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Cited by 307Neuroendocrine regulation and behaviorNeurotransmitter Receptor Influence on BehaviorAttachment and Relationship Dynamics

The behavioral, anatomical and pharmacological parallels between social attachment, love and addiction

Open access
Burkett, James P., Young, Larry J.·2012·Psychopharmacology
DOI 10.1007/s00213-012-2794-xPMC PMC3469771
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Abstract

Love has long been referred to as an addiction in literature and poetry. Scientists have often made comparisons between social attachment processes and drug addiction, and it has been suggested that the two may share a common neurobiological mechanism. Brain systems that evolved to govern attachments between parents and children and between monogamous partners may be the targets of drugs of abuse and serve as the basis for addiction processes. Here, we review research on drug addiction in parallel with research on social attachments, including parent–offspring attachments and social bonds between mating partners. This review focuses on the brain regions and neurochemicals with the greatest overlap between addiction and attachment and, in particular, the mesolimbic dopamine (DA) pathway. Significant overlap exists between these two behavioral processes. In addition to conceptual overlap in symptomatology, there is a strong commonality between the two domains regarding the roles and sites of action of DA, opioids, and corticotropin-releasing factor. The neuropeptides oxytocin and vasopressin are hypothesized to integrate social information into attachment processes that is not present in drug addiction. Social attachment may be understood as a behavioral addiction, whereby the subject becomes addicted to another individual and the cues that predict social reward. Understandings from both fields may enlighten future research on addiction and attachment processes.

AI snapshot

Written from the paper’s full text.

TL;DR

This review argues that social attachment and love share behavioral, anatomical, and pharmacological parallels with drug addiction, especially within the mesolimbic dopamine pathway. It proposes a “partner addiction” hypothesis in which attachment can be understood as addiction to another individual and the cues predicting social reward.

Key findings

  • Behavioral criteria for substance dependence overlap with normal romantic attachment phenomena such as tolerance, withdrawal, craving, relapse, and neglect of other responsibilities; the authors note that adult pair bonds occur in only 3–5% of mammalian species, whereas up to 90% of bird species are monogamous.
  • Dopamine, opioids, and CRF systems overlap between addiction and attachment: in prairie voles, pair bonding is prevented by DA receptor blockade in the nucleus accumbens shell, D2R activation induces bonding without mating, D1R activation prevents bonding, and cohabitation increases D1R expression in the NAC by 60% relative to same-sex sibling cohabitation.
  • Opioid findings include that MOR in the caudate-putamen is necessary for pair bond formation, KOR in the NAC shell mediates pair bond maintenance/selective aggression, and the human MOR A118G variant is associated with altered attachment style and increased alcohol dependence risk.
  • Oxytocin and vasopressin are proposed to integrate social information into attachment: OT in the NAC is necessary and sufficient for female prairie vole pair bonding and interacts with D2R; human OT is higher during early romantic relationships and predicts which couples stay together after 6 months, while a human AVP V1aR variant is associated with males being half as likely to be married and twice as likely to report major relationship problems.

Methods

  • Narrative review, not a systematic review or meta-analysis, synthesizing animal and human research on drug addiction and social attachment with focus on the mesolimbic dopamine pathway and DA, opioids, CRF, oxytocin, and vasopressin.
  • Animal evidence includes drug self-administration, conditioned place preference, reinstatement, and compulsive drug-seeking paradigms, plus maternal attachment models in sheep, rats, mice, and Rhesus macaques and pair-bonding studies in prairie voles using partner preference and selective aggression.
  • Human parallel evidence includes PET, genetic association, plasma oxytocin, and other human studies; the main analysis is conceptual and neurobiological comparison leading to the partner addiction hypothesis.

Limitations

  • Stated in the paper: extrapolation from animal maternal attachment and pair bonding to human love relies on a debated supposition; complete animal models of addiction are considered impossible by some; not all researchers agree that tolerance and withdrawal represent addiction-related phenomena; sources of CRF projections into the VTA and NAC shell are unknown; CRF action in the NAC during social loss remains a hypothesis to be tested; and there is a relative lack of studies showing that OT antagonists alone affect drug-related processes.
  • Evident from the text: as a narrative review, it does not report a systematic search strategy, quality assessment, or meta-analysis, and many conclusions rest on animal models plus correlational human genetic or physiological findings rather than experimental causal evidence in humans.

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

Integrity signals

Registry

Crossref · 10.1007/s00213-012-2794-x

Cites

401 of 401 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

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  • Interpretationnone given

    Attachment and addiction share strong commonality in the roles and sites of action of dopamine, opioids, and CRF.

    “Significant overlap exists between these two behavioral processes. In addition to conceptual overlap in symptomatology, there is a strong commonality between the two domains regarding the roles and sites of action of dopamine, opioids, and corticotrophin-releasing factor (CRF).”
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  • Interpretationcitation

    Dopamine has a well-validated role in addiction processes.

    “DA has a well-validated role in addiction processes.”
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  • Premisecitation

    Paradigms have been developed to model compulsive drug-seeking despite negative consequences by pairing drug delivery with aversive stimuli.

    “More recently, paradigms have been developed to model compulsive drug-seeking despite negative consequences, which pair drug delivery with various aversive stimuli such as electric shock (Deroche-Gamonet et al 2004), bitter taste (Wolffgramm and Heyne 1995), or fear-conditioned cues (Vanderschuren and Everitt 2004).”
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  • Interpretationreasoning

    There is significant overlap between social attachment and drug addiction in behavior and symptomatology.

