-8 · Cited by 139 · cited 2× · see citing works
“Group dynamics and partner choice Initial studies of play in rats involved observing them in their home enclosures with the whole litter present to assess the occurrence of RTP (Baeninger, 1967; Meaney and Stewart, 1981; Pellis and Pellis, 1983).”
BackgroundPellis, S. M. (2002) · Archives of Sexual Behavior · doi:10.1023/a:1014070916047 · Cited by 124 · cited 1× · see citing works
“For example, under some rearing and testing conditions, males initiate more nape attacks (Thor and Holloway, 1984), and in early adulthood are more likely to use defensive tactics that makes their RTP seem rougher (Pellis, 2002).”
BackgroundSelf-citationPellis, S. M., Hastings, E., Shimizu, T., Kamitakahara, H., Komorowska, J., Forgie, M. L., & Kolb, B. (2006) · Behavioral Neuroscience · doi:10.1037/0735-7044.120.1.72 · Cited by 116 · cited 2× · see citing works
“Not all partners are equally attractive as play mates (Holloway and Suter, 2004; Pellis et al., 2006).”
BackgroundSelf-citationThe 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 · cited 1× · see citing works
“, 2016), and the mPFC has a crucial role in coordinating actions with partners as juveniles and as adults in both playful and non-playful social interactions (Bell et al., 2009; Van Kerkhof et al., 2013; Himmler et al., 2014; Stark and Pellis, 2020).”
BackgroundSelf-citationA brain motivated to play: insights into the neurobiology of playfulness
Siviy, S. M. (2016) · Behaviour · doi:10.1163/1568539x-00003349 · Cited by 113 · cited 1× · see citing works
“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).”
Measuring behavior in the home cage: study design, applications, challenges, and perspectives
Grieco, F., Bernstein, B. J., Biemans, B., Bikovski, L., Burnett, C. J., Cushman, J. D., van Dam, E. A., Fry, S. A., Richmond-Hacham, B., Homberg, J. R., et al. (2021) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2021.735387 · Cited by 109 · cited 1× · see citing works
“Indeed, more sophisticated methods are becoming available for continuous recording and scoring of behavior in a home cage (Greico et al., 2021).”
BackgroundDevelopmental 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 · cited 2× · see citing works
“For example, under some rearing and testing conditions, males initiate more nape attacks (Thor and Holloway, 1984), and in early adulthood are more likely to use defensive tactics that makes their RTP seem rougher (Pellis, 2002).”
BackgroundAchterberg, E. J. M., van Kerkhof, L. W. M., Servadio, M., van Swieten, M. M. H., Houwing, D. J., Aalderink, M., Driel, N. V., Trezza, V., & Vanderschuren, L. J. M. J. (2016) · Neuropsychopharmacology · doi:10.1038/npp.2015.212 · Cited by 103 · cited 1× · see citing works
“That mesolimbic dopamine activity does not just affect the launching of nape attacks, but also the motivation to engage in play, has been shown by operant conditioning methods, in which dopamine manipulations affect how hard rats will work for the reward of access to a playmate (Achterberg et al., 2016).”
BackgroundSelf-citationvan Hasselt, F. N., Tieskens, J. M., Trezza, V., Krugers, H. J., Vanderschuren, L. J. M. J., & Joëls, M. (2012) · Physiology & Behavior · doi:10.1016/j.physbeh.2011.12.007 · Cited by 91 · 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 · cited 1× · see citing works
“For example, the development of the medial prefrontal cortex (mPFC) and associated socio-cognitive skills is influenced by the experience of RTP with peers in the juvenile period (about 28–40 days post birth; Bell et al., 2010; Baarendse et al., 2013; Schneider et al., 2016), and the mPFC has a crucial role in coordin…”
BackgroundPeering 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 · cited 1× · see citing works
“A variety of tactics are used to attack and defend the nape, including launching counterattacks following a successful defense (Himmler et al., 2013; Pellis et al., 2022).”
