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Cited by 131Neuroendocrine regulation and behaviorNeurotransmitter Receptor Influence on BehaviorEvolutionary Psychology and Human Behavior

Variation in the β-endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality

Open access
Pearce, Eiluned, Wlodarski, Rafael, Machin, Anna, Dunbar, Robin I. M.·2017·Proceedings of the National Academy of Sciences
DOI 10.1073/pnas.1700712114PMC PMC5441808
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Abstract

Significance: Social behavior in mammals is underpinned by a number of social neuropeptides. Most studies, however, focus on a single neuropeptide (often oxytocin), and invariably only in the context of reproductive relationships. Here, we examine the associations between the six main social neuropeptides (endorphins, oxytocin, vasopressin, dopamine, serotonin, and testosterone) and social indices in three separate social domains (disposition, dyadic/romantic relationships, and social network). We show that each neuropeptide is quite specific in its domain of influence, although endorphins and dopamine influence all three social domains. The results suggest that the social importance of endorphins may have been underestimated.

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Written from the paper’s full text.

TL;DR

Using 33 candidate SNPs across nine genes in 757 white British adults, this study tests associations between six social neurochemical systems and three separate domains of sociality (social disposition, dyadic relationships, and wider social network). It finds that β-endorphin, oxytocin, and dopamine each map predominantly onto a different social domain, with endorphins and dopamine showing broad effects across all three domains while oxytocin's influence is largely confined to romantic/reproductive relationships.

Key findings

  • The distribution of significant SNP–behavior associations across neuropeptides and domains was highly nonrandom (χ² = 51.7, df = 10, P < 0.0001), with disproportionately high frequencies for endorphins, oxytocin, and dopamine; vasopressin, serotonin, and testosterone played only limited roles.
  • Domain specificity differed by neuropeptide: partitioning χ² gave nonsignificant (i.e., random) distributions across the three domains for endorphins (χ² = 4.7, df = 2, P = 0.097) and dopamine (χ² = 0.7, df = 2, P = 0.716), but a significantly nonrandom distribution for oxytocin (χ² = 7.5, df = 2, P = 0.023).
  • Percentage of SNPs significantly associated with each domain (heat map): β-endorphin 30.0% disposition / 20.0% dyadic / 10.0% network; oxytocin 4.0% / 45.0% / 10.0%; dopamine 12.5% / 37.5% / 50.0%; testosterone 25.0% / 0.0% / 0.0%; vasopressin 12.5% / 0.0% / 0.0%; serotonin 0.0% / 0.0% / 25.0%.
  • Seven of the 10 OXTR SNPs were significantly associated with SOI-R scores (e.g., rs237887 add×sex P = 0.014; rs53576 domdev P = 0.035), and OXTR SNPs remained significant independently of DRD1 rs265981. OPRM1 SNPs were linked to anxious attachment (rs3778151 geno_2df P = 0.025) and empathy (rs3778151 domdev P = 0.025 for EQ; add×sex P < 0.001 for RMET), and remained significant when controlling for OXTR/AR variants. Network size was associated with OXTR rs237887 (P = 0.025), DRD2 rs4648317 (add×sex P = 0.006), and HTR1a rs6295 (domdev×sex P = 0.012).
  • Fewer than 18% of possible gene–behavior relationships were significant, and many did not survive controlling for other neuropeptides; the authors note that oxytocin's significance often disappears when controlling for endorphins, even for romantic relationships.

Methods

  • Cross-sectional candidate-gene association study of 757 individuals (423 female), aged 18–65 (mean 40.6 years), self-identified Caucasian, physically healthy, not using recreational drugs or drug replacement therapy, and not taking medication for a psychological condition; ethics approval from the University of Oxford Central Ethics Committee.
  • Saliva DNA (OrageneDNA kit, Kleargene extraction) was genotyped for 33 candidate SNPs across nine genes (OXTR, AVPR1a, OPRM1, DRD1, DRD2, ANKK1, HTR1A, HTR2A, AR); after LD pruning and exclusion of low-minor-allele-frequency SNPs, the final set comprised 10 OXTR, 2 AVPR1a, 5 OPRM1, 1 AR, 1 DRD1, 2 DRD2, 1 ANKK1, and the HTR1a and HTR2a SNPs.
  • Self-report measures indexed three domains: social disposition (RMET, EQ, Experiences of Close Relationships scale), dyadic relationship quality (SOI-R, RAS), and wider network engagement (number of close friends and family, Aron's Inclusion of Other in Self scale). Analysis used PLINK v1.9 with additive, dominance, and genotypic (2df) models plus age, sex, and sex×genotype interactions, an optimized permutation test for multiple testing, and post hoc models controlling for other significantly associated SNPs.

