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Cited by 31Neuroendocrine regulation and behaviorPrimate Behavior and EcologyHuman-Animal Interaction Studies

Helping behavior in prairie voles: a model of empathy and the importance of oxytocin

Open access
Kitano, Kota, Yamagishi, Atsuhito, Horie, Kengo, Nishimori, Katsuhiko, Sato, Nobuya·2022·iScience
DOI 10.1016/j.isci.2022.103991PMC PMC8931361

Published version of a preprint

Open this paper in CoChat — free accountOpen PDF

Abstract

Several studies suggest that rodents show empathic responses and helping behavior toward others. We examined whether prairie voles would help conspecifics who were soaked in water by opening a door to a safe area. Door-opening latency decreased as task sessions progressed. Female and male voles stayed close to the soaked voles' side at equal rates and opened the door with similar latencies. When the conspecific was not soaked in water, the door-opening latency did not decrease. This suggests that the distress of the conspecific is necessary for learning to open the door and that the door-opening performed by prairie voles corresponds to helping behavior. Additionally, we examined the helping behavior in prairie voles in which oxytocin receptors were genetically knocked out. Oxytocin receptor knockout voles demonstrated less learning of the door-opening behavior and less interest in soaked conspecifics. This suggests that oxytocin is important for the emergence of helping behavior.

AI snapshot

Written from the paper’s full text.

TL;DR

Prairie voles learned to open a door to rescue a soaked cagemate, and this helping behavior was reduced when the cagemate was not distressed and in oxytocin receptor knockout voles. The authors conclude that conspecific distress is important for learning door-opening and that oxytocin is important for helping behavior.

Key findings

  • Helper voles’ door-opening latency decreased across seven sessions (session effect: F(6, 144) = 23.35, p < 0.001; sessions 1–2 longer than sessions 6–7, ps < 0.01), with no main effect of sex combination or helper’s sex.
  • When cagemates were not soaked, door-opening did not substantially decrease; Distress vs No Distress differed by group (F(1, 17) = 5.98, p = 0.003) and session (F(6, 102) = 5.97, p < 0.001), and Distress voles had more total openings (t(16.26) = 3.30, p = 0.004) and consecutive openings (t(16.83) = 2.98, p = 0.009).
  • Oxtr KO helpers showed less learning and less interest in soaked voles: latency group effect F(1, 20) = 27.93, p < 0.001; total openings t(8.57) = 4.41, p = 0.002; consecutive openings t(12.35) = 4.74, p < 0.001; time on soaked vole’s side F(1, 20) = 41.87, p < 0.001; huddling F(1, 20) = 9.49, p = 0.006.
  • Opposite-sex pairs tended to be less consistent: total door-opening sessions showed a sex-combination effect (F(1, 24) = 4.32, p = 0.049), and consecutive sessions were close to significant (F(1, 24) = 3.32, p = 0.081).

Methods

  • Used 80 experimentally naïve wildtype prairie voles (52 males, 28 females) and 16 Oxtr KO voles (8 males, 8 females); mean age 21.1 weeks, mean weight 38.3 g; helper and soaked voles were littermates.
  • Door-opening task: helper in ground area, soaked cagemate in pool area with 25 mm water; helper could open a circular door; one trial/day for 7 days, max 10 min, followed by 2-min interaction if opened; habituation was 10 min/day for 2 days.
  • Measured door-opening latency, time on soaked vole’s side, huddling, and habituation time in ground/pool areas; analyzed with ANOVAs, Holm post hoc tests, and Welch’s t-tests in RStudio.

Limitations

  • The paper states that Oxtr KO deficits could reflect increased water aversion, reduced social motivation, or hypoactivity; the Oxtr KO voles were not tested in a No Distress condition, and they were compared with wildtype voles of a different lineage rather than littermate controls.
  • The paper states that all helper and soaked voles were littermates, so effects of familiarity and kinship on helping behavior remain unclear.
  • Evident from the text: sample sizes were small, especially for Oxtr KO helpers (n = 8), and some key comparisons were trends or close to significance rather than definitive.

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

Integrity signals

Registry

Crossref · 10.1016/j.isci.2022.103991

Cites

88 of 88 references verified0 citation errors · 0 not checkable

Statistics

44 tests recomputed · 0 with the reported and recomputed p on opposite sides of .05

Claims the paper makes

0 of 35 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    There was no main effect of sex combination on door-opening latency in prairie voles.

