Cited by 83Neuroendocrine regulation and behaviorPrimate Behavior and EcologyMemory and Neural Mechanisms

Peering into the Dynamics of Social Interactions: Measuring Play Fighting in Rats

Open access
Himmler, Brett T., Pellis, Vivien C., Pellis, Sergio M.·2013·Journal of Visualized Experiments

Abstract

Play fighting in the rat involves attack and defense of the nape of the neck, which if contacted, is gently nuzzled with the snout. Because the movements of one animal are countered by the actions of its partner, play fighting is a complex, dynamic interaction. This dynamic complexity raises methodological problems about what to score for experimental studies. We present a scoring schema that is sensitive to the correlated nature of the actions performed. The frequency of play fighting can be measured by counting the number of playful nape attacks occurring per unit time. However, playful defense, as it can only occur in response to attack, is necessarily a contingent measure that is best measured as a percentage (#attacks defended/total # attacks X 100%). How a particular attack is defended against can involve one of several tactics, and these are contingent on defense having taken place; consequently, the type of defense is also best expressed contingently as a percentage. Two experiments illustrate how these measurements can be used to detect the effect of brain damage on play fighting even when there is no effect on overall playfulness. That is, the schema presented here is designed to detect and evaluate changes in the content of play following an experimental treatment.

AI snapshot

Written from the paper’s full text.

TL;DR

This paper presents a contingent, dynamic scoring schema for rat play fighting that measures initiating nape attacks, probability of defense, and defensive tactics as percentages rather than absolute outcomes. Two illustrative experiments show the schema can detect brain-damage effects on play content even when overall playfulness is unchanged.

Key findings

  • Play fighting frequency is measured by playful nape attacks per unit time; defense is contingent and best expressed as percentage of attacks defended (# attacks defended/total attacks × 100%), and defensive tactic type as a percentage of defenses.
  • In decorticate vs intact juvenile male rats, pinning was reduced by about 50% (40 vs 18), but playful attacks did not differ significantly (53 vs 49, ns), probability of defense did not differ (.91 vs .93, ns), and evasion did not differ (20% vs 25%, ns); however, complete rotation was lower in decorticates (60% vs 20%, p < 0.05) and partial rotation was higher (20% vs 60%, p < 0.05).
  • In subordinate adult males with orbital frontal cortex (OFC) lesions, there was no significant difference before or after surgery in frequency of playful attacks, probability of defense, or use of facing defense, but OFC-damaged rats showed virtually no asymmetry in defensive tactics toward the dominant male vs the female, whereas controls continued to show a high degree of asymmetry.
  • With puberty, male rats show a relative decline in complete rotation and an increase in partial rotation; subordinate males use partial rotation more with another subordinate male or a female, but complete rotation more with the dominant colony male.

Methods

  • Animals were weaned between postnatal days 21–23, housed in groups of 2–4, habituated for three consecutive 30-min sessions, isolated 24 hr before testing, and tested as pairs for 10 min in a clear Plexiglas box (50 cm × 50 cm × 50 cm) with corncob bedding, filmed in the dark with a night-shot camcorder at a 45° downward angle.
  • Scoring used a sequential analysis: initiating playful attacks were counted when the snout contacted or came within 1–2 cm of the partner’s nape; defense was scored as the proportion of attacks defended; defensive tactics were categorized within the first 2–3 frames of the defensive action; tactic types were expressed as percentages of defenses.
  • Representative experiments compared neonatally decorticated rats with sham-treated controls, and tested subordinate adult males in triads (two males and one female with fallopian tubes tied) before and after OFC lesions or sham surgery, using an asymmetry index based on the percentage difference in complete rotation when playing with the dominant male vs the female.

Limitations

  • The paper states that the framework may be inadequate if used alone; in a wild-type vs Long-Evans comparison, defensive actions began at a bigger inter-animal distance, so measuring nape attacks alone underestimated play, and scoring outcome measures such as chasing and boxing was important for a more balanced picture.
  • The paper states that counterattacks during extended wrestles are more difficult to score, especially regarding the partner’s defensive reaction, and need not be scored unless there is good reason.
  • The paper notes that Table 1 shows only mean values for illustrative purposes and that researchers should establish baseline values for their own rat strain; the text also makes evident that the representative results are illustrative rather than full primary analyses with variance reported.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.3791/4288

Cites

42 of 42 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 51 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    As adults, decorticates seemed to fail to modify their play when confronted with the dominant member of the group.

    “As adults, decorticates seemed to fail to modify their play when confronted with the dominant member of the group.”
    Quote in the paperSubstantial claim
  • Findingcitation

    Measuring nape contacts or nape attacks, there is no difference in the frequency of initiating play between intact and decorticate rats.