Limitations

  • The paper contains no dedicated limitations section; the authors do state that running a large number of statistical tests risks inflating significance levels, which they address with an optimized permutation test they describe as generally conservative.
  • The authors note that, for rs1076560 and rs1800497, smaller numbers of minor allele homozygotes mean greater variance for that group than for major-allele carriers (stated in the text).
  • Evident from the text but not framed as a limitation by the authors: the sample is restricted to white British, physically healthy, non-drug-using adults without treated psychological conditions, and the design is a candidate-gene association study of 33 SNPs, so the findings are correlational and of uncertain generalizability.

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Integrity signals

Registry

Crossref · 10.1073/pnas.1700712114

Cites

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Statistics

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Claims the paper makes

0 of 60 cited sources checked · 0 supported · 0 flagged

  • Findingown data
    § Results · p. 5

    There were significant associations (P < 0.05) between SNPs and all of the sociality measures across the three domains.

    “There were significant associations ( P &lt; 0.05) between SNPs and all of the sociality measures across the three domains ( SI Appendix , Table S2).”
    Quote in the paperSubstantial claim
  • Findingown data
    § Results · p. 5

    OPRM1 rs3778151 had a significant nonlinear relationship with Empathy Quotient (EQ) (domdev, P = 0.025), with the minor allele having lower EQ scores than expected from an additive model.

    “OPRM1 rs3778151 was found to have a significant nonlinear relationship with Empathy Quotient (EQ) ( domdev , P = 0.025), with the negative effect suggesting that the minor allele has lower EQ scores than expected from an additive model.”
    Quote in the paperNot a substantial claim
  • Findingown data
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Three neuropeptides (β-endorphin, oxytocin, and dopamine) play particularly important roles in human sociality.

    “We use variation in 33 SNPs for the receptor genes for six well-known social neuropeptides in relation to three separate domains of sociality (social disposition, dyadic relationships, and social networks) to show that three neuropeptides ( β -endorphin, oxytocin, and dopamine) play particularly important roles, with each being associated predominantly with a different social domain.”
    Quote in the paperSubstantial claim
  • Findingown data
    § Results · p. 5

    OPRM1 rs3778151 had a significant additive effect with sex on Reading the Mind in the Eyes Test (RMET) scores (add x sex, P < 0.001).

    “OPRM1 rs3778151 also had a significant additive effect with sex on Reading the Mind in the Eyes Test (RMET) scores ( add x sex , P &lt; 0.001)”
    Quote in the paperSubstantial claim
  • Findingown data
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Vasopressin, serotonin, and testosterone play only limited roles in human sociality.

    “In contrast, vasopressin, serotonin, and testosterone play only limited roles.”
    Quote in the paperSubstantial claim
  • May combine two claims. 26
  • Classified as probably not a substantial claim. 30

  • May combine two claims.

    Findingown data
    § Results · p. 6

    Partitioning χ2 yields nonsignificant (i.e., random) distributions across the three social domains for endorphins (χ2 = 4.7, df = 2, P = 0.097) and dopamine (χ2 = 0.7, df = 2, P = 0.716).

    “Partitioning χ 2 for these three neuropeptides yields nonsignificant (i.e., random) distributions across the three social domains for endorphins ( χ 2 = 4.7, df = 2, P = 0.097) and dopamine ( χ 2 = 0.7, df = 2, P = 0.716), but a significantly nonrandom distribution for oxytocin ( χ 2 = 7.5, df = 2, P = 0.023).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data
    § Results · p. 5

    OPRM1 SNPs remained significantly associated with RAS independently of the OXTR SNPs, but the opposite was not true.

    “The OPRM1 SNPs remained significantly associated with RAS independently of the OXTR SNPs, but the opposite was not true.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data
    § Results · p. 5

    DRD1 rs265981 interacted with sex and was linked to SOI-R scores (domdev x sex, P = 0.039).

    “DRD1 rs265981 was also found to interact with sex and be linked to SOI-R scores ( domdev x sex , P = 0.039).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data
    § Results · p. 5

    Feelings of connection to local community (IOS) had significant associations with OPRM1 rs1799971 (domdev, P = 0.027), OXTR rs53576 (domdev, P = 0.025), DRD2 rs1076560 (add x sex, P = 0.029), and ANKK1 rs1800497 (add x sex, P = 0.029).

    “For feelings of connection to their local community [measured using Aron ' s Inclusion of Other in Self (IOS) scale] (49), significant associations were found for OPRM1 rs1799971 ( domdev , P = 0.027), OXTR rs53576 ( domdev , P = 0.025), and DRD2 rs1076560 ( add x sexp = 0.029, Fig. 1) and ANNK1 rs1800497 (downstream from DRD2 ; add x sex , P = 0.029).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data
    § Results · p. 6

    Oxytocin is very specific in its functional domain (romantic relationships), whereas endorphins and dopamine both have broad spectrum effects.

    “In sum, oxytocin is very specific in its functional domain (romantic relationships), whereas endorphins and dopamine both have broad spectrum effects.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingnone given
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Oxytocin's effects are confined to romantic/reproductive relationships and often do not survive control for other neuropeptides.