    “A three-way ANOVA for mixed design with between-subjects factors of sex combination (same-sex, opposite-sex) and helper’s sex (male, female), and a within-subject factor of session (7) revealed no main effect of sex combination (F(1, 24) = 2.51, p = 0.126) and helper’s sex (F(1, 24) < 0.01, p = 0.987).”
    Quote in the paperSubstantial claim
  • Findingown data

    Door-opening latency decreased over the seven sessions in both same-sex and opposite-sex pairs of prairie voles.

    “Door-opening latency decreased as task sessions progressed.”
    • Also stated in: § Prairie voles demonstrate door-opening behavior regardless of the sex · plain
      “The latency of door-opening decreased over the seven sessions in both the same-sex and opposite-sex pairs (Figures 1C, 1D, 1E, and S1).”
    Quote in the paperSubstantial claim
  • Findingown data

    The total number of door-opening sessions was larger in wild-type voles than in Oxtr KO voles.

    “Both the total number of door-opening sessions (t(8.57) = 4.41, p = 0.002) and the number of consecutive door-opening sessions (t(12.35) = 4.74, p < 0.001) were larger in the wild-type group than in the Oxtr KO group (Figure 3D).”
    Quote in the paperSubstantial claim
  • Findingown data

    There was no main effect of group on huddling duration between Distress and No Distress groups.

    “A two-way ANOVA revealed neither a main effect of group (F(1, 14) = 0.13, p = 0.729), session (F(6, 126) = 1.44, p = 0.208), nor a group × session interaction (F(6, 84) = 0.66, p = 0.681), suggesting that the huddling of prairie voles after the door is opened is not affected by whether the cagemate is soaked or not.”
    Quote in the paperNot a substantial claim
  • Findingown data

    During habituation, helper voles stayed longer in the ground area than the expected 300 s.

    “The time spent in the ground area was significantly longer than the expected value (300 s, half of the habituation period, t(27) = 23.58, p < 0.001).”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim. 14
  • May combine two claims. 2

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

    Premisecitation

    Rats demonstrate helping behavior toward distressed conspecifics, such as those in a narrow tube or in water.

    “Recent studies suggest that rats also demonstrate helping behavior toward distressed conspecifics, such as those in a narrow tube (Bartal et al., 2011, 2016) and in water (Sato et al., 2015; Yamagishi et al., 2019, 2020).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Bartal et al., 2011, 2016

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

    • cites Sato et al., 2015 source

      Not checked in the free run.

    • cites Yamagishi et al., 2019, 2020

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

  • May combine two claims.

    Premisecitation

    Previous studies suggest that prairie voles show empathy-like behavior, such as freezing caused by emotional contagion of conspecifics' fear, and consolation.

    “Previous studies suggest that prairie voles show empathy-like behavior, such as freezing caused by emotional contagion of conspecifics' fear, and consolation (Burkett et al., 2016; Stetzik et al., 2018; Wardwell et al., 2020).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Burkett et al., 2016 source

      Not checked in the free run.

    • cites Stetzik et al., 2018 source

      Not checked in the free run.

    • cites Wardwell et al., 2020 source

      Not checked in the free run.

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

    Study design and conductown data

    The present study did not examine helping behavior when the effects of social interaction are excluded.

    “However, that condition was not examined in the present study.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    Oxtr KO voles demonstrate less helping behavior and less interest in the soaked voles than wild-type voles.

    “These results suggest that the Oxtr KO voles demonstrate less helping behavior and less interest in the soaked voles than the wild-type voles.”
    Quote in the paperSubstantial claim

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

    Interpretationreasoning

    Such an avoidance tendency might have caused the delay in the appearance of helping behavior toward opposite-sex conspecifics.

    “Although the details are issues for future research, such an avoidance tendency might have caused the delay in the appearance of helping behavior.”
    Quote in the paperNot a substantial claim

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

    Premisecitation

    Oxytocin receptor knockout voles demonstrated autism-like behavior such as a lack of interest in social novelty.

    “A previous study reported that oxytocin receptor knockout voles demonstrated autism-like behavior such as a lack of interest in social novelty (Horie et al., 2019).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Horie et al., 2019 source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductcitation

    Huddling was used as an index for social attachment in the present study in accordance with a previous report.

