Cited by 139Neuroendocrine regulation and behaviorNeuropeptides and Animal PhysiologyStress Responses and Cortisol

Extrahypothalamic oxytocin neurons drive stress-induced social vigilance and avoidance

Open access
Duque-Wilckens, Natalia, Torres, Lisette Y., Yokoyama, Sae, Minie, Vanessa A., Tran, Amy M., Petkova, Stela P., Hao, Rebecca, Ramos-Maciel, Stephanie +8 more·2020·Proceedings of the National Academy of Sciences

Published version of a preprint

Abstract

Significance: The neuropeptide oxytocin is an important regulator of social behavior and is widely considered to reduce anxiety-related behaviors. However, growing evidence suggests that sometimes oxytocin increases anxiety. How can the same molecule have such different effects on behavior? Here we provide evidence that oxytocin produced outside of the hypothalamus is necessary and sufficient for stress-induced social anxiety behaviors. This suggests that the diverse effects of oxytocin on anxiety-related behaviors are mediated by circuit-specific oxytocin action.

AI snapshot

Written from the paper’s full text.

TL;DR

In female California mice, oxytocin synthesis in medioventral bed nucleus of the stria terminalis (BNSTmv) is necessary for social defeat–induced reductions in social approach and increases in social vigilance. BNSTmv oxytocin neurons project to anteromedial BNST (BNSTam), where oxytocin infusion is sufficient to induce social avoidance and vigilance in stress-naïve males and females.

Key findings

  • BNSTmv antisense morpholino reduced oxytocin-immunoreactive cells compared with missense controls (F1,28=5.4, p=0.02) and prevented stress-induced reductions in social approach (stress×treatment F1,31=5.7, p=0.02) and increases in social vigilance (stress×treatment F1,31=6.2, p=0.02) in female California mice.
  • Male California mice tested 10 min after a third social defeat episode showed reduced social approach (t14=2.46, p=0.02, d=1.23) and increased social vigilance (t14=2.36, p=0.03, d=1.18) versus naïve males.
  • Viral tracing showed BNSTmv oxytocin neurons project to BNSTam and anterior/lateral hypothalamus; on average 63% of oxytocin-immunoreactive cells expressed Venus, and all Venus-positive cells coexpressed oxytocin.
  • Oxytocin infusion into BNSTam in stress-naïve mice reduced social approach and increased social vigilance; in females, 1.0 ng reduced approach (p<0.001, d=4.8) and increased vigilance (p<0.001, d=2.3), and in males, 1.0 ng reduced approach (p=0.00, d=1.93) and increased vigilance (p<0.001, d=1.27). In C57BL/6 mice, vicarious defeat increased vigilance in females (p<0.0001, d=1.9) and BNST oxytocin cell number (t14=2.721, p=0.01, d=1.4), but not PVN oxytocin cells.

Methods

  • California mice underwent social defeat or control handling; one week later, females received BNSTmv antisense or missense morpholino injections, followed by social interaction testing. Male California mice were tested 10 min after a third defeat episode.
  • C57BL/6 mice underwent vicarious social defeat or control handling and social interaction testing; BNST and PVN oxytocin cells were assessed immunohistochemically.
  • Circuit mapping used AAV-OTpr-Venus tracing from BNSTmv; qPCR and fluorescent in situ hybridization assessed Oxtr/Gad1/vGlut2 in BNSTam; site-specific oxytocin infusions into BNSTam were tested in stress-naïve females and males. Analyses used 2-way ANOVAs, t-tests, planned comparisons, and Cohen’s d.

Limitations

  • Stated: the authors do not know whether BNST oxytocin neurons synapse within target regions, and non-synaptic/periaxonal or en passant release is possible; a role of PVN oxytocin is not excluded; future single-cell analysis is needed to understand how Oxtr activation in BNSTam modulates behavior.
  • Evident: oxytocin synthesis knockdown in BNSTmv was tested only in female California mice, while males received acute defeat testing and BNSTam oxytocin infusion but not antisense knockdown.
  • Evident: the study does not directly measure oxytocin release in BNSTam during stress or social interaction.

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

Integrity signals

Registry

Crossref · 10.1073/pnas.2011890117

Cites

87 of 87 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 34 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    Viral tracing of BNSTmv oxytocin neurons revealed fibers in regions controlling defensive behaviors including lateral hypothalamus, anterior hypothalamus, and anteromedial BNST (BNSTam).