    “However, endorphins and dopamine have a much wider compass than oxytocin (whose effects are confined to romantic/reproductive relationships and often do not survive control for other neuropeptides).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data
    § Results · p. 6

    The distribution of significant effects shown in Fig. 2 is highly nonrandom (χ2 = 51.7, df = 10, P < 0.0001), with disproportionately higher frequencies than expected for endorphins, oxytocin, and dopamine.

    “The distribution of significant effects shown in Fig. 2 is highly nonrandom ( χ 2 = 51.7, df = 10, P &lt; 0.0001), with disproportionately higher frequencies than expected for endorphins, oxytocin, and dopamine.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data
    § Significance · p. 3

    Each neuropeptide is quite specific in its domain of influence, although endorphins and dopamine influence all three social domains.

    “We show that each neuropeptide is quite specific in its domain of influence, although endorphins and dopamine influence all three social domains.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data
    § Discussion · p. 6

    OPRM1 variations were most strongly linked to individual differences in attachment style, the ability to identify emotional facial expressions (RMET), and a self-report measure of empathy (EQ).

    “OPRM1 variations were most strongly linked to individual differences in attachment style and the ability to identify emotional facial expressions (RMET), as well as a selfreport measure of empathy (EQ).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • May combine two claims.

    Findingown data
    § Discussion · p. 6

    Anxious attachment, but not avoidant attachment, is associated with variation in OPRM1, AR, and OXTR.

    “Our results suggest that anxious, but not avoidant, attachment is associated with OPRM1 , AR , and OXTR variation.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • May combine two claims.

    Findingown data
    § Discussion · p. 6

    Both measures of empathy (RMET and EQ) were related to OPRM1 genotypes.

    “Both measures of empathy were related to OPRM1 genotypes”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • May combine two claims.

    Findingown data
    § Discussion · p. 7

    One of the OPRM1 SNPs and DRD2 variation correlate with degree of community integration.

    “one of the OPRM1 SNPs and DRD2 variation correlates with degree of community integration”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • May combine two claims.

    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Most social neuroscience concentrates on a very narrow spectrum of social behavior or has used meaninglessly broad definitions of sociality.

    “Most social neuroscience suffers from a second important limitation: It invariably concentrates on a very narrow spectrum of social behavior (mainly romantic and parent -child bonding, empathy, and dyadic interactions in economic games) or has used meaninglessly broad definitions of sociality (as in a recent metaanalysis that failed to find a significant association between the two most-studied OX”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [24] Bakermans-Kranenburg MJ, van Ijzendoorn MH (2014) A sociability gene? Metaanalysis of oxytocin receptor genotype effects in humans. Psychiatr Genet 24:45 -51.

      Not checked in the free run.

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

    Premisenone given
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    The impact of genetic variation on individual differences in social behavior and cognition occurs mainly through altering the function of a key set of neuropeptides.

    “The impact of genetic variation on individual differences in social behavior and cognition occurs mainly through altering the function of a key set of neuropeptides.”
    Quote in the paperNot a substantial claim

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

    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    The number and quality of social relationships have substantial impacts on health, well-being, and longevity, and, at least in animals, on reproductive fitness.

    “There is growing evidence that the number and quality of social relationships have substantial impacts on health, well-being, and longevity, and, at least in animals, on reproductive fitness.”
    Quote in the paperNot a substantial claim0 of 4 sources checked
    • cites [1] source

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    • cites [4]

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

    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Oxytocin may be a universal feature of mammalian sociality, but primates' unusual lifelong bonded relationships require a more robust neurochemical underpinning, and the endorphin system was exapted for this purpose.

    “It has specifically been argued that, whereas oxytocin may be a universal feature of mammalian sociality, primates ' unusual lifelong bonded relationships requires a more robust neurochemical underpinning, and the endorphin system was exapted for this purpose (23).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [23] Broad KD, Curley JP, Keverne EB (2006) Mother-infant bonding and the evolution of mammalian social relationships. Philos Trans R Soc Lond B Biol Sci 361:2199 -2214. source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Primates are characterized by multilayered social groups in which individuals are embedded in an inner core of intense relationships, which in turn are embedded in larger, more loosely structured social networks.

    “Primates, in particular, are characterized by multilayered social groups in which individuals are embedded in an inner core of intense relationships, which in turn are embedded in larger, more loosely structured social networks (25, 26).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [25] Kudo H, Dunbar RIM (2001) Neocortex size and social network size in primates. Anim Behav 62:711 -722.

      Not checked in the free run.

    • cites [26] Hill RA, Bentley RA, Dunbar RIM (2008) Network scaling reveals consistent fractal pattern in hierarchical mammalian societies. Biol Lett 4:748 -751.

      Not checked in the free run.

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

    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    Variation in OXTR and AVPR1a alleles has been associated with individual capacities for affective and cognitive empathy, respectively.