    “We used huddling as an index for social attachment in the present study in accordance with a previous report (Barrett et al., 2015).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Barrett et al., 2015 source

      Not checked in the free run.

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

    Premisecitation

    Huddling in prairie voles has generally been explored in the context of a partner preference test.

    “However, huddling in prairie voles has generally been explored in the context of a partner preference test (Walum and Young, 2018).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Walum and Young, 2018 source

      Not checked in the free run.

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

    Premisecitation

    Various factors of social context are involved in empathy.

    “Various factors of social context are involved in empathy (Bartal et al., 2014; Yamagishi et al., 2019).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Bartal et al., 2014 source

      Not checked in the free run.

    • cites Yamagishi et al., 2019 source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    In the present study, door-opening behavior in Oxtr KO voles was not examined with the No Distress condition in which the cagemate is not soaked in water.

    “In the present study, however, the door-opening behavior in the Oxtr KO voles was not examined with the No Distress condition in which the cagemate is not soaked in water.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Water at a depth of 25 mm is aversive to prairie voles.

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

    Interpretationnone given

    The present study implies a relationship between helping behavior and oxytocin in prairie voles.

    “The present study implies a relationship between helping behavior and oxytocin in prairie voles.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: Water at a depth of 25 mm is aversive to prairie voles.

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

    Study design and conductnone given

    In the present study, helping behavior in Oxtr KO voles was compared with that in wild-type voles with a different lineage.

    “In addition, in the present study, we compared helping behavior in the Oxtr KO voles with that in wild-type voles with a different lineage.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: Water at a depth of 25 mm is aversive to prairie voles.

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

    Premisecitation

    Prairie voles exhibit allogrooming that functions as social buffering to relieve the fear of their partner.

    “Prairie voles exhibit allogrooming that functions as social buffering to relieve the fear of their partner.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Burkett et al., 2016 source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    Helper and soaked voles were randomly assigned to individuals in the same cage.

    “Helper and soaked voles were randomly assigned to individuals in the same cage.”
    Quote in the paperSubstantial claim

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

    Premisecitation

    A previous study examining emotional contagion of pain in mice suggests that the pain of familiar individuals is more contagious than that of strangers.

    “A previous study examining the emotional contagion of pain in mice suggests that the pain of familiar individuals is more contagious than that of strangers (Langford et al., 2006).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Langford et al., 2006 source

      Not checked in the free run.

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

    Premisecitation

    Rats display as much helping behavior toward strangers as toward cagemates.

    “Rats display as much helping behavior toward strangers as toward cagemates (Bartal et al., 2014; Yamagishi et al., 2019).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Bartal et al., 2014 source

      Not checked in the free run.

    • cites Yamagishi et al., 2019 source

      Not checked in the free run.

16 of 16

  • The paper relies on this without support.

    Helping behavior that rescues conspecifics from distressed situations has not been elucidated in prairie voles.

    “However, helping behavior that rescues conspecifics from distressed situations has not been elucidated in prairie voles.”

  • The paper relies on this without support.

    Water at a depth of 25 mm is aversive to prairie voles.

    “During the experiment, the pool area contained water with a depth of 25 mm to create an aversive situation for the prairie voles.”
2 of 2

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?

Statistics that don’t add up

Each reported p-value recomputed from its test statistic and degrees of freedom.

  • p-value fits an unstated one-tailed test: t(19.09) = 3.73, p < .001

    Minor

    The paper reports p < .001 here; recomputing from the test statistic and its degrees of freedom gives p = .001 (two-tailed). The reported value is what a one-tailed test gives, but the paper does not say near this result that a one-tailed test was used.

    Reported p < .001 · recomputed p = .001

  • Inconsistent p-value: F(1, 17) = 5.98, p = .003

    Minor

    The paper reports p = .003 here; recomputing from the test statistic and its degrees of freedom gives p = .026. The check cannot tell whether the statistic or the p-value is the misprint.

    Reported p = .003 · recomputed p = .026

  • Inconsistent p-value: F(6, 126) = 1.44, p = .208

    Minor

    The paper reports p = .208 here; recomputing from the test statistic and its degrees of freedom gives p = .204. The check cannot tell whether the statistic or the p-value is the misprint.