BackgroundSelf-citationFair play and its connection with social tolerance, reciprocity and the ethology of peace
Palagi, E., Cordoni, G., Demuru, E., & Bekoff, M. (2016) · Behaviour · doi:10.1163/1568539x-00003336 · Cited by 72 · see citing works
Pellis, S. M., & Pellis, V. C. (1983) · Developmental Psychobiology · doi:10.1002/dev.420160403 · Cited by 71 · cited 2× · see citing works
“Group dynamics and partner choice Initial studies of play in rats involved observing them in their home enclosures with the whole litter present to assess the occurrence of RTP (Baeninger, 1967; Meaney and Stewart, 1981; Pellis and Pellis, 1983).”
BackgroundSelf-citationPlay, variation in play and the development of socially competent rats
Himmler, S. M., Himmler, B. T., Pellis, V. C., & Pellis, S. M. (2016) · Behaviour · doi:10.1163/1568539x-00003307 · Cited by 57 · cited 2× · see citing works
“Similarly, there are differences across strains in the relative frequency of nape attacks and in the use of defensive tactics that lead to evading contact or promoting close-quarter wrestling (Himmler et al., 2016).”
BackgroundSelf-citationKisko, T. M., Wöhr, M., Pellis, V. C., & Pellis, S. M. (2015) · Current Topics in Behavioral Neurosciences · doi:10.1007/7854_2015_432 · Cited by 56 · cited 1× · see citing works
“As the use of ultrasonic vocalizations (USV) can be critical for communication during play, facilitating role reversals and avoiding escalation to aggression (Kisko et al., 2017; Burke et al., 2020), rats with different styles of RTP may need to modify their use of such calls (Pellis et al.”
BackgroundSelf-citationPlay deprivation without social isolation: Housing controls
Holloway, K. S., & Suter, R. B. (2004) · Developmental Psychobiology · doi:10.1002/dev.10151 · Cited by 52 · cited 1× · see citing works
“Not all partners are equally attractive as play mates (Holloway and Suter, 2004; Pellis et al., 2006).”
BackgroundOpioid modulation of social play reward in juvenile rats
Achterberg, E. J. M., van Swieten, M. M. H., Houwing, D. J., Trezza, V., & Vanderschuren, L. J. M. J. (2019) · Neuropharmacology · doi:10.1016/j.neuropharm.2018.09.007 · Cited by 52 · cited 1× · see citing works
“While opioid systems have been implicated in the rewards derived from engaging in play (Vanderschuren et al., 2016; Achterberg et al., 2019), exactly which neural circuits are associated with what types of playful actions and styles of play remains to be investigated.”
BackgroundSelf-citationThe role of the medial prefrontal cortex in regulating interanimal coordination of movements.
Himmler, B. T., Bell, H. C., Horwood, L., Harker, A., Kolb, B., & Pellis, S. M. (2014) · Behavioral Neuroscience · doi:10.1037/bne0000010 · Cited by 37 · cited 1× · see citing works
“, 2016), and the mPFC has a crucial role in coordinating actions with partners as juveniles and as adults in both playful and non-playful social interactions (Bell et al., 2009; Van Kerkhof et al., 2013; Himmler et al., 2014; Stark and Pellis, 2020).”
BackgroundSelf-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 · cited 1× · see citing works
“These differences have provided valuable tools for using sex and strain differences to identify the roles of specific neural systems and neural networks in regulating particular aspects of play (Siviy, 2020; VanRyzin et al., 2020).”
Measuring play fighting in rats: a multilayered approach
Pellis, S. M., Pellis, V. C., Burke, C. J., Stark, R. A., Ham, J. R., Euston, D. R., & Achterberg, E. J. M. (2022) · Current Protocols · doi:10.1002/cpz1.337 · Cited by 33 · cited 6× · see citing works
“Within a strain, high players not only initiate more nape attacks, but also preferentially use a different suite of the rat-typical defensive tactics than do low players (Pellis et al., 2022).”