    “and variation in OXTR and AVPR1a alleles has been associated with individual capacities for affective and cognitive empathy, respectively (30).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [30] Uzefovsky F, et al. (2015) Oxytocin receptor and vasopressin receptor 1a genes are respectively associated with emotional and cognitive empathy. Horm Behav 67:60 -65. source

      Not checked in the free run.

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    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    The μ-opioid receptor is the target of β-endorphin, which seems to play an important role in maintaining social bonds in primates (including humans).

    “The μ -opioid receptor (16) has been of particular interest because it is the target of β -endorphin, which seems to play an important role in maintaining social bonds in primates (including humans) (17 -19).”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [16] Inoue K, Burkett JP, Young LJ (2013) Neuroanatomical distribution of μ -opioid receptor mRNA and binding in monogamous prairie voles ( Microtus ochrogaster ) and non-monogamous meadow voles ( Microtus pennsylvanicus ). Neuroscience 244: 122 -133.

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    • cites [17] Fabre-Nys C, Meller RE, Keverne EB (1982) Opiate antagonists stimulate affiliative behaviour in monkeys. Pharmacol Biochem Behav 16:653 -659. source

      Not checked in the free run.

    • cites [18] Keverne EB, Martensz ND, Tuite B (1989) Beta-endorphin concentrations in cerebrospinal fluid of monkeys are influenced by grooming relationships. Psychoneuroendocrinology 14:155 -161. source

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    • cites [19] Machin AJ, Dunbar RIM (2011) The brain opioid theory of social attachment: A review of the evidence. Behaviour 148:985 -1025. source

      Not checked in the free run.

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    Premisecitation
    § Variation in the β -endorphin, oxytocin, and dopamine receptor genes is associated with different dimensions of human sociality · p. 3

    β-Endorphin is a likely candidate in the maintenance of long-term relationships in humans because, unlike oxytocin, it can underpin relationships outside of the context of intercourse or parturition and operates effectively at both the dyadic and group levels.

    “β -Endorphin is a likely candidate in the maintenance of long-term relationships in humans because, unlike oxytocin, it can underpin relationships outside of the context of intercourse or parturition and operates effectively at both the dyadic and group levels (20 -22).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [20] Nummenmaa L, et al. (2016) Social touch modulates endogenous μ -opioid system activity in humans. Neuroimage 138:242 -247.

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    • cites [21] Weinstein D, Launay J, Pearce E, Dunbar RI, Stewart L (2016) Group music performance causes elevated pain thresholds and social bonding in small and large groups of singers. Evol Hum Behav 37:152 -158.

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    • cites [22] Dunbar RIM, et al. (2012) Social laughter is correlated with an elevated pain threshold. Proc Biol Sci 279:1161 -1167.

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    Premisecitation
    § Significance · p. 4

    DRD1 and DRD2 genotypes have been linked to age at first sexual intercourse.

    “and DRD1 and DRD2 genotypes have been linked to age at first sexual intercourse (40)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [40] Miller WB, et al. (1999) Dopamine receptor genes are associated with age at first sexual intercourse. J Biosoc Sci 31:43 -54.

      Not checked in the free run.

  • May combine two claims.

    Interpretationcitation
    § Discussion · p. 6

    The neurochemical mechanism underpinning long-term relationships is the endogenous opioid system rather than oxytocin.

    “the neurochemical mechanism underpinning long-term relationships is the enodgenous opioid system rather than oxytocin”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [35] Schneiderman I, Kanat-Maymon Y, Ebstein RP, Feldman R (2014) Cumulative risk on the oxytocin receptor gene (OXTR) underpins empathic communication difficulties at the first stages of romantic love. Soc Cogn Affect Neurosci 9:1524 -1529.

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    • cites [51] Dunbar RIM (2010) The social role of touch in humans and primates: Behavioural function and neurobiological mechanisms. Neurosci Biobehav Rev 34:260 -268. source

      Not checked in the free run.

    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • Classified as probably not a substantial claim.

    Study design and conductcitation
    § Results · p. 6

    We used an optimized permutation test that is generally considered to be conservative to control for multiple comparisons.

    “Running a large number of statistical tests risks inflating significance levels; however, we used an optimized permutation test that is generally considered to be conservative ( SI Appendix ).”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Interpretationreasoning
    § Discussion · p. 6

    The association of OPRM1 variation with dyadic relationship quality and community integration may reflect the way social dispositions affect these higher-level outcomes.

    “which may reflect the way social dispositions affect these higher-level outcomes”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • Classified as probably not a substantial claim.

    Premisecitation
    § Significance · p. 5

    Variation in OXTR can predict the size and diversity of social networks via its influence on social temperament.

    “What little work has been done suggests that variation in OXTR can predict the size and diversity of social networks via its influence on social temperament (45)”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [45] Creswell KG, et al. (2015) OXTR polymorphism predicts social relationships through its effects on social temperament. Soc Cogn Affect Neurosci 10:869 -876.