    “Significant overlap exists between these two behavioral processes. In addition to conceptual overlap in symptomatology, there is a strong commonality between the two domains regarding the roles and sites of action of dopamine, opioids, and corticotrophin-releasing factor (CRF).”
    Quote in the paperSubstantial claim
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    A wide range of animal paradigms model individual features of addiction behavior.

    “a wide range of paradigms have been developed that model individual features of addiction behavior (Sanchis-Segura and Spanagel 2006).”
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    Addiction as a behavioral phenomenon includes drug consumption, reinforcement learning, drug seeking, relapse, tolerance, and withdrawal.

    “Addiction as a behavioral phenomenon is complex and has several components or phases, including drug consumption, reinforcement learning, drug seeking, relapse, tolerance, and withdrawal (Sanchis-Segura and Spanagel 2006).”
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    All known drugs of abuse cause dopamine release in the nucleus accumbens, preferentially in the shell, and more broadly throughout the mesolimbic dopamine pathway.

    “All known drugs of abuse cause DA release in the nucleus accumbens (NAC), preferentially in the nucleus accumbens shell region (NACs), and also more broadly throughout the mesolimbic DA pathway (Swanson 1982; Di Chiara and Imperato 1988; Koob and Bloom 1988; Pontieri et al 1995; Pontieri et al 1996; Tanda et al 1997; for a review, see Di Chiara et al 2004).”
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    Reinstatement of drug-seeking after extinction is widely considered a valid model for drug-seeking and relapse.

    “Reinstatement can occur as a result of stress, cues that predict the drug, or a dose of the drug itself, and is widely considered to be a valid model for drug-seeking and for relapse in general.”
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    Rodents will not self-administer selective D1R agonists, which by themselves induce conditioned place aversion.

    “Nonetheless, rodents will not self-administer selective D1R agonists, which, by themselves, induce conditioned place aversion (Woolverton et al 1984; Hoffman and Beninger 1988).”
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    D2R is necessary for reward and incentive learning and maintenance of reward- and incentive-related responding, and rats will self-administer drugs that activate these receptors.

    “D2R is also necessary for reward and incentive learning, as well as maintenance of reward- and incentive-related responding, and rats will readily self-administer drugs that activate these receptors (Woolverton et al 1984; Sanger 1986; Woolverton 1986; Hoffman et al 1988; for an early review of the roles of D1R and D2R in reward and reinforcement, see Beninger et al 1989).”
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    Most rodents are not spontaneously maternal, and naïve females often attack or ignore pups.

    “Most rodents are not spontaneously maternal, and naïve females will often attack or ignore pups (Wiesner and Sheard 1933).”
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    Cocaine primarily activates D1R-containing neurons, which are necessary for reward-related learning and maintenance of reward-related behaviors.

    “Cocaine primarily activates D1R-containing neurons, which are necessary for reward-related learning and maintenance of reward-related behaviors (Nakajima 1986; Nakajima and Mckenzie 1986; Beninger et al 1987; Bertran-Gonzalez et al 2008).”
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    Rats and mice do not appear to be selective in their maternal care, as experienced mothers will direct maternal behavior toward any pups they encounter.

    “Nonetheless, rats and mice do not appear to be selective in their maternal care, as experienced mothers will direct maternal behavior toward any pups they encounter (Wiesner and Sheard 1933).”
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    Rat mothers will press a lever repeatedly to gain access to pups and will cross an electrified grid to retrieve them.

    “Rat mothers will press a lever repeatedly to gain access to pups, and will cross an electrified grid in order to retrieve them (Nissen 1930; Wilsoncroft 1969).”
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    Dopamine release in the nucleus accumbens and subsequent D1R activation is necessary for normal maternal behavior and sufficient to induce maternal behavior under conditions where it would otherwise not occur.

    “This DA release in the NAC, and the subsequent activation of D1R, is both necessary for the normal expression of maternal behavior and sufficient to induce maternal behavior under conditions where it would otherwise not occur (Gaffori and Le Moal 1979; Hansen et al 1991a; Hansen et al 1991b; Keer and Stern 1999; Numan et al 2005; Stolzenberg et al 2007; Stolzenberg et al 2010; for a review, see St”
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    The mesolimbic dopamine pathway is generally thought to be involved in motivation to work for rewards and in the salience of incentives.

    “the mesolimbic DA pathway is generally thought to be involved in both the motivation to act or work for rewards, and the salience of incentives (Mogenson 1987; Blackburn et al 1992; Ikemoto and Panksepp 1999; for a complete discussion of the many competing theories regarding the role of the mesolimbic DA pathway, see Wise 2004; Berridge 2007).”
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    Acute stressors are encoded by dopamine-releasing neurons in the VTA, cause dopamine sensitization in the nucleus accumbens, and facilitate drug taking; mesolimbic dopamine release also encodes negative motivational states and stress responses.

    “Recent experiments also show that acute stressors are encoded by DA-releasing neurons in the VTA, cause DA sensitization in the NAC, and facilitate drug taking; demonstrating that mesolimbic DA release also encodes negative motivational states and stress responses (Miczek et al 2011; Wang and Tsien 2011).”
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    Chronic cocaine exposure is associated with increased phasic D1R signaling involved in reward prediction, sensitized reward responses, and dampening further reinforcing effects of drugs of abuse.