    Reported p = .208 · recomputed p = .204

3 of 3

Statistics recomputed

Every reported test result, recomputed from its test statistic and degrees of freedom. A result shows as not adding up only once an independent check has confirmed it.

  • Doesn’t add upF(1, 17) = 5.98reported p = .003 · recomputed p = .026

    “F(1, 17) = 5.98, p = .003”

  • Doesn’t add upF(6, 126) = 1.44reported p = .208 · recomputed p = .204

    “F(6, 126) = 1.44, p = .208”

  • Doesn’t add upt(19.09) = 3.73reported p < .001 · recomputed p = .001

    “t(19.09) = 3.73, p < .001”

  • Not confirmedF(6, 102) = 1.17reported p = .325 · recomputed p = .328

    “F(6, 102) = 1.17, p = .325”

  • ConsistentF(6, 144) = 23.35reported p < .001 · recomputed p < .001

    “F(6, 144) = 23.35, p < .001”

  • ConsistentF(1, 24) = 2.51reported p = .126 · recomputed p = .126

    “F(1, 24) = 2.51, p = .126”

  • ConsistentF(1, 24) < 0.01reported p = .987 · recomputed p = .921

    “F(1, 24) < 0.01, p = .987”

  • ConsistentF(6, 144) = 1.88reported p = .087 · recomputed p = .088

    “F(6, 144) = 1.88, p = .087”

  • ConsistentF(1, 24) = 4.32reported p = .049 · recomputed p = .049

    “F(1, 24) = 4.32, p = .049”

  • ConsistentF(1, 24) = 0.04reported p = .852 · recomputed p = .843

    “F(1, 24) = 0.04, p = .852”

  • ConsistentF(1, 24) < 0.01reported p > .999 · recomputed p = .921

    “F(1, 24) < 0.01, p > .999”

  • ConsistentF(1, 24) = 3.32reported p = .081 · recomputed p = .081

    “F(1, 24) = 3.32, p = .081”

  • ConsistentF(1, 24) = 0.01reported p = .924 · recomputed p = .921

    “F(1, 24) = 0.01, p = .924”

  • Consistentt(27) = 23.58reported p < .001 · recomputed p < .001

    “t(27) = 23.58, p < .001”

  • Consistentt(22.57) = 0.63reported p = .536 · recomputed p = .535

    “t(22.57) = 0.63, p = .536”

  • ConsistentF(1, 24) = 2.11reported p = .16 · recomputed p = .159

    “F(1, 24) = 2.11, p = .160”

  • ConsistentF(1, 24) = 0.1reported p = .756 · recomputed p = .755

    “F(1, 24) = 0.10, p = .756”

  • ConsistentF(6, 144) = 0.39reported p = .885 · recomputed p = .885

    “F(6, 144) = 0.39, p = .885”

  • ConsistentF(6, 144) = 2.39reported p = .031 · recomputed p = .031

    “F(6, 144) = 2.39, p = .031”

  • ConsistentF(1, 24) = 0.95reported p = .34 · recomputed p = .339

    “F(1, 24) = 0.95, p = .340”

  • ConsistentF(1, 24) = 0.65reported p = .428 · recomputed p = .428

    “F(1, 24) = 0.65, p = .428”

  • ConsistentF(6, 102) = 5.97reported p < .001 · recomputed p < .001

    “F(6, 102) = 5.97, p < .001”

  • Consistentt(16.26) = 3.3reported p = .004 · recomputed p = .005

    “t(16.26) = 3.30, p = .004”

  • Consistentt(16.83) = 2.98reported p = .009 · recomputed p = .008

    “t(16.83) = 2.98, p = .009”

  • ConsistentF(1, 14) = 0.89reported p = .362 · recomputed p = .361

    “F(1, 14) = 0.89, p = .362”

  • ConsistentF(6, 84) = 1.44reported p = .208 · recomputed p = .209

    “F(6, 84) = 1.44, p = .208”

  • ConsistentF(6, 84) = 1.36reported p = .239 · recomputed p = .240

    “F(6, 84) = 1.36, p = .239”

  • ConsistentF(1, 14) = 0.13reported p = .729 · recomputed p = .724

    “F(1, 14) = 0.13, p = .729”

  • ConsistentF(6, 84) = 0.66reported p = .681 · recomputed p = .682

    “F(6, 84) = 0.66, p = .681”