BackgroundMethodInfluentialSelf-citationDomestication and the Role of Social Play on the Development of Sociocognitive Skills in Rats
Pellis, S. M., Pellis, V. C., Himmler, B. T., Modlinska, K., Stryjek, R., Kolb, B., & Pisula, W. (2019) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2019.32.00.17 · Cited by 32 · cited 1× · see citing works
“Indeed, identifying that turn taking, as shown by the occurrence of role reversals, is a key feature of RTP that promotes the development of socio-cognitive skills (Pellis et al., 2019; Stark et al., 2021), has been important for engineering therapeutic play contexts that similarly promote the development of those ski…”
BackgroundSelf-citationStark, R., & Pellis, S. M. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1720142 · Cited by 32 · cited 2× · see citing works
“, 2016), and the mPFC has a crucial role in coordinating actions with partners as juveniles and as adults in both playful and non-playful social interactions (Bell et al., 2009; Van Kerkhof et al., 2013; Himmler et al., 2014; Stark and Pellis, 2020).”
BackgroundSelf-citationContext-dependent individual differences in playfulness in male rats
Lampe, J. F., Burman, O., Würbel, H., & Melotti, L. (2017) · Developmental Psychobiology · doi:10.1002/dev.21509 · Cited by 30 · cited 1× · see citing works
“Even within members of the same sex and same strain, not all individuals play to the same degree—some rats consistently play more than others (Lampe et al., 2017; Lesscher et al., 2021).”
BackgroundDifferential effects of amphetamine on the attack and defense components of play fighting in rats
Field, E. F., & Pellis, S. M. (1994) · Physiology & Behavior · doi:10.1016/0031-9384(94)90202-x · Cited by 29 · see citing works
Self-citationReppucci, C. J., Brown, L. A., Chambers, A. Q., & Veenema, A. H. (2020) · Physiology & Behavior · doi:10.1016/j.physbeh.2020.113162 · Cited by 28 · see citing works
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 · cited 1× · see citing works
“As the use of ultrasonic vocalizations (USV) can be critical for communication during play, facilitating role reversals and avoiding escalation to aggression (Kisko et al., 2017; Burke et al., 2020), rats with different styles of RTP may need to modify their use of such calls (Pellis et al.”
BackgroundSelf-citationLesscher, H. M. B., Achterberg, E. J. M., Siviy, S. M., & Vanderschuren, L. J. M. J. (2021) · Psychopharmacology · doi:10.1007/s00213-021-05929-1 · Cited by 23 · cited 1× · see citing works
“Even within members of the same sex and same strain, not all individuals play to the same degree—some rats consistently play more than others (Lampe et al., 2017; Lesscher et al., 2021).”
BackgroundSelf-citationStark, R. A. M., Ramkumar, R., & Pellis, S. M. (2021) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2021.34.5592 · Cited by 20 · cited 1× · see citing works
“Indeed, identifying that turn taking, as shown by the occurrence of role reversals, is a key feature of RTP that promotes the development of socio-cognitive skills (Pellis et al., 2019; Stark et al., 2021), has been important for engineering therapeutic play contexts that similarly promote the development of those ski…”
BackgroundSelf-citationHimmler, B. T., Himmler, S. M., Stryjek, R., Modlińska, K., Pisula, W., & Pellis, S. M. (2015) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2015.28.00.10 · Cited by 18 · see citing works
Self-citationTreatment with low doses of nicotine but not alcohol affects social play reward in rats
Achterberg, E. J. M., & Vanderschuren, L. J. M. J. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1720121 · Cited by 13 · see citing works
Self-citationSiviy, S. M. (2020) · International Journal of Play · doi:10.1080/21594937.2020.1721024 · Cited by 10 · cited 3× · see citing works
“There is strong evidence that the difference in launching nape attacks is linked to differences in mesolimbic dopamine activity (Vanderschuren et al., 2016; Siviy, 2020).”