    “Chronic exposure to cocaine is associated with an increase in phasic D1R signaling, and these increases are involved in reward prediction, sensitized responses to reward, and in dampening further reinforcing effects of drugs of abuse (Ljungberg et al 1992; Henry and White 1995; Self et al 1996; Anderson et al 2008; Bertran-Gonzalez et al 2008; Zweifel et al 2009).”
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    Premisecitation

    D2R also seems to be necessary for maternal behavior, though the site of action has not been determined.

    “D2R also seems to be necessary for maternal behavior, though the site of action has not been determined (Silva et al 2001).”
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    Lesions that disrupt dopamine release from the VTA into the nucleus accumbens, or simultaneous antagonism of D1R and D2R in the nucleus accumbens, disrupt motivated maternal behaviors such as retrieval without disrupting passive maternal behaviors such as nursing.

    “Similarly, lesions that disrupt the release of DA from the VTA into the NAC, or simultaneous antagonism of D1R and D2R in the NAC, all disrupt motivated maternal behaviors such as retrieval without disrupting passive maternal behaviors such as nursing (Gaffori and Le Moal 1979; Hansen et al 1991a; Hansen et al 1991b; Keer and Stern 1999).”
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    Cocaine seeking behavior is inhibited by pharmacological D1R activation, suggesting that experimental D1R activation may reduce the drive to seek out rewards.

    “Furthermore, cocaine seeking behavior is inhibited by pharmacological D1R activation (Self et al 1996), suggesting that this experimental activation of D1R may reduce the drive to seek out rewards.”
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    Interpretationreasoning

    D2R activation in the nucleus accumbens shell enhances, while D1R activation inhibits, pair bond formation.

    “These results suggest that D2R activation in the NAC shell enhances, while D1R activation inhibits, pair bond formation. (For a review see Aragona and Wang 2009.)”
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    Prairie voles reliably form a pair bond after 24 hours of mating, but not after 6 hours of cohabitation where mating is prevented.

    “Although mating is not required for pair bonding, prairie voles will reliably form a pair bond with a partner after 24 hours of mating, but not after 6 hours of cohabitation where mating is prevented (Wang et al 1999).”
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    In prairie voles, mating partners form highly selective pair bonds, share a nest, coordinate care of offspring, and display high levels of affiliative behavior toward each other and their young.

    “In this species, mating partners form highly selective pair bonds, share a nest, coordinate care of offspring, and display high levels of affiliative behavior toward each other and their young (Thomas and Birney 1979; Getz et al 1981; Ahern et al 2011).”
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    Pair bonding in prairie voles is operationally defined by two observable behaviors: partner preference and selective aggression.

    “Pair bonding in prairie voles is operationally defined based on two observable behaviors: preference for the partner over the stranger in a choice test (or “partner preference”); and intense aggression toward prairie voles other than the mate (or “selective aggression”) (Williams et al 1992; Winslow et al 1993; Insel et al 1995; Wang et al 1997b; Ahern et al 2009).”
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    The mixed D1R and D2R agonist apomorphine enhances pair bonding when injected into the nucleus accumbens shell in low doses, but fails to enhance pair bonding at high doses.

    “Furthermore, the mixed D1R and D2R agonist apomorphine enhances pair bonding when injected into the NAC shell in low doses, where it would be expected to bind preferentially to D2R; but fails to enhance pair bonding at high doses where it would be expected to bind to D1R as well (Aragona et al 2003).”
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    Interpretationcitation

    Selective activation of D1R disrupts several kinds of reward and incentive learning and may disrupt learning of associations between the social reward provided by the partner and the partner's specific identity cues.

    “Selective activation of D1R also disrupts several kinds of reward and incentive learning (Beninger et al 1989) and therefore may disrupt the learning of associations between the social reward provided by the partner, and the partner’s specific identity cues.”
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    Prairie voles are spontaneously parental, with offspring often assisting in the care of siblings.

    “They are also spontaneously parental, with offspring often assisting in the care of siblings (Carter and Roberts 1997).”
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    Premisecitation

    Partner preference measures the motivational force bringing partners together, while selective aggression measures mate-guarding and rejection of new potential partners.

    “These two observable behavioral measures describe different dimensions of the pair bond: partner preference measures the motivational force bringing the partners together, while selective aggression measures mate-guarding and the rejection of new, potential partners (Carter et al 1995).”
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    Interpretationcitation

    Selective aggression serves in part to reject potential new partners and represents an ongoing mechanism of maintenance of the pair bond.

    “Since this selective aggression serves, in part, to reject potential new partners, it represents an ongoing mechanism of maintenance of the pair bond (Carter et al 1995).”
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    Amphetamine injected directly into the nucleus accumbens shell creates a 20-fold increase in local extracellular dopamine, which enhances pair bonding only if a D1R antagonist is also given.

    “Finally, amphetamine injected directly into the NAC shell creates a 20-fold increase in local extracellular dopamine, which enhances pair bonding only if a D1R antagonist is also given (Curtis and Wang 2007).”
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    Mating-induced pair bonding is prevented by selective blockade of D2R in the nucleus accumbens shell, while activation of these receptors induces bonding in the absence of mating.

    “Mating-induced pair bonding is prevented by selective blockade of D2R in the NAC shell, while activation of these receptors induces bonding in the absence of mating (Wang et al 1999; Gingrich et al 2000; Aragona et al 2006).”
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    Some have proposed that maternal attachment is the evolutionary precursor for monogamous bonding.

