Rough-and-Tumble Play and the Development of the Social Brain

“…involved in play and structured games that involve learning and group cooperation may adapt differently than children who are involved in individual physical activities that are performed in relative isolation (e.g., treadmill running or cycling on a stationary ergometer) (Pellis and Pellis 2007).”

“…mammals, energizing them to develop and refine their physical, emotional, and social competencies in a safe context Frontiers in Human Neuroscience | www.frontiersin.org 10 March 2017 | Volume 11 | Article 145 (Panksepp, 1998; Pellis and Pellis, 2007; Trezza et al., 2010; Panksepp and Biven, 2012).”

“…the social brain and social behavior, and many studies involving humans and rodents have illustrated the negative consequences associated with limited or abnormal play exposure in adolescence (Bell et al., 2010; Darwish et al., 2001; Hol et al., 1999; Pellis and Pellis, 2007; Pellis et al., 2010b).”

“* Andreas Alexiou A.Alexiou@uva.nl Michaéla C. Schippers mschippers@rsm.nl 1 Amsterdam Business School, University of Amsterdam, Amsterdam, the Netherlands 2 Rotterdam School of Management, Erasmus University, Rotterdam, the Netherlands studies in neuroscience (e.g. Pellis and Pellis 2007).”

“Laboratory rats display spontaneous play fighting, which seems to have important role in the development of social skills—like coping with aggressive individuals.(105,106) This type of social interaction— contrary to serious fighting—typically does not involve any biting nor threat postures, but instead is based on tr…”

“Unstructured play is therefore thought to be a crucial context for the development of independence and emotional/behavioural self-regulation (Pellis & Pellis, 2007).”

“They also do not copulate normally, which is attributed to a failure to coordinate their bodies with others’ bodies [44].”

“One of these “fertilized” brain regions, the orbitofrontal cortex, is commonly associated with the regulation of social behavior and is important for RTP competence in rats (Pellis and Pellis 2007).”

“Humor appreciation facilitates rough-and-tumble play, the most common source of laughter in nonhuman primates, which promotes social development later in life (Pellis & Pellis, 2007).”

“…studies in rodents manipulating these variables show that in at least some cases , interlitter variation in size , sex ratios and play - fighting behaviours are not only correlated with , but actually cause , differences in adult behaviours ( Laviola & Terranova 1998 ; Pellis & Pellis 2007 ) .”

“For example, play may help the young of a species to develop important social skills and knowledge (Bekoff & Byers, 1981; Connoly & Doyle, 1984; Pellis & Pellis, 2007; Piaget, 1962; Vygotsky, 1978).”

“(Bekoff & Byers, 1998; Pellis & Pellis, 2007).”

“play during childhood (Hart, 2002; Elkind, 2007; Pellis and Pellis, 2007).”

“Rough and tumble play (R&T), is characterized by fighting elements (performed in non-serious way), which involve more than one player [12–16].”

“In particular, rough and tumble play could exert a critical effect on shaping young minds as in learning adaptive limits for interacting socially (Panksepp, 2007; Pellis & Pellis, 2007).”

“In addition, subcortical circuits may mediate the execution of the appropriate locomotor acts and the integration of sensory stimuli, as well as encode the emotional and motivational properties of social play (Pellis and Pellis 2007; Siviy and Panksepp 2011; Vanderschuren et al. 1997).”

“The specific neural activity observed indicates that connections are being made within areas of the brain that deal with emotion and sociability (Gordon, Burke, Akil, Watson, & Panksepp, 2003; Pellis & Pellis, 2007).”

“Deprivation from play behavior during this critical period leads to social incompetence later in life (Pellis and Pellis 2007).”

“…role of play in the above outcomes comes from experimental manipulations: immature rats deprived of play partners (but not other social partners) do not develop normal neural circuitry underlying socio-cognitive and motor skills (Pellis & Pellis, 2007; Bell et al., 2010; Schneider et al., 2016).”