      Not checked in the free run.

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

    Premisecitation
    § Significance · p. 4

    HTR1A variation is associated with romantic relationship formation.

    “whereas HTR1A variation is associated with romantic relationship formation (41).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [41] Liu J, Gong P, Zhou X (2014) The association between romantic relationship status and 5-HT1A gene in young adults. Sci Rep 4:7049.

      Not checked in the free run.

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

    Study design and conductown data
    § Results · p. 5

    Due to smaller numbers of minor allele homozygotes, the variance for this group is greater than for carriers of the major allele for both rs1076560 and rs1800497.

    “It should be noted that due to smaller numbers of minor allele homozygotes, the variance for this group is greater than for carriers of the major allele for both rs1076560 (see Fig. 1) and rs1800497.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Premisecitation
    § Significance · p. 5

    OXTR variation is related to feelings of social connectedness and 'general sociality'.

    “as well as being related to feelings of social connectedness (46) and ' general sociality ' (47).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [46] Chang SC, et al. (2014) Are genetic variations in OXTR, AVPR1A, and CD38 genes important to social integration? Results from two large U.S. cohorts. Psychoneuroendocrinology 39:257 -268.

      Not checked in the free run.

    • cites [47] Li J, et al. (2015) Association of oxytocin receptor gene (OXTR) rs53576 polymorphism with sociality: A meta-analysis. PLoS One 10:e0131820.

      Not checked in the free run.

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

    Premisecitation
    § Significance · p. 4

    AR variation has been associated with relationship status and testosterone response in opposite-sex interactions.

    “Furthermore, AR variation has been associated with relationship status and testosterone response in opposite-sex interactions (43, 44).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [43] Zitzmann M, Depenbusch M, Gromoll J, Nieschlag E (2004) X-chromosome inactivation patterns and androgen receptor functionality influence phenotype and social characteristics as well as pharmacogenetics of testosterone therapy in Klinefelter patients. J Clin Endocrinol Metab 89:6208 -6217.

      Not checked in the free run.

    • cites [44] Roney JR, Simmons ZL, Lukaszewski AW (2010) Androgen receptor gene sequence and basal cortisol concentrations predict men ' s hormonal responses to potential mates. Proc Biol Sci 277:57 -63.

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing). May combine two claims.

    Interpretationown data
    § Discussion · p. 6

    An individual's ability to empathize and their attachment style likely feed into their romantic relationships.

    “An individual ' s ability to empathize, and their typical way of relating in close relationships (their attachment style), is also likely to feed into their romantic relationships”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

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

    Premisecitation
    § Significance · p. 4

    OPRM1 and HTR2A have been associated with variation in mate-choice success differentially in the sexes in 'speed-dating' situations.

    “In addition, OPRM1 and HTR2A have been associated with variation in mate-choice success differentially in the sexes in ' speed-dating ' situations (42).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [42] Wu K, Chen C, Moyzis RK, Greenberger E, Yu Z (2016) Gender interacts with opioid receptor polymorphism A118G and serotonin receptor polymorphism -1438 A/G on speed-dating success. Hum Nat 27:244 -260.

      Not checked in the free run.

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

    Premisecitation
    § Significance · p. 4

    DRD2 and HTR2A variation has been linked to different 'loving styles'.

    “In humans, DRD2 and HTR2A variation has been linked to different ' loving styles ' (39)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [39] Emanuele E, Brondino N, Pesenti S, Re S, Geroldi D (2007) Genetic loading on human loving styles. Neuroendocrinol Lett 28:815 -821.

      Not checked in the free run.

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

    Study design and conductcitation
    § Results · p. 6

    These data are not subject to multiple comparisons effects because the P value is simply a criterion for a binary partition of the dataset.

    “These data are not subject to multiple comparisons effects because the P value is simply a criterion for a binary partition of the dataset ( SI Appendix ).”
    Quote in the paperNot a substantial claim

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

    Premisecitation
    § Significance · p. 5

    Other studies have provided indirect evidence for a relationship between endorphins and social network size.

    “Other studies have, however, provided indirect evidence for a relationship between endorphins and social network size (48).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [48] Johnson KVA, Dunbar RIM (2016) Pain tolerance predicts human social network size. Sci Rep 6:25267.

      Not checked in the free run.

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

    Premisecitation
    § Discussion · p. 6

    The density of μ-opioid receptors has previously been found to be related to attachment style, mainly with the avoidance dimension.

    “The density of μ -opioid receptors has previously been found to be related to attachment style, though mainly with the avoidance dimension (32).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [32] Nummenmaa L, et al. (2015) Adult attachment style is associated with cerebral μ -opioid receptor availability in humans. Hum Brain Mapp 36:3621 -3628.

      Not checked in the free run.