    “Nonetheless, monogamy has evolved independently in multiple orders of the class Mammalia, suggesting that the evolutionary precursor for monogamous bonding is present in most if not all mammals; and some have proposed that maternal attachment is that precursor (Getz and Hofmann 1986; Ross and Young 2009; for a review of the evolution of monogamy, see Freeman and Young, in press).”
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    Sexually naïve male prairie voles are highly social and show very little aggressive behavior toward novel conspecifics.

    “Sexually naïve male prairie voles are highly social and show very little aggressive behavior toward novel conspecifics (Insel et al 1995).”
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    Expression of D1R in the nucleus accumbens is 60% higher in males that cohabitate with a female than in males that cohabitate with a same-sex sibling; D1R in dorsal striatum, and D2R in both areas, remains unchanged.

    “Expression of D1R in the NAC is 60% higher in males that cohabitate with a female than it is in males that cohabitate with a same-sex sibling; while D1R in dorsal striatum, and D2R in both areas, remains unchanged.”
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    D2 antagonists have long been used to treat aggression in schizophrenia and other psychiatric disorders in humans despite nonspecific effects on behavior in general.

    “and to the treatment of aggression in schizophrenia and other psychiatric disorders in humans, where D2 antagonists have long been used despite the nonspecific effects on behavior in general (Yudofsky et al 1987; Buckley 1999).”
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    Interpretationcitation

    Activation of D2R in the nucleus accumbens shell is rewarding and enhances reward-based learning, which aligns with the role of D2R in pair bonding.

    “Meanwhile, activation of D2R in the NAC shell is rewarding and enhances reward-based learning (Beninger et al 1989), which aligns well with the role of D2R in pair bonding.”
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    Prairie voles are socially monogamous rodents indigenous to most of the Midwest United States and Canada.

    “Prairie voles are socially monogamous rodents indigenous to most of the Midwest United States and Canada (Tamarin 1985).”
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    Sexually naïve male prairie voles that receive this treatment reject potential female partners and fail to bond with them, exactly as is observed in pair-bonded males.

    “Sexually naïve male prairie voles that receive this treatment reject potential female partners and fail to bond with them, exactly as is observed in pair-bonded males (Liu et al 2010).”
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    All three opioid receptor types are Gi protein-coupled receptors that are inhibitory when activated.

    “All three receptor types are GI protein-coupled receptors that are inhibitory when activated (Burns et al 1983; Tsunoo et al 1986; North et al 1987; for a review see Knapp et al 1995).”
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    These same males are able to form a pair bond if D1R in the nucleus accumbens shell is blocked with an antagonist, a manipulation that prevents the expression of selective aggression in drug-naïve, pair bonded males.

    “These same males are able to form a pair bond if D1R in the NAC shell is blocked with an antagonist, a manipulation that prevents the expression of selective aggression in drug-naïve, pair bonded males (Aragona et al 2006; Liu et al 2010).”
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    Activation of MOR in the VTA, and DOR in the nucleus accumbens, is directly rewarding in rats, producing conditioned place preferences.

    “Activation of MOR in the VTA, and DOR in the NAC, is directly rewarding in rats, producing conditioned place preferences (Goeders et al 1984; Olmstead and Franklin 1997b).”
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    The endogenous opiate system comprises three opioid receptor types (MOR, DOR, KOR) and their endogenous ligands endorphin, enkephalin, and dynorphin.

    “This system is comprised of three types of opioid receptors: μ (MOR), δ (DOR), and κ (KOR); and their endogenous ligands, endorphin, enkephalin and dynorphin (Hughes et al 1975; Birdsall and Hulme 1976; Goldstein et al 1979; Evans et al 1992; Kieffer et al 1992; Chen et al 1993; Yasuda et al 1993).”
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    The three opioid receptor types have distinct patterns of expression throughout the brain, with significant inter-species variability.

    “These three receptor types have distinct patterns of expression throughout the brain, and these patterns have significant inter-species variability (Khachaturian et al 1985; Robson et al 1985; Mansour et al 1987; Mansour et al 1988; Mansour et al 1994a; Mansour et al 1994b; Curran and Watson 1995; Resendez et al 2012).”
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    • cites Khachaturian et al 1985 source

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    • cites Robson et al 1985 source

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    • cites Mansour et al 1987

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  • May combine two claims.

    Premisecitation

    The A118G variant of the MOR gene has enhanced binding and signaling properties but low expression of mRNA.

    “The A118G variant has enhanced binding and signaling properties, but low expression of mRNA (Zhang et al 2005; Kroslak et al 2007).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Zhang et al 2005 source

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    • cites Kroslak et al 2007 source

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  • May combine two claims.

    Premisecitation

    Human subjects possessing the A118G variant of the MOR gene have altered reinforcement learning and increased risk for alcohol dependence.

    “Human subjects possessing this variant of the MOR gene have altered reinforcement learning and increased risk for alcohol dependence (Lee et al 2011; Koller et al 2012).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Lee et al 2011 source

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    • cites Koller et al 2012

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  • May combine two claims.

    Premisecitation

    Opioid antagonists reduce cravings and are now being used to treat an increasing variety of disorders including alcoholism, opioid dependence, and obesity.

