Sex differences in 50 kHz call subtypes emitted during tickling-induced playful behaviour in rats.
Open accessAbstract
‘Tickling’ induces positive affective states in laboratory rats as evidenced by the production of 50-kHz ultrasonic vocalisations (USVs), although this has mostly been investigated in males. Juvenile rats emit distinctive 50-kHz USV subtypes. Frequency-modulated (FM) 50-kHz USVs are thought to be associated with positive affect and flat 50-kHz USVs with social communication. FM and flat USVs are produced by both sexes during tickling, but it is unclear whether these calls are produced in relation to particular play-related behaviours, and whether USV subtypes are used in a sexually dimorphic manner during tickling. We tested the hypotheses that FM USVs are associated with tickle-induced play behaviours in a sex-specific way, and that flat USVs are associated with non-play activities. Rats were allocated to one of two treatment groups: tickling (tickled, n = 16/sex) or no hand contact (control, n = 16/sex). Play behaviours (hopping, darting and hand approaches) and FM and flat USVs emitted during the testing session were quantified for each rat, with the frequency of FM and flat USVs made in anticipation of, and during, each behaviour analysed. In females, play behaviours were associated with more flat USVs than in males (before and during; p < 0.001), irrespective of treatment. FM USVs were paired with hopping and darting (before and during; p < 0.001), and in anticipation of hand approaches (p < 0.001) in both tickled females and males compared to controls (both sexes) suggesting that FM USVs are linked with play behaviour. The higher call rate of flat USVs paired with play behaviour in females suggests that there may be sex differences in the role of flat USVs during play. This result is evidence of sex differences in tickle-induced behaviours and has implications for our understanding of the function of different USVs in juvenile female and male rats.
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Female rats produce more USVs than males in response to tickling.
“We have also recently found evidence of sex differences with female rats producing more USVs than males in response to tickling ( Tivey et al., 2022).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Tivey et al., 2022
The Discussion and Conclusions (page not specified) report that tickled female rats produced more flat 50 kHz USVs associated with play-related behaviours than males. The claim omits those subtype and behavioural qualifiers, implying that females produced more USVs overall; the paper does not establish that broader comparison. (LLM, conf=0.88)
Experimental — accuracy not yet measuredTivey et al. (2022) found that playfully handled and control females produced more flat calls associated with play behaviours, performed during the playful handling sessions, than did males; in contrast both sexes produced trills in response to playful handling.
“Tivey et al., (2022) found that playfully handled and control females produced more flat calls associated with play behaviours, performed during the playful handling sessions, than did males; in contrast both sexes produced trills in response to playful handling.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Tivey et al., (2022)
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Experimental — accuracy not yet measuredTivey et al. (2022) found that after 10 days of playful handling, females produced more USVs than males with an indication that female controls also increased calling relative to male controls.
“Tivey et al., (2022) also found that after 10 days of playful handling, females produced more USVs than males with an indication that female controls also increased calling relative to male controls.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Tivey et al., (2022)
The Methods describe ten days of testing, and the Results (USV production associated with play-related behaviours; page unavailable) report more flat 50 kHz USVs in females than males around certain play behaviours. They also report more flat calls from female than male controls in the second before hopping. The claim omits those subtype and behaviour limits: the paper does not establish that females produced more USVs overall or that female controls’ calling increased over time. (LLM, conf=0.84)
Experimental — accuracy not yet measuredBased on previous data from the authors' own and other work where treatments come closest to matching the proposed treatments, the most variable response measure is total 50 kHz USVs.
“Based on our previous data, the most variable response measure is total 50 kHz USVs with data taken from our own and other work ( LaFollette et al., 2018; Tivey et al., 2022) where treatments come closest to matching the proposed treatments.”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as Tivey et al., 2022
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- Authors
- Tivey, E. K. L., Martin, J. E., Brown, S. M., Bombail, V., Lawrence, A. B., & Meddle, S. L.
- Affiliations
- Roslin Institute (GB)
- Scotland's Rural College (GB)
- Corresponding author
- Tivey, Emma K. L.
- ORCID
- Venue
- Scientific Reports
- Published
- 2022-09-12
- Volume
- 12
- Issue
- 1
- Pages
- 15323-15323
- ISSN
- 2045-2322 (ISSN-L)
- PMC
- PMC9468157
- OpenAlex
- W4295292862
- Cited by
- 13
- References
- 63
- Citations by year
- 2026: 4 · 2025: 5 · 2024: 1 · 2023: 3
- FWCI
- 2.07
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Scientific Reports (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Edinburgh Research Explorer (University of Edinburgh) (submittedVersion, cc-by)
- Newcastle University ePrints (Newcastle Univesity) (cc-by)
- Funders
- University of Edinburgh
- Biotechnology and Biological Sciences Research Council (BB/P013759/1)
- Rural and Environment Science and Analytical Services Division
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Keywords
- tickling, 50-kHz ultrasonic vocalizations, sex differences, frequency-modulated calls, positive affect, social communication, young rats
- MeSH terms
- Animals, Female, Male, Sex Characteristics, Ultrasonics, Vocalization, Animal, Rats
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