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

    Premisecitation
    § Significance · p. 4

    Both the dopamine and opioid systems have been shown to interact to maintain monogamous pair bonds in prairie voles.

    “For instance, both the dopamine and opioid systems have been shown to interact to maintain monogamous pair bonds in prairie voles (15, 38).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [15] Resendez SL, et al. (2013) μ -Opioid receptors within subregions of the striatum mediate pair bond formation through parallel yet distinct reward mechanisms. J Neurosci 33: 9140 -9149. source

      Not checked in the free run.

    • cites [38] Resendez SL, et al. (2016) Dopamine and opioid systems interact within the nucleus accumbens to maintain monogamous pair bonds. eLife 5:15325. source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation
    § Significance · p. 4

    Romantic relationship status, relationship quality, and sexual behaviors have been linked to OXTR variation in women and AVPR1a variation in men.

    “Romantic relationship status, relationship quality, and sexual behaviors have been linked to OXTR variation in women and AVPR1a variation in men, although this sexual division is not always straighforward (12, 35 -37).”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [12] Donaldson ZR, Young LJ (2008) Oxytocin, vasopressin, and the neurogenetics of sociality. Science 322:900 -904. source

      Not checked in the free run.

    • cites [35] Schneiderman I, Kanat-Maymon Y, Ebstein RP, Feldman R (2014) Cumulative risk on the oxytocin receptor gene (OXTR) underpins empathic communication difficulties at the first stages of romantic love. Soc Cogn Affect Neurosci 9:1524 -1529.

      Not checked in the free run.

    • cites [36] Walum H, et al. (2012) Variation in the oxytocin receptor gene is associated with pairbonding and social behavior. Biol Psychiatry 71:419 -426. source

      Not checked in the free run.

    • cites [37] Walum H, et al. (2008) Genetic variation in the vasopressin receptor 1a gene (AVPR1A) associates with pair-bonding behavior in humans. Proc Natl Acad Sci USA 105: 14153 -14156. source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation
    § Discussion · p. 6

    Prior to this study, dopamine function had never been studied in relation to social interaction beyond the dyad; past research showed links only to romantic relationships or parenting.

    “To our knowledge, dopamine function has never before been studied in relation to social interaction beyond the dyad, with past research showing links only to romantic relationships or parenting (15, 39, 40).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [15] Resendez SL, et al. (2013) μ -Opioid receptors within subregions of the striatum mediate pair bond formation through parallel yet distinct reward mechanisms. J Neurosci 33: 9140 -9149. source

      Not checked in the free run.

    • cites [39] Emanuele E, Brondino N, Pesenti S, Re S, Geroldi D (2007) Genetic loading on human loving styles. Neuroendocrinol Lett 28:815 -821.

      Not checked in the free run.

    • cites [40] Miller WB, et al. (1999) Dopamine receptor genes are associated with age at first sexual intercourse. J Biosoc Sci 31:43 -54.

      Not checked in the free run.

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

    Premisecitation
    § Discussion · p. 6

    Much previous work on β-endorphin has focused on 'social pain' resulting from social rejection.

    “much previous work on β -endorphin has focused on ' social pain ' resulting from social rejection (53, 54)”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [53] Way BM, Taylor SE, Eisenberger NI (2009) Variation in the mu-opioid receptor gene (OPRM1) is associated with dispositional and neural sensitivity to social rejection. Proc Natl Acad Sci USA 106:15079 -15084. source

      Not checked in the free run.

    • cites [54] Bonenberger M, Plener PL, Groschwitz RC, Grön G, Abler B (2015) Polymorphism in the μ -opioid receptor gene (OPRM1) modulates neural processing of physical pain, social rejection and error processing. Exp Brain Res 233:2517 -2526.

      Not checked in the free run.

  • May combine two claims.

    Interpretationcitation
    § Discussion · p. 6

    The vasopressin association with RAS confirms an earlier finding that vasopressin is predictive of high levels of promiscuity.

    “confirming an earlier finding that it is predictive of high levels of promiscuity (39)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [39] Emanuele E, Brondino N, Pesenti S, Re S, Geroldi D (2007) Genetic loading on human loving styles. Neuroendocrinol Lett 28:815 -821.

      Not checked in the free run.

    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

  • Classified as probably not a substantial claim.

    Study design and conductown data
    § Materials and Methods · p. 7

    To test the independence of associations between particular SNPs and specific social variables, we ran post hoc models that controlled for the effect of additional SNPs that also showed significant associations with the same social variable.

    “To test the independence of associations between particular SNPs and specific social variables, we ran post hoc models that controlled for the effect of additional SNPs that also showed significant associations with the same social variable.”
    Quote in the paperSubstantial claim

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

    Premisecitation
    § Discussion · p. 7

    Poor relationships and network disintegration lead to morbidity and mortality risks even greater than those of alcohol abuse, lack of exercise, and smoking.