    “Furthermore, opioid antagonists are known to reduce cravings, and are now being used to treat an increasing variety of disorders including alcoholism, opioid dependence, and obesity (O’Malley et al 1992; Volpicelli et al 1992; Greenway et al 2010; Minozzi et al 2011).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites O’Malley et al 1992

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    • cites Volpicelli et al 1992 source

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    • cites Greenway et al 2010 source

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  • May combine two claims.

    Premisecitation

    MOR and DOR mediate generally positive motivation and affect, while KOR mediates generally negative motivation and aversion.

    “Nonetheless, the three receptor types have different behavioral domains, with MOR and DOR mediating generally positive motivation and affect, while KOR mediates generally negative motivation and aversion (Pfeiffer et al 1986; Shippenberg et al 1987; Pecina and Berridge 2000; McLaughlin et al 2003; for reviews see Van Ree et al 2000; Le Merrer et al 2009).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Pfeiffer et al 1986 source

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    • cites Shippenberg et al 1987 source

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    • cites Pecina and Berridge 2000 source

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  • May combine two claims.

    Premisecitation

    Opioids interact with other addiction-related neurochemical systems, including dopamine and glutamate, though these interactions do not entirely explain the role of opioids in addiction.

    “Opioids also interact with other addiction-related neurochemical systems, including DA and glutamate, though these interactions do not entirely explain the role of opioids in addiction (Dichiara and Imperato 1988; Johnson and North 1992; Scavone et al 2011).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Dichiara and Imperato 1988 source

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    • cites Johnson and North 1992 source

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    • cites Scavone et al 2011 source

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  • May combine two claims.

    Premisecitation

    Both human A118G subjects and Rhesus C77G subjects are more likely to respond positively to opioid antagonist therapy for alcoholism, suggesting that differences in receptor properties may be a direct physiological mechanism for the increased risk for alcoholism.

    “Interestingly, both human A118G subjects and Rhesus C77G subjects are more likely to respond positively to opioid antagonist therapy for alcoholism, which suggests that the differences in receptor properties may be a direct physiological mechanism for the increased risk for alcoholism (Oslin et al 2003).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Oslin et al 2003 source

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  • May combine two claims.

    Premisecitation

    Drugs of abuse cause dynorphin release and up-regulation of dynorphin in the dorsal and ventral striatum, which inhibits dopamine release by acting on KOR receptors and is therefore thought to be compensatory.

    “Drugs of abuse cause dynorphin release and up-regulation of dynorphin in the dorsal and ventral striatum, which inhibits DA release by acting on KOR receptors and is therefore thought to be compensatory (Hanson et al 1988; Sivam 1989; Spanagel et al 1990; Hurd et al 1992; Daunais et al 1993; El Daly et al 2000; Isola et al 2009).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Hanson et al 1988 source

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    • cites Sivam 1989 source

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    • cites Spanagel et al 1990 source

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  • May combine two claims.

    Premisecitation

    Mice that lack the MOR gene do not develop place preferences or withdrawal symptoms in response to morphine.

    “Mice that lack the MOR gene do not develop place preferences or withdrawal symptoms in response to morphine (Matthes et al 1996).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Matthes et al 1996 source

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  • May combine two claims.

    Premisecitation

    Individuals with the A118G MOR polymorphism show increased likelihood of affectionate relationships, increased sensitivity to social rejection, and increased brain activity in a rejection task, possibly indicating an altered attachment style.

    “Individuals possessing the A118G polymorphism of the MOR gene show increased likelihood to engage in affectionate relationships, increased sensitivity to social rejection, and increased brain activity in a rejection task, possibly signaling an altered attachment style (Way et al 2009; Troisi et al 2011a).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Way et al 2009 source

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    • cites Troisi et al 2011a

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  • May combine two claims.

    Premisecitation

    In rats, guinea pigs, sheep, and Rhesus macaques, acute opioid antagonists increase social need and care solicitation by offspring, while agonists decrease solicitation in rats and Rhesus macaques.

    “In rats, guinea pigs, sheep, and Rhesus macaques, acute administration of opioid antagonists increases social need and the solicitation for care by offspring, while agonists decrease solicitation in rats and Rhesus macaques (Panksepp et al 1980; Kalin et al 1988; Martel et al 1993; Panksepp et al 1994; Martel et al 1995; Shayit et al 2003).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Panksepp et al 1980 source

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    • cites Kalin et al 1988 source

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    • cites Martel et al 1993 source

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  • May combine two claims.

    Premisecitation

    MOR knockout mouse pups emit fewer distress vocalizations after maternal separation and show less selectivity for their mother's cues.

    “In MOR knockout mice, pups emit fewer DVs in response to maternal separation and show less selectivity for their mothers’ cues (Moles et al 2004).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Moles et al 2004 source

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  • May combine two claims.

    Premisecitation

    In puppies, opioid antagonists increase social solicitation for attention but reduce the comfort obtained from later social contact.

    “Similarly, social solicitation for attention in puppies is increased with opioid antagonists, although this same treatment reduces the comfort received from subsequent social contact (Panksepp et al 1980).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Panksepp et al 1980 source

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  • May combine two claims.

    Premisecitation

    In the striatum, DA D2R is expressed in neurons containing enkephalin, the endogenous MOR ligand; both receptors are involved in pair bond formation but not maintenance.

    “In the striatum, DA D2R is expressed in striatal neurons containing enkephalin, the endogenous ligand for MOR (Gerfen and Young 1988); both of which receptors are involved in pair bond formation but do not have a clear role in maintenance.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Gerfen and Young 1988 source

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  • May combine two claims.