    “Viral tracing of BNSTmv oxytocin neurons revealed fibers in regions controlling defensive behaviors including lateral hypothalamus, anterior hypothalamus, and anteromedial BNST (BNSTam).”
    Quote in the paperSubstantial claim
  • Findingown data

    Oxytocin knock-down in BNSTmv prevented stress-induced increases in social vigilance and decreases in social approach.

    “Oxytocin knock-down prevented stress-induced increases in social vigilance and decreases in social approach.”
    Quote in the paperSubstantial claim
  • Findingown data

    There were no basal or stress-induced sex differences in Oxtr expression within the BNSTam.

    “First, we used real-time PCR to assess whether there were basal or stress-induced sex differences in Oxtr expression within the BNSTam and there were no differences (fig., 3I).”
    Quote in the paperNot a substantial claim
  • Findingown data

    The effect of social defeat on social approach was dependent on antisense treatment.

    “The effect of social defeat on social approach, measured by time spent in the interaction zone, was dependent on antisense treatment (F 1,31 =5.7, p=0.02, fig.1E).”
    Quote in the paperSubstantial claim
  • Findingown data

    BNST antisense infusions had no effects on PVN oxytocin cells.

    “There were no effects of BNST antisense infusions on PVN oxytocin cells (supplementary fig.1F), indicating oxytocin knockdown in the BNST was site-specific.”
    Quote in the paperSubstantial claim
  • May combine two claims. 21
  • Classified as probably not a substantial claim. 11
  • The quote only partly states this claim. 2
  • May combine two claims.

    Findingown data

    Oxytocin infusion into BNSTam in stress naïve mice increased social vigilance and reduced social approach.

    “Oxytocin infusion into BNSTam in stress naïve mice increased social vigilance and reduced social approach.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.
  • May combine two claims.

    Findingown data

    There were no effects of defeat on approach behavior when target mouse was absent or in open field in males.

    “There were no effects of defeat on approach behavior when the target mouse was absent (supplementary fig.2B) or in the open field (supplementary fig.2C, 2D).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    On average, 63% of oxytocin-immunoreactive cells expressed Venus and all Venus positive cells coexpressed oxytocin.

    “Immunohistochemical analyses in the BNST (fig.3C) and the PVN (supplementary fig.3A) showed that on average, 63% of oxytocin-immunoreactive cells expressed Venus and that all Venus positive cells coexpressed oxytocin.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Male California mice show reduced social approach and increased social vigilance 10 min after a third episode of defeat compared to naïve males.

    “When stressed males were tested 10 min after a third episode of defeat (Fig. 1H), social approach was reduced (t 14 =2.46, p=0.02, d=1.23, fig.1I), and social vigilance was increased (t 14 =2.36, p=0.03, d=1.18, fig.1J) compared to naïve males.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    There were no effects of stress or treatment on approach behavior during acclimation, time spent in center, or distance traveled.

    “There were no effects of stress or treatment on approach behavior during the acclimation phase when the target mouse was absent (supplementary fig.1C), or time spent in center (supplementary fig.1D) and distance traveled (supplementary fig.1D) during the open field phase.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    Stress increased social vigilance in missense treated females but not antisense treated females.

    “Stress increased social vigilance in missense treated females (p=0.03,d=0.96) but not antisense treated females.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Vicarious stress increased the number of oxytocin cells in the BNST but not in the PVN in female C57BL/6J mice.

    “Importantly, vicarious stress increased the number of oxytocin cells in the BNST (t 14 =2.721, p=0.01, d=1.4,fig 2D), but not in the PVN (fig.2E), comparable to what was previously reported in female California mice.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Defeat reduced social approach in females receiving missense, but not in females receiving antisense.

    “Defeat reduced social approach in females receiving missense (p=0.02,d=1.1), but not in females receiving antisense.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingcitation

    Social stress increases oxytocin cell mRNA and counts in the BNST of female California mice and C57BL6/J mice.

    “Social stress increases oxytocin cell mRNA and counts in the BNST of female California mice (Steinman et al., 2016) and C57BL6/J mice.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Steinman et al., 2016

      Not checked in the free run.

  • May combine two claims.

    Findingown data

    Stressed male California mice show reduced social approach and increased social vigilance at the time point after third episode of defeat.

    “We found that comparable to females, stressed male California mice show reduced social approach and increased social vigilance at this time point.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    In males, oxytocin increased social vigilance, with 1.0 ng increasing vigilance compared to vehicle.