    “However, measuring nape contacts 10 , or in our conception, nape attacks 26 , there is no difference in the frequency of initiating play between intact and decorticate rats.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [10]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [26]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Findingnone given

    The ability of the OFC rats to play differently with the two types of partners was compromised.

    “However, the ability of the OFC rats to play differently with the two types of partners was compromised.”
    Quote in the paperSubstantial claim
  • Findingcitation

    When the effects of decortication were measured using pinning, there was about a 50% reduction in this measure in the juvenile period.

    “When the effects of decortication were measured using pinning, there was about a 50% reduction in this measure in the juvenile period 10,26 .”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [10]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [26]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Findingcitation

    Damage limited to the motor cortex was sufficient to replicate the effect on juvenile rats.

    “Subsequent analyses showed that damage limited to the motor cortex was sufficient to replicate this effect on juvenile rats 6 .”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [6]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims. 18
  • Classified as probably not a substantial claim. 18
  • May combine two claims.

    Findingown data

    When a defender executes a complete rotation, the movement is quick and fluid; on a standard 30 fps recording, a rotation to fully supine takes 2-3 frames and the movement is continuous.

    “When a defender executes a complete rotation, the movement is quick and fluid -if videotaped on a standard 30 fps recording, a rotation to fully supine takes 2-3 frames and the movement is continuous (Video).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    A 'forced' supine from partial rotation occurs in a staccato manner, with incremental rotation of the body, and can take substantially more than 3 video frames.

    “Such a 'forced' supine occurs in a staccato manner, with incremental rotation of the body, and can take substantially more than 3 video frames (Video showing a partial rotation leading to the defender adopting a supine position).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingreasoning

    A partial rotation can also end in a fully supine position, but this typically involves the concerted actions of the attacker.

    “A partial rotation can also end in a fully supine position, but this typically involves the concerted actions of the attacker.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    Following the lesion and re-establishment of social relationships, the controls continued to show a high degree of asymmetry, whereas the OFC damaged rats showed virtually no difference in their pattern of defense when attacked by the dominant male or by the female.

    “Following the lesion and the re-establishment of social relationships, the controls continued to show a high degree of asymmetry, whereas the OFC damaged rats showed virtually no difference in their pattern of defense when attacked by the dominant male or by the female (Figure 4).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    There was no significant difference either before or after surgery in the frequency of playful attacks, the probability of defense or in the use of facing defense between the subordinates in the experimental and control conditions.

    “There was no significant difference either before or after surgery in the frequency of playful attacks, the probability of defense or in the use of facing defense between the subordinates in the experimental and control conditions.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingnone given

    The control rats in Table 1 show typical values for these measures when using Long-Evans hooded rats.

    “The column in Table 1 for the control rats shows the typical values for these measures when using Long-Evans hooded rats.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    The tactic attempted by the subject can be judged by examining the movements of the defender within the first 2-3 frames of the commencement of a defensive action.

    “This judgment can be made by examining the movements of the defender within the first 2-3 frames of the commencement of a defensive action (Video highlighting the first 2-3 frames, frame-by-frame).”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    The relative defensiveness of the recipient of attacks can be measured as the proportion of attacks defended.

    “Third, given that defense is contingent on attack, the relative defensiveness of the recipient of attacks can be measured as the proportion of attacks defended.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    Brief periods of social isolation increase the motivation to engage in social play while not having a similar effect on other social behavior such as social investigation.

    “Brief periods of social isolation increases the motivation to engage in social play while not having a similar effect on other social behavior such as social investigation 9 .”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [9]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    Play fighting in the rat involves attack and defense of the nape of the neck, which if contacted is gently nuzzled with the snout.

    “Rats, for example, attack and defend the nape of the neck 15 , which if contacted is gently nuzzled with the snout 11,32 .”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [15]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [11]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [32]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Study design and conductcitation

    By 15 min, animals appear fatigued by vigorous play; trials lasting 10 min are a good compromise between economy of protocol and ability to detect experimental effects.

    “Even so, by 15 min, the animals appear fatigued by the vigorous play and consequently, trials lasting 10 min are a good compromise between economy of protocol and the ability to detect experimental effects 15,16,27 .”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [15]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [16]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [27]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    The scoring schema is designed to detect and evaluate changes in the content of play following an experimental treatment.

    “That is, the schema presented here is designed to detect and evaluate changes in the content of play following an experimental treatment.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim.

    Study design and conductnone given

    Playful attacks can be measured when the snout of one rat either contacts or comes within 1-2 cm of its partner's nape.