    Premisecitation

    Infant Rhesus macaques with the C77G MOR polymorphism show increased attachment to mothers, and Rhesus mothers with it show enhanced maternal care and higher OT release during maternal behavior.

    “Infant Rhesus macaques possessing the C77G polymorphism of the MOR gene show increased attachment to their mothers, while Rhesus mothers possessing the same polymorphism show enhanced maternal care and higher OT release during maternal behaviors (Barr et al 2008; Higham et al 2011).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Barr et al 2008 source

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    • cites Higham et al 2011 source

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  • May combine two claims.

    Premisereasoning

    Under the opioid hypothesis, high opioid receptor activation signals a social reward state, whereas low activation produces a drive to seek social rewards.

    “According to this hypothesis, high activation of opioid receptors signals a social reward state, and low activation induces a drive to seek social rewards.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationcitation

    The role of MOR in maternal behavior is conserved in humans, and mechanisms encoding altered attachment style overlap directly with risk of alcohol dependence.

    “These studies suggest that the role of MOR in maternal behavior is conserved in humans, and also show a direct overlap between mechanisms that encode for altered attachment style and risk of alcohol dependence.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    CRF administration into rat brain reproduces many stress-like behavioral responses, while CRF antagonists reduce or prevent stress responses.

    “Administration of CRF into the brain of rats reproduces many different behavioral responses analogous to stress, while CRF antagonists reduce or prevent stress responses (Dunn and Berridge 1990; Heinrichs et al 1994; Menzaghi et al 1994).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Dunn and Berridge 1990 source

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    • cites Heinrichs et al 1994 source

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    • cites Menzaghi et al 1994 source

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  • May combine two claims.

    Premisecitation

    Opioid agonists promote mother-offspring bond formation and later maternal behavior in sheep, while opioid antagonists prevent both mother-offspring and offspring-mother bonding.

    “Opioid agonists potentiate the formation of mother-offspring bonds and subsequent maternal behavior in sheep, while opioid antagonists prevent both mother-offspring and offspring-mother bonding (Kendrick and Keverne 1989; Keverne and Kendrick 1994; Shayit et al 2003).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Kendrick and Keverne 1989 source

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    • cites Keverne and Kendrick 1994 source

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    • cites Shayit et al 2003 source

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  • May combine two claims.

    Premisecitation

    When rats mate, they associate ejaculation reward with non-social cues and can form preferences for location or non-social odors.

    “Instead, when rats mate, they associate the reward from ejaculation with non-social cues and can form preferences for the location or for non-social odors (Miller and Baum 1987; Mehrara and Baum 1990; Ismail et al 2009).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Miller and Baum 1987 source

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    • cites Mehrara and Baum 1990 source

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    • cites Ismail et al 2009 source

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  • May combine two claims.

    Premisecitation

    In prairie voles, a peripherally administered non-selective opioid antagonist prevents pair bond formation but also reduces mating.

    “In this species, a peripherally administered, non-selective opioid antagonist prevents pair bond formation between prairie voles, but also reduces mating (Burkett et al 2011).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Burkett et al 2011 source

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  • May combine two claims.

    Premisecitation

    D1R is expressed in striatal neurons containing dynorphin, the endogenous KOR ligand; both are necessary for pair bond maintenance but not formation.

    “Conversely, D1R is expressed in striatal neurons containing dynorphin, the endogenous ligand for KOR; and both of these receptors are necessary for pair bond maintenance, but not for formation.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Opioids mediate many aspects of maternal behavior, and the opiate system was the first brain system implicated in animal social attachment.

    “Opioids also mediate many aspects of maternal behavior, and in fact, the opiate system was the first brain system to be implicated in social attachment in animals (Panksepp et al 1978).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Panksepp et al 1978

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Low non-sedative doses of opioid agonists suppress separation distress calls in young puppies, guinea pigs, and chicks, while opioid antagonists usually elicit such calls.

    “Distress vocalizations are effectively blocked by low, non-sedative doses of opioid agonists, and, in most cases, induced by opioid antagonists (Panksepp et al 1978; Warnick et al 2005).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Panksepp et al 1978

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    • cites Warnick et al 2005 source

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  • May combine two claims.

    Interpretationreasoning

    Opioid antagonists produce a social drive while simultaneously blocking the rewarding effects of social contact.

    “Thus, opioid antagonists induce a social drive while simultaneously blocking the rewarding effects of social contact.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    The relationships between MOR and reward/formation, and between KOR and maintenance, are the same as in drug addiction.

    “Furthermore, it is interesting to note that the relationships between MOR and reward/formation, and between KOR and maintenance, are the same as in drug addiction.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    The caudate-putamen is activated in humans when viewing faces of loved ones, and this activation correlates with romantic love and passion scores.

    “Additionally, the CP is activated in humans when viewing the faces of loved ones, and this activation is correlated with romantic love and passion scores (Bartels and Zeki 2000; Aron et al 2005; Acevedo et al 2012).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Bartels and Zeki 2000 source

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    • cites Aron et al 2005 source

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    • cites Acevedo et al 2012 source

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  • May combine two claims.

    Premisecitation

    Pair bonding in male prairie voles is enhanced by activation of CRF receptors in the brain, an effect localized to the NAC shell.