  • ConsistentF(1, 20) = 27.93reported p < .001 · recomputed p < .001

    “F(1, 20) = 27.93, p < .001”

  • ConsistentF(6, 120) = 18.84reported p < .001 · recomputed p < .001

    “F(6, 120) = 18.84, p < .001”

  • ConsistentF(6, 120) = 2.01reported p = .07 · recomputed p = .069

    “F(6, 120) = 2.01, p = .070”

  • Consistentt(8.57) = 4.41reported p = .002 · recomputed p = .002

    “t(8.57) = 4.41, p = .002”

  • Consistentt(12.35) = 4.74reported p < .001 · recomputed p < .001

    “t(12.35) = 4.74, p < .001”

  • ConsistentF(1, 20) = 41.87reported p < .001 · recomputed p < .001

    “F(1, 20) = 41.87, p < .001”

  • ConsistentF(6, 120) = 0.57reported p = .757 · recomputed p = .754

    “F(6, 120) = 0.57, p = .757”

  • ConsistentF(6, 120) = 1.55reported p = .169 · recomputed p = .168

    “F(6, 120) = 1.55, p = .169”

  • ConsistentF(1, 20) = 9.49reported p = .006 · recomputed p = .006

    “F(1, 20) = 9.49, p = .006”

  • ConsistentF(6, 120) = 1.51reported p = .181 · recomputed p = .181

    “F(6, 120) = 1.51, p = .181”

  • ConsistentF(6, 120) = 0.66reported p = .685 · recomputed p = .682

    “F(6, 120) = 0.66, p = .685”

  • Consistentt(7) = 179.29reported p < .001 · recomputed p < .001

    “t(7) = 179.29, p < .001”

  • Consistentt(14.42) = 5.45reported p < .001 · recomputed p < .001

    “t(14.42) = 5.45, p < .001”

  • Consistentt(13.68) = 0.98reported p = .342 · recomputed p = .344

    “t(13.68) = 0.98, p = .342”

  • ConsistentF(1, 18) = 1reported p = .331 · recomputed p = .331

    “F(1, 18) = 1.00, p = .331”

44 of 44

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.

  • Huddling ANOVA reports incompatible denominator degrees of freedom

    Minor

    The session effect and group-by-session interaction are reported from the same two-way mixed ANOVA, but their denominator degrees of freedom are 126 and 84. With the same seven-session factor and observations, both effects should use the same denominator degrees of freedom.

1 of 1

Cites

Flagged first, then by citations

Research works printed but not linked: 1

  1. Getz, 1996, Prairie-vole partnerships: this rodent forms social groups that appear to have evolved as an adaptation for living in a low-food habitat, Am. Sci., 84, 56

Cited by

Sorted by citations

Related

Authors
Kitano, K., Yamagishi, A., Horie, K., Nishimori, K., & Sato, N.
Affiliations
  • Kwansei Gakuin University (JP)
  • Emory University (US)
  • Fukushima Medical University (JP)
Corresponding author
Sato, Nobuya
ORCID
Venue
iScience
Publisher
Elsevier BV
Published
2022-04
Volume
25
Issue
4
Pages
103991
ISSN
2589-0042 (ISSN-L)
DOI
10.1016/j.isci.2022.103991
PMC
PMC8931361
OpenAlex
W4220830117
Semantic Scholar
2e1586e714864f7e458753869fa0258187a54e06
Cited by
31
References
88
Citations by year
2026: 5 · 2025: 8 · 2024: 8 · 2023: 4 · 2022: 3 · 2021: 3
FWCI
4.41
Type
journal article
Language
en
Open access
Licence
Locations
  • iScience (publishedVersion, cc-by)
  • SSRN Electronic Journal (publishedVersion)
  • bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Funders
  • Ministry of Education, Culture, Sports, Science and Technology (S1511032, 2015-2019)
  • Kwansei Gakuin University
  • Japan Society for the Promotion of Science (21H00965, 18K07357, 21H05242)
Topics
  • Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
  • Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Human-Animal Interaction Studies
Keywords
helping behavior, Microtus ochrogaster, empathy, oxytocin, social learning, sex differences, rodent social behavior

About these checks

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.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-09.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-09.

Statistics and arithmetic are recomputed, not judged.

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