    “Playful attacks can be measured when the snout of one rat either contacts or comes within 1-2 cm of its partner's nape (Video showing a playful attack compared to non-playful movements near partner rat).”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S2 (checkable).

    Study design and conductreasoning

    The type of defense used against a playful attack is best expressed contingently as a percentage because it is contingent on defense having taken place.

    “How a particular attack is defended against can involve one of several tactics, and these are contingent on defense having taken place; consequently, the type of defense is also best expressed contingently as a percentage.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    Playful defense, because it can only occur in response to attack, is best measured as a percentage (#attacks defended/total # attacks X 100%).

    “However, playful defense, as it can only occur in response to attack, is necessarily a contingent measure that is best measured as a percentage (#attacks defended/total # attacks X 100%).”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Interpretationreasoning

    A low frequency of complete rotations could arise because of low rates of attack or because subjects used the complete rotation infrequently even if attacked just as frequently as controls.

    “For example, a low frequency of complete rotations could arise because of low rates of attack or because the subjects in that particular experimental condition used the complete rotation infrequently even if they had been attacked just as frequently as the animals in the control condition.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Study design and conductreasoning

    Trials of 5 min can detect isolation-induced changes but are too short to detect subtle differences such as those between the sexes due to a ceiling effect; the greater propensity for males to play mostly emerges after the first 5 min.

    “While trials of 5 min can detect such isolation-induced changes, they are too short to detect subtle differences such as those between the sexes due to a ceiling effect -in this example, the greater propensity for males to play mostly emerges after the first 5 min.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Study design and conductreasoning

    When scoring play, there is a sequential analysis: how many attacks are launched by the subjects (number/unit time), how many attacks are defended against (probability of defense), and when defending, the likelihood of using different tactics (%tactics).

    “Thus, when scoring play, there is a sequential analysis: how many attacks are launched by the subjects (number/unit time), how many attacks are defended against (i.e., probability of defense), and when defending, what is the likelihood of using the different tactics of defense (%tactics).”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Study design and conductreasoning

    In some situations, the current framework is likely to be inadequate if used alone.

    “In some situations, the current framework is likely to be inadequate if used alone.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data

    Even though the OFC damaged subordinates played just as much as the controls, they were no longer able to discriminate between their pair mates.

    “That is, even though the OFC damaged subordinates played just as much as the controls, they were no longer able to discriminate between their pair mates, supporting the hypothesis that partner-related modulation of play and other social behaviors is mediated by the OFC”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The asymmetry index (difference in probability of complete rotation with the dominant male versus the female) reflects the subordinate male's ability to discriminate between its pair mates.
  • Probably not a substantial claim: may not pass S2 (checkable).

    Study design and conductreasoning

    A scoring scheme that is sensitive to those dynamics and the contingent nature of the actions performed is needed.

    “As a consequence, a scoring scheme that is sensitive to those dynamics and the contingent nature of the actions performed is needed.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Premisecitation

    The orbital frontal cortex (OFC) was tested as a likely candidate structure in such modulation, given its role in regulating social behavior generally.

    “The orbital frontal cortex (OFC) was tested as a likely candidate structure in such modulation, given its role in regulating social behavior generally 7 .”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [7]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The distance at which rats respond to the approach of an attacker can influence the type and the success of the tactics executed.

    “For example, the distance at which rats respond to the approach of an attacker can influence the type and the success of the tactics executed 25 .”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [25]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Complete removal of the cortex can lead to adult rats that can function relatively normally in laboratory environments.

    “Complete removal of the cortex can lead to adult rats that can function relatively normally in laboratory environments 42 , however, given the presumed role of large brains in promoting play in mammals 3 , we wanted to test whether play is reduced in rats without a cortex.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [42]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S4 (based).

    Premisenone given

    Play fighting is a complex, dynamic interaction in which the participants continually influence each other's behavior.

    “Play fighting is a complex, dynamic interaction in which the participants continually influence each other's behavior.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    In males, with the onset of puberty, there is a relative decline in use of the complete rotation tactic and an increase in use of the partial rotation tactic, with other tactics remaining at about the same frequency at all ages.

    “In males, with the onset of puberty, there is a relative decline in their use of the complete rotation tactic (Different Video of Full Rotation as shown in above parts) and an increase in their use of the partial rotation tactic (Different Video of Partial Rotation as show in above parts), with the other tactics remaining at about the same frequency at all ages 16 .”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [16]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductcitation

    In play fighting, this can be when the attacker is approaching the neck from a perpendicular angle and the recipient is at least one body length away from a wall, thus removing confounds that could arise from opportunity rather than capacity.