    “Pair bonding in males is also enhanced by activation of CRF receptors in the brain, an effect which can be localized to the NAC shell (DeVries et al 2002; Lim et al 2007).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites DeVries et al 2002 source

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    • cites Lim et al 2007 source

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  • May combine two claims.

    Premisecitation

    This CRF release produces a withdrawal-induced anxiety state that is reversed by CRF-R1 antagonists.

    “This CRF release produces a withdrawal-induced anxiety state that is reversed by CRF-R1 antagonists (Sarnyai et al 1995; Tucci et al 2003; Knapp et al 2004; Skelton et al 2007).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Sarnyai et al 1995 source

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    • cites Tucci et al 2003 source

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    • cites Knapp et al 2004 source

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  • May combine two claims.

    Premisecitation

    In male prairie voles, forced swim stress or peripheral corticosterone injections enhance subsequent pair bond formation, while these treatments inhibit pair bonding in females.

    “In male prairie voles, forced swim stress or peripheral corticosterone injections both enhance the subsequent formation of pair bonds, while these treatments inhibit pair bonding in females (DeVries et al 1996).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites DeVries et al 1996 source

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  • May combine two claims.

    Premisecitation

    The NAC shell is activated during stress in rats and mice, and this activation in mice is blocked by a CRF-R1 antagonist.

    “The NAC shell is activated during stress in rats and mice, and this same activation in mice is blocked by CRF-R1 antagonist (Pliakas et al 2001; Kreibich et al 2009).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Pliakas et al 2001 source

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    • cites Kreibich et al 2009 source

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  • May combine two claims.

    Interpretationreasoning

    CRF within the NAC shell may have a role in the stress-induced increase in incentive value of cues previously associated with reward, which may complement or create its more general effect on depression in that nucleus.

    “Taken together, these findings indicate that CRF within the NAC shell may have a role in the stress-induced increase in incentive value of cues previously associated with reward, which may complement or create its more general effect on depression in that nucleus.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Both OT and AVP are synthesized in the hypothalamus and released from the posterior pituitary into the peripheral circulation.

    “Both OT and AVP are synthesized in the hypothalamus and released from the posterior pituitary into the peripheral circulation (Gainer and Wray 1994; Burbach et al 2005).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Gainer and Wray 1994

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    • cites Burbach et al 2005

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  • May combine two claims.

    Premisecitation

    A CRF-R2 agonist also blocks excessive alcohol consumption in dependent rats, suggesting opposite roles for CRF-R1 and CRF-R2.

    “However, a CRF-R2 agonist also blocked excessive alcohol consumption in dependent rats, suggesting opposite roles for these two receptors (Valdez et al 2004).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Valdez et al 2004 source

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  • May combine two claims.

    Premisecitation

    CRF injected into the NAC shell induces depressive-like behavior in rats, reversed by a CRF-R1 antagonist.

    “Furthermore, CRF injected into in the NAC shell induces depressive-like behavior in rats, and this effect is reversed by a CRF-R1 antagonist (Chen et al 2012b).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Chen et al 2012b

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  • May combine two claims.

    Premisecitation

    Total social isolation in prairie voles leads to depressive-like behavior and an increase in CRF-immunoreactive neurons in the PVN.

    “Studies have shown that total social isolation in prairie voles leads to depressive-like behavior and an increase in CRF-immunoreactive neurons in the PVN (Grippo et al 2007a; Grippo et al 2007b).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Grippo et al 2007a

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    • cites Grippo et al 2007b

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  • May combine two claims.

    Premisecitation

    Monogamous and non-monogamous vole species differ greatly in brain distribution of CRF-R1 and CRF-R2, while CRF peptide distribution is highly conserved.

    “Monogamous and non-monogamous species of voles differ greatly in the distribution of CRF-R1 and CRF-R2 in the brain, even while the distribution of CRF peptide is highly conserved (Lim et al 2005; Lim et al 2006).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Lim et al 2005 source

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    • cites Lim et al 2006 source

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  • May combine two claims.

    Premisecitation

    Stress causes CRF release into the VTA and NAC shell, and this CRF acts on CRF receptors to induce reinstatement of drug seeking.

    “Using stress-induced reinstatement of drug seeking, it has been demonstrated that stress causes CRF release into the VTA and NAC shell (Wang et al 2005; Chen et al 2012a), and this CRF release acts on CRF receptors to induce reinstatement (Ungless et al 2003; Wang et al 2005; Wang et al 2007; Blacktop et al 2011).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Wang et al 2005 source

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    • cites Chen et al 2012a

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    • cites Ungless et al 2003 source

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  • May combine two claims.

    Premisecitation

    Permanent social loss can be traumatic and cause deterioration of physical health and susceptibility to depression.

    “This permanent social loss can be traumatic and cause both deterioration of physical health and susceptibility to depression (Prigerson et al 1997; Ott 2003).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Prigerson et al 1997 source

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    • cites Ott 2003 source

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  • May combine two claims.

    Premisereasoning

    Pair bonding induces an increase in CRF mRNA in the bed nucleus of the stria terminalis, suggesting this region may be the source of separation-induced CRF release and is primed during pair bond formation.

    “The study also showed that pair bonding induces an increase in CRF mRNA in the bed nucleus of the stria terminalis, a part of the extended amygdala; suggesting that this region may be the source of separation-induced CRF release, and is primed for such release during pair bond formation.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Four days of separation from a pair-bonded mate leads to increases in passive coping strategies in male prairie voles, and this does not occur after separation from a male sibling.