“The increase in motivated behaviors, such as mating and aggressive behaviors, and the decrease in social behaviors, such as play, are a few examples (Spear, 2000; Romeo et al., 2002; Pellis and Pellis, 2007).”

“Social play during adolescence, with play fighting being the most commonly expressed form, is crucial for the development of adult social competence (Einon and Morgan, 1977; van den Berg et al., 1999; Pellis and Pellis, 2007).”

“[46,49,60,66] on rough and tumble play in rats adds compelling new dimensions to our understanding of this pervasive form of play.”

“2017; lemurs, Pellis and Pellis 2007; Palagi et al.”

“This behavior might even be considered to be crucial for the development of social and other competencies (Pellis & Pellis, 2007).”

“Indeed, research with animal models has suggested that play is modulated by the brain’s reward system (Manduca et al., 2016) and promotes brain and sensorimotor development (Pellis & Pellis, 2007).”

“Peer play is an important early experience that is believed to refine neural networks of the frontal cortex (Pellis and Pellis, 2007;Pellis et al., 2010).”

“activities, explore their social world and make their own decisions [24,25].”

“consensus that removing the opportunity to play can have a number of consequences on later behavior and social functioning [10-13].”

“Given these findings, the availability of social partners, and particularly exposure to older and more experienced peers, has the potential to affect the structure, amount, and behavioral diversity of play (Bender, 2012; Kuczaj et al., 2006; Mackey et al., 2014; Pellis & Pellis, 2007).”

“…social play experiences during early development are necessary for the establishment of competent social skills and the normal expression of social behavior in adulthood (Einon et al. 1978; Jones et al. 1991; Da Silva et al. 1996; Hol et al. 1999; von Frijtag et al. 2002; Pellis and Pellis 2007).”

“…in rodents have been shown to improve the integration of the pre-frontal cortex for learning complex behaviours like self-regulation and planning (Bell, Pellis, & Kolb, 2010; Pellis & Pellis, 2007; Pellis, Pellis, & Himmler, 2014), and enhance neural plasticity (Himmler, Pellis, & Kolb, 2013).”

“A major difference, however, between human play and that of non-human mammals is the greater role for human parents in playful interactions with juveniles (Pellis & Pellis, 2007).”

“Juvenile play is a highly vigorous and dynamic form of social interaction (Pellis and Pellis, 2007; Trezza et al., 2010) and amphetamine may therefore reduce juvenile play behavior secondary to its effects on hyperactivity or by rendering animals more impulsive (Barendsee and Vanderschuren, 2012).”

“Thus, the social behavior repertoire differs between pubescent and adult rats (Panksepp and Beatty, 1980; Pellis and Pellis, 2007; Graham and Burghardt, 2010).”

“…the nature and function of social play in myriad species (e.g., Bekoff, 1995; Connolly & Doyle, 1984; Durand & Schank, 2015; Pelligrini, 1992; Pellis & Pellis, 2007), the goal of this paper is to demonstrate that, at least for humans, social play is not limited to instances in which multiple…”

“For example, studies indicate that fear of novel situations is a hindrance for optimal performance, but that play experiences in childhood might dampen such fear, mainly by improving self-regulation (Pellis & Pellis, 2007; Riksen-Walraven & van Aken, 1997).”

“For example, all mammalian species engage in early behaviors, such as certain patterns of play, which are designed in part to develop inhibitory control, and have been shown to affect prefrontal development (Panksepp, Burgdorf, Turner, and Gordon, 2003; Pellis and Pellis, 2007).”

“Similarly, it is well established that engaging in play with peers is important for the development and practice of social and communication skills (Howes et al. 1994, Pellis and Pellis 2007).”

“It has been observed that pair-housed juvenile rats spend approximately 60 min (2 × 30 min) daily in playful roughhousing (Panksepp, 1993; Panksepp et al., 1984; Pellis and Pellis, 2007), and that, when occurring following a stressful situation, social play and other affiliative behaviours are associated with a reduct…”

“Second, research indicates that fear of novel situations is a negative factor for learning and functioning in adolescence and adulthood, and that play experiences in childhood might dampen such fear (Pellegrini et al., 2007; Pellis and Pellis, 2007).”