    “Similarly, oxytocin also increased social vigilance (F 2,21 =5.3, p=0.01, fig.4E), with 1.0 ng of oxytocin increasing vigilance compared to vehicle (p<0.001,d=1.27).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.
  • May combine two claims.

    Findingown data

    In females, oxytocin treatment increased social vigilance, with 1.0 ng but not 0.2 ng dose increasing vigilance compared to vehicle.

    “Treatment with oxytocin also increased social vigilance in females (F 3,21 =8.8, p=0.001, fig.4C).The 1.0 ng (p<0.001,d=2.3) but not the 0.2 ng dose increased vigilance in stress naïve females compared to vehicle.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.
  • May combine two claims.

    Findingown data

    In males, oxytocin reduced social approach, with 1.0 ng reducing social approach compared to vehicle.

    “In males, oxytocin also reduced social approach (F 2,21 =6.7, p=0.00), with 1.0 ng of oxytocin reducing social approach compared to vehicle (p=0.00,d=1.93 fig.4D).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.
  • May combine two claims.

    Findingown data

    There were no vGlut2+ cells in the BNSTam, although there were vGlut2+ cells in the adjacent lateral septum.

    “We found no vGlut2+ cells in the BNSTam, although there were vGlut2+ cells in the adjacent lateral septum (supplementary fig.3B).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    Stress increased BNSTmv oxytocin cell number in animals receiving missense, but this increase was prevented by morpholino injections (as stated, missense).

    “Stress increased BNSTmv oxytocin cell number in animals receiving missense, but this was prevented by injections of morpholino missense.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    There were no differences in cycle thresholds between groups for B2m.

    “There were no differences in cycle thresholds between groups for B2m.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    No effects of stress or sex were found on OxtR expression in BNSTam after social defeat.

    “No effects of stress or sex were found.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    Social defeat did not affect approach, time in center, or distance traveled when there was no social target present.

    “Social defeat did not affect approach, time in center, or distance traveled when there was no social target present.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    No Glut+ cells were found in the BNSTam.

    “No Glut+ cells were found in the BNSTam, but there were Glut+ cells in the adjacent lateral septum.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    There were no effects of antisense injections on approach, time spent in center, or distance traveled when no social target was present.

    “There were no effects of antisense injections on approach, time spent in center, or distance traveled when no social target was present.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    There were Glut+ cells in the adjacent lateral septum.

    “No Glut+ cells were found in the BNSTam, but there were Glut+ cells in the adjacent lateral septum.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Premisecitation

    Oxytocin acting in hypothalamic-mesolimbic circuits promotes social approach.

    “Thus, while oxytocin acting in hypothalamic-mesolimbic circuits promotes social approach (Dölen et al., 2013;Marlin et al., 2015;Song et al., 2016;Wang and Aragona, 2004;Williams et al., 2020), our data in the BNST support the hypothesis that divergent effects of oxytocin on social approach are mediated by discrete brain circuits (Grinevich et al., 2016;Shamay-Tsoory and Abu-Akel, 2016;Steinman et”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Dölen et al., 2013

      Not checked in the free run.

    • cites Marlin et al., 2015

      Not checked in the free run.

    • cites Song et al., 2016

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    Stress-sensitive oxytocin neurons located in BNSTmv project to BNSTam, where oxytocin acts to drive a social anxiety-related phenotype.

    “Together, these data suggest that stress-sensitive oxytocin neurons located in the BNSTmv project to the BNSTam, where oxytocin acts to drive a social anxiety-related phenotype.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.
  • Classified as probably not a substantial claim. The quote only partly states this claim.

    Study design and conductcitation

    Female C57BL/6 mice were randomly assigned to control handling or vicarious social defeat stress.

    “Female mice were randomly assigned to control handling or vicarious social defeat stress as described previously (Iñiguez et al., 2018).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Iñiguez et al., 2018

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    The key role of BNST oxytocin neurons in long-term changes in female-biased social avoidance and vigilance suggests these cells may be related to symptoms associated with social anxiety disorder and depression, which are more common in women than men.

    “However, the key role of BNST oxytocin neurons play in long term changes in female-biased social avoidance and vigilance suggests that these cells may be related to symptoms associated with social anxiety disorder (Asher et al., 2017) and depression (Kessler et al., 2005), which are more common in women than men.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Asher et al., 2017

      Not checked in the free run.