BackgroundAldis, O. (1975) · Play Fighting · doi:10.1016/b978-0-12-049450-7.50014-2
- Self-citation
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Flagged first, then by citations
Research works printed but not linked: 3
- Pellis, 1997, 31, 193
- Pellis, 2009, The Playful Brain: Venturing to the Limits of Neuroscience
- Pellis, 2016, Play and cognition: the final frontier, 201
Cited by
Sorted by citations
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-citationHam, J., & Pellis, S. (2023) · Animal Behavior and Cognition · doi:10.26451/abc.10.04.02.2023 · Cited by 22 · see citing works
Self-citationAchterberg, E. J. M., Burke, C. J., & Pellis, S. M. (2023) · Behavioural Processes · doi:10.1016/j.beproc.2023.104933 · Cited by 20 · see citing works
Self-citationHam, J. R., Szabo, M., Annor‐Bediako, J., Stark, R. A., Iwaniuk, A. N., & Pellis, S. M. (2024) · Developmental Psychobiology · doi:10.1002/dev.22456 · Cited by 18 · see citing works
Self-citationHome-cage behavior is impacted by stress exposure in rats
Eraslan, E., Castelhano-Carlos, M. J., Amorim, L., Soares‐Cunha, C., Rodrigues, A. J., & Sousa, N. (2023) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2023.1195011 · Cited by 17 · see citing works
“They engage in play behavior, such as chasing and rough-and-tumble, which helps them to learn and practice important social skills (Vanderschuren et al., 1997; Schweinfurth et al., 2017; Pellis et al., 2022).”
BackgroundPellis, S. M., Pellis, V. C., & Ham, J. R. (2024) · Frontiers in Ethology · doi:10.3389/fetho.2024.1362052 · Cited by 15 · see citing works
Self-citationPlay 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
Self-citationAssociations of the immune system in aggression traits and the role of microglia as mediators
Takahashi, A. (2024) · Neuropharmacology · doi:10.1016/j.neuropharm.2024.110021 · Cited by 11 · see citing works
Social 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
“5,14 17 Playactivityisdefinedasthe summedtimeofpouncing,pinning,andboxingdividedbythetotal time of observation.”
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
“Since social play behavior in rats strongly depends on the playfulness of its partner (Pellis et al. 2022; Trezza and Vanderschuren 2008a), both rats in a pair were from the same treatment group and did not differ more than 10 g in body weight.”
Self-citationHam, J. R., & Pellis, S. M. (2025) · iScience · doi:10.1016/j.isci.2025.112562 · Cited by 5 · see citing works
“32 With that said, it is clear from our results that the choice of partners should be provided to rats, and that rats should be tested on more than one day as individual idiosyncrasies in play change day to day.”
Self-citationJuvenile 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-citationComparative analysis of tickling and conspecific play in tame mice and golden hamsters
Dagher, S., DeAngelo, D., Sato, R. Y., Norimoto, H., Koide, T., & Ishiyama, S. (2025) · Behavioural Brain Research · doi:10.1016/j.bbr.2025.115849 · Cited by 5 · see citing works
Križalkovičová, Z., Szabó, P., Kumli, K., Štefanovský, M., Makai, A., & Szentpéteri, J. L. (2024) · Frontiers in Psychology · doi:10.3389/fpsyg.2024.1457515 · Cited by 6 · see citing works
“…childhood, judo may even resemble rough and tumble play among mammals which is a crucial part of their developmental process, with usually the playfight continuing until one of the participants is on their back on the ground (Blomqvist and Stylin, 2021; Palagi et al., 2016; Pellis et al., 2022).”
González‐González, S., Gutiérrez‐Pérez, M., Guzmán‐Ruiz, M. A., Espítia‐Bautista, E., Pavón, R. M., Estrada‐Rodriguez, K. P., Alejandro Díaz-Infante, R., Gándara, C. G. G., Escobar, C., & Guerrero‐Vargas, N. N. (2025) · Frontiers in Physiology · doi:10.3389/fphys.2024.1520160 · Cited by 3 · see citing works
“Social interactions during play behavior are critical for social and cognitive development in adulthood (Pellis et al., 2022).”