“Juvenile play is believed to provide an array of benefit to the participants, including development of the social brain (Pellis et al., 2010; Pellis and Pellis, 2007).”

“This form of play likely results in improved social competencies and later social-competitive advantage (Graham and Burghardt, 2010), as well as establishing dominance relationships before play merges into potentially harmful fighting (Pellis and Pellis, 2007).”

“It has also been concluded that children who have a diverse and rich exposure to various forms of play are more likely to be able to navigate unpredictable and varied social situations when they are adults (Pellis and Pellis, 2007; Brown, 2009).”

“Play is a natural way to enhance self-regulation, solve problems and express oneself; it helps children build a set of mental representations of the world (Bodrova and Leong 2005; Pellis and Pellis 2007).”

“One common element of recent descriptions and definitions of rough-and-tumble play is the inclusion of a ‘play face’ where participants are smiling and laughing (Reed & Brown, 2000). This play face is an important characteristic as it distinguishes rough-and-tumble play from aggression. According to Reed and Brown (20…”

“As the most prevalent and ancient type of social play in mammals (Panksepp, 1998; Pellis & Pellis, 2007), RTP has been argued to have clear evolutionary roots and functions, such as the safe practice of fighting skills and the establishment of social dominance (Boulton & Smith, 1995; Pellegrini &…”

“…of motor and brain functions (e.g. Bekoff & Byers, 1998; Bergh€anel, Schülke, & Ostner, 2015; Byers & Walker, 1995; Montgomery, 2014; Pellis & Pellis, 2007), social play might also be vital for preparing the immature for unpredictable outcomes with unfamiliar conspecifics later in life…”

“Effect of similarity or dissimilarity in playfulness on social play development Play in rats peaks between 30 and 40 days of age and starts to decline during the 8 to 9 week [56, 8, 44].”

“Data has shown that increased social play positively correlates with improved socio-cognitive functioning in rodents (Pellis and Pellis, 2007).”

“Although the true function of play is 94 still unclear from an evolutionary perspective, play is likely to provide an animal with psychological 95 improving skills such as social interaction (Pellis and Pellis, 2007).”

“This contrasts with rat play, which is much more than just pinning, and includes scampering, chasing, sparring, pouncing, darting, dorsal grooming, and wrestling (10, 14, 25, 26).”

“Social play deprivation in rats is also associated with difficulty coordinating movements with a partner in adult social interactions and a tendency to exhibit hyperdefensiveness in response to an approaching rat (Pellis & Pellis, 2007; Pellis, Field, & Whishaw, 1999).”

“In rats, such rough-and-tumble play has been related to the development of social and emotional competences (Pellis & Pellis, 2007).”

“Even before puberty, interaction with peers is fundamental for the proper functioning of mental processes (Pellis and Pellis, 2007; Vicedo, 2010), as considered above.”

“…largely of the order Rodentia, including ground squirrels, hamsters, and prairie dogs (Nunes et al., 2004; Cooper et al., 2023) and especially rats (Pellis and Pellis, 2007), with the occasional reference to some marine mammals, of which dolphins are probably the most playful (Kuczaj and…”

“…including superhero play (Bauer & Dettore, 1997), ‘bad guy’ play (Logue & Detour, 2011), active pretend play (Logue & Harvey, 2010), play fighting (Pellis & Pellis, 2007), physically active and imaginative play (Parsons & Howe, 2006), and rough-andtumble play (Jarvis, 2007; Pellegrini, 1987;…”

“The drive for play is strong, as evidenced by the increased intensity that humans and animals exhibit when they play after a period of restriction—almost as if making up for lost time (L. A. Barnett, 1984; Pellegrini & Smith, 1998; Pellis & Pellis, 2007).”