Rat tickling: A systematic review of applications, outcomes, and moderators.
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INTRODUCTION: Rats initially fear humans which can increase stress and impact study results. Additionally, studying positive affective states in rats has proved challenging. Rat tickling is a promising habituation technique that can also be used to model and measure positive affect. However, current studies use a variety of methods to achieve differential results. Our objective was to systematically identify, summarize, and evaluate the research on tickling in rats to provide direction for future investigation. Our specific aims were to summarize current methods used in tickling experiments, outcomes from tickling, and moderating factors. METHODS: We systematically evaluated all articles about tickling identified from PubMed, Scopus, Web of Science, and PsychInfo. Our inclusion criteria were publication in a peer-reviewed journal and collection of original, empirical data on rats using the handling method of tickling. One researcher extracted information from each article. Bias was assessed by 2 investigators using the SYRCLE bias assessment tool. RESULTS: We identified 32 articles (56 experiments) published in peer-reviewed journals about rat tickling for inclusion. A wide variety of strains, sexes, and ages of rats were included. The most common method used for tickling was cycling through 15 seconds of tickling and 15 seconds of rest for 2 minutes for 3 to 5 days. Experiments with a control for tickling (N = 22) showed that tickling increases positive vocalization, approach behavior, decreases anxiety measures, improves handling, and in some cases decreases stress hormones. Tickling juvenile, individually housed rats with a trait predisposition to respond more positively to tickling, results in the most positive outcomes. Methods to reduce bias were insufficiently reported. CONCLUSIONS: We conclude that tickling is a promising method for improving rat welfare and investigating positive affect. However, the establishment of tickling best practices is essential as the outcomes from tickling can be moderated by several factors.
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Rats find tickling rewarding, as tickled rats show reduced latency for approaching the human hand and can be trained to perform operant responses for tickling.
“Other evidence suggests that rats find tickling rewarding as in addition to production of USVs, tickled rats show a reduced latency for approaching the human hand and can be trained to perform operant responses for tickling ( Burgdorf & Panksepp, 2001; LaFollette et al., 2017).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as LaFollette et al., 2017
The Results section (‘Approach behavior’; page unavailable) reports that nine of ten experiments assessing approach found positive results, including decreased approach latency. The review also discusses tickling’s reward value, but the passages in our copy do not report rats being trained to perform operant responses for tickling. (LLM, conf=0.87)
Experimental — accuracy not yet measuredOver 60% of papers on tickling used rats within the 3-4 to 7 week age window.
“We purchased the rats from CRL (16 at a time), aged 3-4 weeks and completed the research prior to their puberty around 7 weeks of age as over 60% of papers on tickling used rats within this time window ( LaFollette et al., 2017).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as LaFollette et al., 2017
Table 3 (Results, Characteristics of tickling; page unavailable) lists many studies using rats aged 21–49 days, consistent with the stated age window. The Discussion (Outcomes of tickling; page unavailable) reports that 81% of experiments used singly housed rats under 50 days old, but it does not establish that over 60% of papers used rats throughout the specified 3–4-to-7-week window. The reported percentage also counts experiments, not papers. (LLM, conf=0.84)
Experimental — accuracy not yet measuredPlay fighting between rats, as well as play between a rat and a human, are known to elicit intense 50 kHz calling, while threatening scenarios, such as an intruder or exposure to a predator, are more commonly associated with 22 kHz calls.
“Play fighting between rats, as well as play between a rat and a human, are known to elicit intense 50 kHz calling,20,25 while threatening scenarios, such as an intruder or exposure to a predator is more commonly associated with 22 kHz calls.46”
From Ultrasonic Vocalization Analysis as a Novel Metric to Assess Cage Enrichment in Rats (2022) · cites this paper as [25]
Wistar rats have been shown to be reliably responsive to tickling and PH.
“This outbred albino strain has been shown to be reliably responsive to tickling and PH ( Bombail et al., 2019; Hammond et al., 2019) and is one of the commonly used strains for such experiments (23% of rat tickling studies used Wistar rats; LaFollette et al., 2017).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as LaFollette et al., 2017
When rats are tickled by the human hand, they often produce 50 kHz USVs, which are understood to indicate positive affect in rats.
“Research (see Bombail et al., 2021; LaFollette et al., 2017 for reviews) has shown that when rats are tickled by the human hand, they often produce 50 kHz USVs, which are understood to indicate positive affect in rats ( e.g., Barker, 2018, Figure 2i).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as LaFollette et al., 2017
Other work has reported significant levels of individual variation in response to tickling.
“Other work has reported significant levels of individual variation in response to tickling ( LaFollette et al., 2017).”
From Stage 2 Registered Report: Refinement of tickling protocols to improve positive animal welfare in laboratory rats (2025) · cites this paper as LaFollette et al., 2017
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“We purchased the rats from CRL (16 at a time), aged 3-4 weeks and completed the research prior to their puberty around 7 weeks of age as over 60% of papers on tickling used rats within this time window (LaFollette et al., 2017).”
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- Authors
- LaFollette, M. R., O’Haire, M. E., Cloutier, S., Blankenberger, W. B., & Gaskill, B. N.
- Affiliations
- Purdue University West Lafayette (US)
- Canadian Council on Animal Care (CA)
- Stanford University (US)
- Corresponding author
- LaFollette, Megan R.
- Venue
- PLOS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2017-04-06
- Volume
- 12
- Issue
- 4
- Pages
- e0175320
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC5383284
- OpenAlex
- W2604855621
- Cited by
- 84
- References
- 59
- Citations by year
- 2026: 4 · 2025: 16 · 2024: 14 · 2023: 10 · 2022: 8 · 2021: 6 · 2020: 6 · 2019: 9 · 2018: 10 · 2017: 1
- FWCI
- 1.61
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- PLoS ONE (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- RePEc: Research Papers in Economics (acceptedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (acceptedVersion)
- Figshare (submittedVersion, cc-by)
- Topics
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Life Sciences › Neuroscience › Behavioral Neuroscience › Stress Responses and Cortisol
- Life Sciences › Neuroscience › Cognitive Neuroscience › Tactile and Sensory Interactions
- Keywords
- positive affect, animal welfare, habituation, handling stress, anxiety, stress hormones, vocalizations, approach behavior, rat strain differences, young rats, individual housing, research bias
- MeSH terms
- Animals, Animal Communication, Anxiety, Behavior, Animal, Fear, Female, Humans, Male, Touch, Rats
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