Who’s laughing? Play, tickling and ultrasonic vocalizations in rats.
Open accessAbstract
Social play in rats is a highly rewarding, energetic form of social interaction and important for development of the brain and social skills. The 50 kHz ultrasonic vocalizations (USV) emitted during social play are thought to be an expression of a positive affective state (laughter), which in some situations may also function as communication signals. Heterospecific play, ‘tickling’ by an experimenter, is thought to simulate conspecific play, and has been used to improve welfare and to study the neurobiology of positive affect. Given that tickling evokes substantial amounts of USV, we investigated whether heterospecific play is simulating conspecific play by comparing USV-behaviour associations in both contexts. If the 50 kHz calls are merely an expression of ‘laughter’ then the pattern and type of emission in both contexts should be similar. By contrast, as playing with a conspecific involves a two-way exchange of signalling, the additional demands on communication should lead to a different pattern of calling. While calling was prevalent in both types of play, how the different types of 50 kHz calls are used in the two contexts differed markedly. The findings suggest that while conspecific and heterospecific play are positive experiences, tickling is not the equivalent of conspecific play. This article is part of the theme issue ‘Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience’.
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Burke et al. (2022) suggested that for male rats being pinned during tickling may be important for production of USVs in comparison to social play where de-vocalisation studies show that it is typically the rat doing the pinning that emits these calls.
“Burke et al. (2022) suggested that for male rats being pinned during tickling may be important for production of USVs in comparison to social play where de-vocalisation studies show that it is typically the rat doing the pinning that emits these calls ( Burke et al., 2018).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al., 2022
Split into 2 atoms — 1 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 0 unverifiable, 1 not checked. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredBurke et al. (2022) propose that the lack of correlation between trills and specific tickling behaviour indicates that trills are produced relatively indiscriminately during tickling, potentially due to the lack of a conspecific play partner.
“Burke et al. (2022) propose that the lack of correlation between trills and specific tickling behaviour indicates that trills are produced relatively indiscriminately during tickling, potentially due to the lack of a conspecific play partner.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al. (2022)
Split into 2 atoms — 1 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 0 unverifiable, 1 not checked. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredIn male rats the strongest associations between USVs and behaviour in social play are between approaching, following and nape contact, whereas with tickling the strongest association is with pinning.
“In male rats the strongest associations between USVs and behaviour in social play are between approaching, following and nape contact whereas with tickling the strongest association is with pinning ( Burke et al., 2022; they applied the standard tickling protocol).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al., 2022
Burke et al. (2022) report that in social play, the strongest associations between behaviour and USVs involve frequency modulated (trill) calls, whereas in tickling the strongest associations involve flat calls.
“Burke et al. (2022) report that in social play, the strongest associations between behaviour and USVs involve frequency modulated (trill) calls, whereas in tickling the strongest associations involve flat calls.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al. (2022)
Another distinctive difference between social play and tickling in the Burke et al. (2022) study were the correlations between behaviours and sub-types of USVs.
“Another distinctive difference between social play and tickling in the Burke et al., (2022) study were the correlations between behaviours and sub-types of USVs.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al., (2022)
The patterns and correlations between USVs and specific behaviours are known to vary across contexts.
“For example, the patterns and correlations between USVs and specific behaviours are known to vary across contexts ( Burke et al., 2022; Melotti et al., 2014).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Burke et al., 2022
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- Authors
- Burke, C. J., Pellis, S. M., & Achterberg, E. J. M.
- Affiliations
- University of Lethbridge (CA)
- Utrecht University (NL)
- Corresponding author
- Burke, C. J.
- Venue
- Philosophical Transactions of the Royal Society B
- Publisher
- The Royal Society
- Published
- 2022-09-21
- Volume
- 377
- Issue
- 1863
- Pages
- 20210184-20210184
- ISSN
- 0962-8436 (ISSN-L)
- 1471-2970
- PMC
- PMC9489288
- OpenAlex
- W4296781468
- Cited by
- 23
- References
- 57
- Citations by year
- 2026: 2 · 2025: 6 · 2024: 6 · 2023: 6 · 2022: 3
- FWCI
- 4.82
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Licence
- Locations
- Philosophical Transactions of the Royal Society B Biological Sciences (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Utrecht University Repository (Utrecht University) (submittedVersion, other-oa)
- Utrecht University Repository (Utrecht University) (submittedVersion)
- Funders
- Natural Science and Engineering Research Council of Canada (2018-03706)
- Natural Sciences and Engineering Research Council of Canada (PGSD3)
- Nederlandse Organisatie voor Wetenschappelijk Onderzoek (016.Veni.181.039)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Social Sciences › Psychology › Social Psychology › Humor Studies and Applications
- Keywords
- social play, tickling, ultrasonic vocalizations, 50-kHz calls, positive affect, animal welfare, communication signals, rats
- MeSH terms
- Animals, Emotions, Reward, Ultrasonics, Vocalization, Animal, Rats
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