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
Self-citationThrough Human Eyes: Owner Insights into the Social Relationships of Pet Rats
Walburn, C., Blackwell, E. J., Mendl, M., Paul, E. S., & Neville, V. (2025) · Animals · doi:10.3390/ani15172579 · Cited by 1 · see citing works
Developmental encoding of natural sounds in the mouse auditory cortex
Zucca, S., Rosa, C. L., Fellin, T., Peretto, P., & Bovetti, S. (2024) · Cerebral Cortex · doi:10.1093/cercor/bhae438 · 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
Self-citationHam, 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-citationKuehnel, I. M., González‐Irarrázabal, C. E., Pardo, R., Julio‐Pieper, M., & Bravo, J. A. (2025) · Journal of Neurochemistry · doi:10.1111/jnc.70224 · Cited by 1 · see citing works
“Both behaviours are known to be mediated by dopaminergic receptors and connections in the prefrontal cortex (PFC) (Pellis et al. 2022; Lynch et al. 2023; Chen et al. 2024).”
Liu, T. H., Sroka, A. W., Zhang, J. J., Jacobson, K. K., Gevirtz, R., Kellman, J., & Radwan, K. (2025) · Developmental Psychobiology · doi:10.1002/dev.70100 · Cited by 1 · 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
“Social play is typically defined as active, voluntary interaction with at least one other peer and is highly rewarding in nature (Achterberg & Vanderschuren, 2023; Melotti et al., 2014; Pellis et al., 2022b).”
Unravelling the Role of the Dopamine D1 Receptor in Anhedonia, Asociality, and Avolition
Witt, K. (2023) · doi:10.26686/wgtn.24646128 · 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
Orsucci, I. C., Becker, K. D., Ham, J. R., Lee, J., Bowden, S., & Veenema, A. H. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.11.14.623692 · 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-citationEconomic and Social Implications of Nanochitosan
Oranusi, S. U., Mameh, E. O., Oyegbade, S. A., et al. (2024) · Springer eBooks · doi:10.1007/978-3-031-52261-1_12
Ham, J. R., & McDonald, R. J. (2026) · iScience · doi:10.1016/j.isci.2026.115459
“11 Kyoto rats also engage in less play—a naturally occurring behavior 12 —than Wistar rats, at all ages, demonstrating anhedonia through emotionally and socially relevant behavior.”
Self-citation
- Authors
- Pellis, S. M., Pellis, V. C., Ham, J. R., & Achterberg, E. J. M.
- Affiliations
- University of Lethbridge (CA)
- Utrecht University (NL)
- Corresponding author
- Pellis, Sergio M.
- Venue
- Frontiers in Behavioral Neuroscience
- Publisher
- Frontiers Media SA
- Published
- 2022-10-18
- Volume
- 16
- Pages
- 1033999-1033999
- ISSN
- 1662-5153 (ISSN-L)
- PMC
- PMC9623181
- OpenAlex
- W4306679605
- Cited by
- 29
- References
- 51
- Citations by year
- 2026: 4 · 2025: 8 · 2024: 13 · 2023: 4
- FWCI
- 5.60
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Frontiers in Behavioral Neuroscience (publishedVersion, cc-by)
- PubMed (publishedVersion)
- PubMed Central (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Utrecht University Repository (Utrecht University) (submittedVersion, other-oa)
- Funders
- Natural Sciences and Engineering Research Council of Canada
- Topics
- Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Social Sciences › Psychology › Developmental and Educational Psychology › Child and Animal Learning Development
- Keywords
- rough-and-tumble play, social play, rats, social brain, social dynamics, competition and cooperation, reciprocity, turn-taking, aggression, individual differences, social skills, social cognition, juvenile development, social communication, evolution of social behavior, social behavior
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