    • cites Kessler et al., 2005

      Not checked in the free run.

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

    Interpretationcitation

    Oxytocin may act on inhibitory neurons in the BNST.

    “About 50% of Oxtr+ cells in the BNSTam coexpress Gad1+, suggesting that oxytocin may act on inhibitory neurons in the BNST as has been observed in the central nucleus of amygdala (Knobloch et al., 2012).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Knobloch et al., 2012

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductcitation

    Mice were randomly assigned to control handling or social defeat.

    “Mice were randomly assigned to control handling or social defeat for 3 consecutive days as described previously (Trainor et al., 2011).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Trainor et al., 2011

      Not checked in the free run.

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

    Study design and conductown data

    Animals were randomly assigned to receive morpholino missense or antisense.

    “One week after the last episode of control handling or social defeat, animals were randomly assigned to receive one bilateral 0.2uL injection of 100pM morpholino missense or antisense aimed a little anterior to medioventral BNST (BNSTmv, AP:0.39,ML: 1.1, DV:-5.85).”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Study design and conductown data

    Vigilance behavior was manually analyzed by an observer blind to treatment.

    “Vigilance behavior, defined as time spent with head oriented toward the target mouse while outside of the interaction zone, was manually analyzed by an observer blind to treatment.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim.

    Study design and conductown data

    Female California mice were randomly assigned to social defeat or control conditions.

    “To test whether stressinduced increases in oxytocin synthesis within the BNSTmv are necessary for stress-induced social deficits, females were first randomly assigned to social defeat or control conditions (Fig. 1B).”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. The quote only partly states this claim.

    Study design and conductown data

    Mice were randomly assigned to receive bilateral infusions of aCSF, 0.2ng, or 1ng oxytocin.

    “After 7 days, mice were randomly assigned to receive bilateral 200nl infusions of either aCSF, 0.2ng (females only), or 1ng (males and females) of oxytocin using internal guides that projected 1 mm past the guides.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationown data

    In Mus musculus, vicarious social stress increases social vigilance and BNST OT cell number in females.

    “In Mus musculus, vicarious social stress increases social vigilance and BNST OT cell number in females.”
    Quote in the paperSubstantial claim
32 of 32
  • The paper relies on this without support.

    Exogenous oxytocin infused into BNSTam acts through oxytocin receptors to produce its effects on social behavior.

    “Next, we tested whether administration of oxytocin within the BNSTam is sufficient to reduce social approach and increase vigilance in stress naïve females and males (fig.4A).”
  • The paper relies on this without support.

    Morpholino oligos inhibit protein synthesis by blocking the translation initiation complex.

    “Morpholino oligos inhibit protein synthesis by blocking the translation initiation complex.”
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?

  • In female California mice, SDS resulted in reduced social approach and increased social vigilance along with increased Oxt mRNA levels and co-localization of OXT with c-fos in the BNST.

    “In female California mice, SDS resulted in reduced social approach and increased social vigilance along with increased Oxt mRNA levels and co-localization of OXT with c-fos (a marker of cellular activity) in the bed nucleus of the stria ter­ minalis (BNST) (Duque-Wilckens et al., 2020).”
  • Knocking down OXT in the BNST prevented the SDS-induced increase in social vigilance and decrease in social approach, while OXT infusion into the BNST induced social vigilance in both stress-naïve female and male California mice.

    “Interestingly, knocking down OXT in the BNST prevented the SDS-induced increase in social vigilance and decrease in social approach, while OXT infusion into the BNST induced social vigilance in both stress-naïve female and male California mice”
  • Previous work in California mice shows changes in BNST drive stress-induced social avoidance and social vigilance.

    “Previous work in California mice (Peromyscus californicus) shows that social defeat stress has stronger effects on BNST structure and function in adult females than males (37, 38) and that these changes drive stress-induced social avoidance and social vigilance (39–41).”
  • Oxytocin neurons in the bed nucleus of the stria terminalis of California mice make local connections within the bed nucleus of the stria terminalis, project to the hypothalamus, and are involved in social vigilance.

    “However, OXT neurons in the BNST of California mice make local connections within the BNST, project to the hypothalamus, and are involved in social vigilance (66).”
  • In California mice, knockdown of oxytocin or administration of an OXTR antagonist in the anterior bed nucleus of the stria terminalis increased social approach and decreased social vigilance, an anxiety-like behavior, suggesting that oxytocin may also drive social anxiety-like behavior depending on its source or context.