    “poor relationships and network disintegration lead to morbidity and mortality risks even greater than those of alcohol abuse, lack of exercise. and smoking (1)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [1] source

      Not checked in the free run.

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

    Interpretationcitation
    § Discussion · p. 7

    The correlation of one OPRM1 SNP and DRD2 variation with community integration may reflect known interactions between the dopamine and endorphin systems in both humans and rodents, whereby endorphin activation may itself trigger a dopamine response.

    “may reflect known interactions between the dopamine and endorphin systems in both humans and rodents (40, 55, 56), whereby endorphin activation may itself trigger a dopamine response”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites [40] Miller WB, et al. (1999) Dopamine receptor genes are associated with age at first sexual intercourse. J Biosoc Sci 31:43 -54.

      Not checked in the free run.

    • cites [55] Peciña M, Love T, Stohler CS, Goldman D, Zubieta J-K (2015) Effects of the Mu opioid receptor polymorphism (OPRM1 A118G) on pain regulation, placebo effects and associated personality trait measures. Neuropsychopharmacology 40:957 -965.

      Not checked in the free run.

    • cites [56] Devine DP, Leone P, Pocock D, Wise RA (1993) Differential involvement of ventral tegmental mu, delta and kappa opioid receptors in modulation of basal mesolimbic dopamine release: In vivo microdialysis studies. J Pharmacol Exp Ther 266:1236 -1246. source

      Not checked in the free run.

    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

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

    Study design and conductnone given
    § Materials and Methods · p. 7

    Participants with less than 90% coverage were removed (11 in total).

    “Participants with less than 90% coverage were removed (11 in total).”
    Quote in the paperNot a substantial claim

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

    Study design and conductown data
    § Materials and Methods · p. 7

    Only 4 of the SNPs differed significantly from Hardy-Weinberg equilibrium in the full sample.

    “Only 4 of the SNPs differed significantly from Hardy -Weinberg equilibrium in the full sample”
    Quote in the paperNot a substantial claim

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

    Study design and conductown data
    § Materials and Methods · p. 7

    All of the SNPs had at least 95% coverage.

    “All of the SNPs had at least 95% coverage.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Study design and conductnone given
    § Materials and Methods · p. 7

    We used an optimized permutation test to control for multiple testing.

    “We used an optimized permutation test to control for multiple testing.”
    Quote in the paperSubstantial claim

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

    Premisecitation
    § Discussion · p. 6

    μ-opioid receptors are activated by touch by a romantic partner.

    “the μ -opioid receptors are activated by touch by a romantic partner (20)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [20] Nummenmaa L, et al. (2016) Social touch modulates endogenous μ -opioid system activity in humans. Neuroimage 138:242 -247.

      Not checked in the free run.

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

    Premisecitation
    § Discussion · p. 7

    Network size and relationship quality are known to have significant impacts on reproduction and fitness in primates and other mammals.

    “network size and relationship quality are known to have significant impacts on reproduction and fitness in primates and other mammals (8 -10)”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites [8] source

      Not checked in the free run.

    • cites [9]

      Not checked in the free run.

    • cites [10] source

      Not checked in the free run.

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

    Premisecitation
    § Discussion · p. 6

    In synchronous group activities, endorphin release is thought to play a key part in bonding group members.

    “in synchronous group activities, endorphin release is thought to play a key part in bonding group members (21, 22)”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [21] Weinstein D, Launay J, Pearce E, Dunbar RI, Stewart L (2016) Group music performance causes elevated pain thresholds and social bonding in small and large groups of singers. Evol Hum Behav 37:152 -158.

      Not checked in the free run.

    • cites [22] Dunbar RIM, et al. (2012) Social laughter is correlated with an elevated pain threshold. Proc Biol Sci 279:1161 -1167.

      Not checked in the free run.

  • May combine two claims.

    Interpretationcitation
    § Discussion · p. 6

    OPRM1 variation is linked to the extent to which individuals experience social rewards rather than to pain per se.

    “OPRM1 variation is linked to the extent to which individuals experience social rewards (31) rather than to pain per se”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [31] Troisi A, et al. (2011) Social hedonic capacity is associated with the A118G polymorphism of the mu-opioid receptor gene (OPRM1) in adult healthy volunteers and psychiatric patients. Soc Neurosci 6:88 -97.

      Not checked in the free run.

    • Rests on an assumption: The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.
    • Rests on an assumption: The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.
    • Rests on an assumption: The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.
51 of 51

  • The authors state this as an assumption.

    Sociality can be meaningfully divided into three separate domains: social disposition, dyadic relationships, and social networks.

    “An important distinction needs to be drawn between social dispositions, intimate dyadic relationships, and the wider social network within which the individual is embedded.”

  • The paper relies on this without support.

    The behavioral measures used (e.g., RAS, SOI-R, EQ, RMET, IOS, network size) validly index the three social domains.

    “Details of the measures used to index these variables are given in Materials and Methods and SI Appendix .”