    “In the case of play fighting, this can be when the attacker is approaching the neck from a perpendicular angle and the recipient is at least one body length away from a wall, thus removing the confounds that could arise from opportunity rather than capacity 22 .”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [22]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    Decorticates infrequently adopted the complete rotation defensive strategy, which is the most common route by which pins arise.

    “Thus, decorticates infrequently adopted a defensive strategy -complete rotation -that is the most common route by which pins arise.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing). May combine two claims.

    Study design and conductreasoning

    Although the currently proposed framework may have limitations when used as the sole method of studying play, modifications can be introduced in a manner that meets specific research goals and does so while retaining the guiding principle of the contingent and dynamic nature of such interactions.

    “Therefore, although the currently proposed framework may have limitations when used as the sole method of studying play, modifications can be introduced in a manner that meets specific research goals and does so while retaining the guiding principle of the contingent and dynamic nature of such interactions 13 .”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [13]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    A recent analysis of juvenile play in wild-type rats compared to Long-Evans rats showed that, because defensive actions began at a bigger inter-animal distance than in the wild rats, measuring nape attacks alone underestimated the measure of play.

    “For example, a recent analysis of juvenile play in wild-type rats compared to Long-Evans rats showed that, because defensive actions began at a bigger inter-animal distance than in the wild rats, measuring nape attacks alone underestimated the measure of play.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Himmler et al., work in progress

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Study design and conductreasoning

    The currently proposed framework for scoring the actions performed during play can be thought of as a skeleton, upon which additional measurements can be incorporated as needed.

    “Thus, the currently proposed framework for scoring the actions performed during play can be thought of as a skeleton, upon which additional measurements can be incorporated as needed.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    The pubertal decline in play fighting is because rats decrease the frequency of launching nape attacks, but maintain the same high probability of defense.

    “This is because they decrease the frequency of their launching of nape attacks, but maintain the same, high probability of defense 17 .”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [17]

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

32 of 32
  • The paper relies on this without support.

    The asymmetry index (difference in probability of complete rotation with the dominant male versus the female) reflects the subordinate male's ability to discriminate between its pair mates.

    “This yields an asymmetry index, so that the closer the score is to 100%, the more asymmetrical, and the closer to 0%, the more symmetrical.”
    “That is, even though the OFC damaged subordinates played just as much as the controls, they were no longer able to discriminate between their pair mates, supporting the hypothesis that partner-related modulation of play and other social behaviors is mediated by the OFC”
1 of 1

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?

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • Decorticate defense percentages exceed 100%

    Minor

    The three listed defense types total 105% for decorticate rats (25% + 20% + 60%). The paper describes these as proportions of defenses classified by tactic, so the listed categories cannot exceed 100% even before including any other tactic.

1 of 1

Cites

Flagged first, then by citations

Research works printed but not linked: 3

  1. Aldis, O. In: Play Fighting. Academic Press, (1975).
  2. Pellis, S.M. & Pellis, V.C. The Playful Brain. In: Venturing to the Limits of Neuroscience. Oneworld Press, Oxford, (2009).
  3. Whishaw, I.Q. In: The decorticate rat. Kolb, B. & Tees, R.C., Eds., Cerebral cortex of the rat, MIT Press, 239-268, (1990).

Cited by

Sorted by citations

Authors
Himmler, B. T., Pellis, V. C., & Pellis, S. M.
Affiliations
University of Lethbridge (CA)
Venue
Journal of Visualized Experiments
Published
2013-01-18
Issue
71
Pages
e4288-e4288
ISSN
1940-087X (ISSN-L)
OpenAlex
W2091955003
Cited by
81
References
39
Citations by year
2026: 1 · 2025: 5 · 2024: 6 · 2023: 4 · 2022: 7 · 2021: 12 · 2020: 4 · 2019: 5 · 2018: 9 · 2017: 5 · 2016: 6 · 2015: 8 · 2014: 7 · 2013: 2
FWCI
10.99
Type
journal article
Language
en
Open access
Locations
  • Journal of Visualized Experiments (publishedVersion)
  • PubMed (publishedVersion)
  • Europe PMC (PubMed Central) (submittedVersion)
  • PubMed Central (submittedVersion)
Funders
  • Harry Frank Guggenheim Foundation
  • John Simon Guggenheim Memorial Foundation
Topics
  • Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
  • Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
Keywords
play fighting, rats, social interaction, defense tactics, brain damage, play behavior, behavioral assessment
MeSH terms
Agonistic Behavior, Animals, Behavior, Animal, Female, Male, Play and Playthings, Social Behavior, Rats

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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.

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Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-12.

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