    “However, in California mice, knockdown of OXT or administration of OXTR antagonist in the anterior BNST increased social approach and decreased social vigilance, an anxiety-like behavior, which suggests that OXT may also drive social anxiety-like behavior depending on its source or context (66, 88).”

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.

  • Morpholino dose is reported as both a concentration and a much larger administered amount

    Major

    The methods specify a 0.2 μL injection of 100 pM morpholino, whereas the supplementary figure describes the same single-injection experiment as 100 pmol. A 0.2 μL injection at 100 pM contains only 0.00002 pmol, not 100 pmol.

  • Supplementary caption names the wrong treatment as preventing the increase

    Minor

    The results say antisense reduced oxytocin-cell number relative to missense. The supplementary caption instead says missense both permitted the stress-induced increase and prevented it.

  • Male testing time conflicts with the stated timing for California-mouse analyses

    Minor

    The methods say all behavioral analyses occurred two weeks after the last defeat, but the male behavioral experiment tested mice 10 minutes after their third defeat.

  • Oxytocin-cell sample size conflicts with the reported t-test degrees of freedom

    Minor

    The figure reports seven mice per group, or 14 total. An independent two-group t-test with those sample sizes has 12 degrees of freedom, not the reported 14.

  • Figure caption identifies oxytocin-positive cells where results identify receptor-positive cells

    Minor

    The results identify the cells quantified in Fig. 3J,K as Oxtr-positive, but the caption calls them Oxt-positive. Those labels refer to different measured markers.

5 of 5

Cites

Flagged first, then by citations

Research works printed but not linked: 1

  1. J. Wahis Oxytocin acts on astrocytes in the central amygdala to promote a positive emotional state. bioRxiv :2020.02.25.963884 (2020). Posted 26 February 2020.

Cited by

Sorted by citations

Authors
Duque-Wilckens, N., Torres, L. Y., Yokoyama, S., Minie, V. A., Tran, A. M., Petkova, S. P., Hao, R., Ramos-Maciel, S., Rios, R. A., Jackson, K., Flores-Ramirez, F. J., Garcia-Carachure, I., Pesavento, P. A., Iñiguez, S. D., Grinevich, V., & Trainor, B. C.
Affiliations
  • Department of Psychology, University of California, Davis, CA 95616;
  • University of California, Davis (US)
  • Department of Psychology, The University of Texas at El Paso, El Paso, TX 79902;
  • The University of Texas at El Paso (US)
  • Heidelberg University (DE)
Venue
Proceedings of the National Academy of Sciences
Published
2020-10-05
Volume
117
Issue
42
Pages
26406-26413
ISSN
  • 0027-8424 (ISSN-L)
  • 1091-6490
OpenAlex
W3091710413
Cited by
139
References
86
Citations by year
2026: 21 · 2025: 29 · 2024: 17 · 2023: 20 · 2022: 17 · 2021: 29 · 2020: 6
FWCI
19.70
Type
journal article
Language
en
Open access
Locations
  • Proceedings of the National Academy of Sciences (publishedVersion)
  • bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc-nd)
  • PubMed Central (publishedVersion)
  • PubMed (publishedVersion)
  • eScholarship (California Digital Library) (submittedVersion)
Funders
  • HHS | NIH | National Institute of Mental Health (R01 MH121829-01)
  • HHS | NIH | National Institute of General Medical Sciences (SC3GM130467)
  • Deutsche Forschungsgemeinschaft (Collaborative Research Center (SFB) 1158, GR 3619/701, GR 3619/4-1)
  • SFB (1158)
  • SNSF-DFG (GR 3619/8-1)
  • Fritz Thyssen Stiftung (10.16.2.018 MN)
Topics
  • Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
  • Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuropeptides and Animal Physiology
  • Life Sciences › Neuroscience › Behavioral Neuroscience › Stress Responses and Cortisol
Keywords
oxytocin neurons, social stress, social vigilance, social avoidance, social anxiety, lateral hypothalamus, anterior hypothalamus, viral tracing, defensive behavior
MeSH terms
Animals, Anxiety, Avoidance Learning, Brain, Brain Mapping, Female, Hypothalamus, Male, Neurons, Oxytocin, Peromyscus, Septal Nuclei, Social Behavior, Stress, Psychological, Receptors, Oxytocin, Mice

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