  • The paper relies on this without support.

    The role of neuropeptides in the wider social context beyond the dyad has been almost totally ignored.

    “By comparison, with work on dyadic relationships, the role of neuropeptides in the wider social context beyond the dyad has been almost totally ignored.”

  • The paper relies on this without support.

    Little has been done to examine the influence of other neuropeptides on wider social network.

    “Aside from these, little has been done to examine the influence of other neuropeptides on wider social network.”

  • The paper relies on this without support.

    The sample of 757 Caucasian, physically healthy individuals is genetically homogeneous enough that the observed associations are not due to population stratification.

    “The final sample consisted of 757 individuals (423 female) aged 18 -65 (mean 40.6) years who identified themselves as Caucasian, were physically healthy, did not use recreational drugs, were not on drug replacement therapy, or taking medication for a psychological condition.”

  • The paper relies on this without support.

    The self-report questionnaires (RMET, EQ, ECR, SOI-R, RAS, number of close friends/family, IOS) validly measure the intended sociality constructs.

    “Participants completed self-report questionnaires presented digitally on mobile devices. These included: the RMET, the EQ scale, and the Experiences of Close Relationships scale as indices of social disposition; the SOI-R and the RAS as indices of dyadic relationship quality; and the number of close friends and family index and the IOS scale as indices of wider network engagement.”

  • The paper relies on this without support.

    The 33 candidate SNPs are in linkage disequilibrium with functional variants that affect the relevant neuropeptide systems.

    “We genotyped 33 candidate SNPs from nine genes ( SI Appendix , Table S1).”

  • The paper relies on this without support.

    The social domains (attachment, empathy, dyadic relationship quality, wider network engagement) are distinct and can be compared.

    “By comparing different social domains, we provide the insight that variation in genes associated with dopamine receptor 2 may be predominantly linked to engagement in wider social groups.”
8 of 8

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?

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.

  • Vasopressin's reported domain reverses the results

    Major

    The discussion says vasopressin was associated only with RAS, a dyadic-relationship measure. The results table instead shows a disposition association and no dyadic-relationship association.

  • Listed SNP counts sum to 31, not 33

    Minor

    The counts listed for the nine genes total 31 candidate SNPs, although the paragraph says 33 were genotyped.

    Reported 33 · recomputed 31

  • Claim of associations with all measures conflicts with avoidant-attachment result

    Minor

    The results say that no SNP was significantly associated with avoidant attachment, contradicting the preceding claim that significant associations were found for all sociality measures.

  • P = 0.46 is called significant

    Minor

    The reported P value exceeds the paper's stated P < 0.05 criterion for significance.

  • OPRM1 is said to remain significant after controls despite the EQ result

    Minor

    The discussion says only OPRM1 remained significant after controlling for other significant genes, but the EQ results explicitly say its association did not remain significant after controlling for ANKK1.

5 of 5

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Related

Authors
Pearce, E., Wlodarski, R., Machin, A., & Dunbar, R. I. M.
Affiliations
University of Oxford (GB)
ORCID
Venue
Proceedings of the National Academy of Sciences
Publisher
National Academy of Sciences
Published
2017-05
Volume
114
Issue
20
Pages
5300-5305
ISSN
  • 0027-8424 (ISSN-L)
  • 1091-6490
DOI
10.1073/pnas.1700712114
PMC
PMC5441808
OpenAlex
W2611261794
Cited by
131
References
58
Citations by year
2026: 8 · 2025: 15 · 2024: 12 · 2023: 11 · 2022: 14 · 2021: 18 · 2020: 13 · 2019: 14 · 2018: 19 · 2017: 7
FWCI
20.76
Type
journal article
Language
en
Update notices
Open access
Licence
www.pnas.org/site/misc/userlicense.xhtml (vor)
Locations
  • Proceedings of the National Academy of Sciences (publishedVersion)
  • Europe PMC (PubMed Central) (publishedVersion, other-oa)
  • PubMed Central (publishedVersion)
  • PubMed (publishedVersion)
  • Aaltodoc (Aalto University) (submittedVersion, cc-by-nc-nd)
Funders
EC | European Research Council (295663)
Topics
  • Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
  • Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
  • Social Sciences › Psychology › Experimental and Cognitive Psychology › Evolutionary Psychology and Human Behavior
Keywords
beta-endorphin, oxytocin, dopamine, receptor genes, single nucleotide polymorphisms, sociality, social networks, dyads, social dispositions, vasopressin, serotonin, testosterone, interpersonal relationships
MeSH terms
Adolescent, Adult, Aged, beta-Endorphin, Brain, Dopamine, Female, Humans, Male, Middle Aged, Neuropeptides, Oxytocin, Receptors, Dopamine, Receptors, Opioid, Saliva, Social Behavior, Social Support, Receptors, Oxytocin, Social Networking

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 2026-10-10.

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

Statistics and arithmetic are recomputed, not judged.

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