    “Of more direct relevance is the finding that 4 days of separation from a pair-bonded mate leads to increases in passive coping strategies, a type of depressive-like behavior, in male prairie voles, and that this increase does not occur in response to separation from a male sibling (Bosch et al 2009).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Bosch et al 2009 source

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  • May combine two claims.

    Premisecitation

    CRF-R1 activation mediates many stress effects in rats; CRF-R2 activation can be the same, opposing, or ineffective depending on assay and brain region.

    “CRF-R1 activation mediates many of these stress effects in rats, while CRF-R2 activation is alternately the same as CRF-R1, opposing CRF-R1, or ineffective in modulating stress, depending on the assay and the brain region (Ho et al 2001; Takahashi et al 2001; Valdez et al 2004; Zhao et al 2007; for a review see Heinrichs and Koob, 2004).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Ho et al 2001

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    • cites Takahashi et al 2001 source

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    • cites Valdez et al 2004 source

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  • May combine two claims.

    Premisecitation

    OT and AVP are paralogous 9-amino-acid peptides unique to mammals, though homologues exist in many vertebrates and invertebrates.

    “These two paralogous 9-amino-acid peptides are unique to mammals, though homologues exist in a wide variety of vertebrates and invertebrates (Archer 1974; van Kesteren et al 1992).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Archer 1974

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    • cites van Kesteren et al 1992 source

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

    Premisenone given

    DA, opioids, and CRF all mediate processing of different aspects of reward, reinforcement, and motivated behavior.

    “DA, opioids, and CRF all mediate the processing of different aspects of reward, reinforcement, and motivated behavior.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Premisecitation

    OT mitigates withdrawal symptoms of morphine and alcohol and reduces drug-primed reinstatement of responding for amphetamine.

    “OT also mitigates the withdrawal symptoms of morphine and alcohol, and reduces drug-primed reinstatement of responding for amphetamine (Kovacs et al 1981; Szabo et al 1988; Carson et al 2010a).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Kovacs et al 1981 source

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    • cites Szabo et al 1988

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    • cites Carson et al 2010a

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  • May combine two claims.

    Premisecitation

    In prairie voles, spontaneous maternal behavior in virgin females is organized by developmental OTR levels in the NAC shell and is prevented by OTR antagonist injected there.

    “Additionally, in prairie voles, the expression of spontaneous maternal behavior in virgin females is organized by developmental (and not adult) levels of OTR in the NAC shell, and is prevented by OTR antagonist injected into this region (Olazabal and Young 2006a; Ross et al 2009b; Keebaugh and Young 2011).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Olazabal and Young 2006a

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    • cites Ross et al 2009b

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    • cites Keebaugh and Young 2011 source

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  • May combine two claims.

    Premisecitation

    OT injected into the NAC is sufficient to induce pair bonds in female prairie voles without mating and interacts with DA D2R.

    “In addition, OT injected into the NAC is sufficient to induce pair bonds in female prairie voles in the absence of mating, and interacts with DA D2R to do so (Liu and Wang 2003).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Liu and Wang 2003 source

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  • May combine two claims.

    Premisecitation

    Mice with reduced AVP release in the LS or lacking AVP V1aR have selective deficits in social memory, and re-introduction of AVP or V1aR in the LS rescues social memory.

    “Mice with reduced AVP release in the LS, or that are lacking AVP V1aR entirely, have similarly selective deficits in social memory, and re-introduction of AVP or V1aR respectively in the LS results in rescue of social memory (Bielsky et al 2004; Bielsky et al 2005; Lukas et al 2011).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Bielsky et al 2004

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    • cites Bielsky et al 2005 source

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    • cites Lukas et al 2011 source

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

    Premisecitation

    AVP is released peripherally in response to osmotic challenge and acts in the kidney to induce water reabsorption and concentrate urine.

    “AVP is released peripherally in response to an osmotic challenge and acts in the kidney to induce the reabsorption of water, concentrating the urine (Gainer and Wray 1994).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Gainer and Wray 1994

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  • May combine two claims.

    Premisecitation

    OT has a single receptor (OTR); AVP has three receptors (V1aR, V1bR, V2R), with V1aR the principal receptor implicated in social processes.

    “OT has a single receptor (OTR) both in the periphery and in the brain. AVP has three receptors (V1aR, V1bR, and V2R); V1bR and V2R are expressed primarily in the periphery, and while V1aR and V1bR are present in the brain, V1aR is the principal receptor implicated in social processes (Gainer and Wray 1994; Burbach et al 2005; for a review of the AVP system and behavior see Caldwell et al 2008; for”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Gainer and Wray 1994

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    • cites Burbach et al 2005

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    • cites Caldwell et al 2008 source

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  • May combine two claims.

    Premisecitation

    OT is released centrally and peripherally during birth; centrally released OT is necessary for rapid onset of maternal behavior in virgin rats and sufficient to induce it.

    “OT is released centrally and peripherally during birth, and centrally released OT is both necessary for the rapid onset of maternal behavior in virgin rats, and sufficient to induce it (Pedersen and Prange 1979; Fahrbach et al 1985; van Leengoed et al 1987; Borrow and Cameron 2012).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Pedersen and Prange 1979 source

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    • cites Fahrbach et al 1985 source

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    • cites van Leengoed et al 1987 source

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  • May combine two claims.

    Premisecitation

    OT KO mice show no deficits in spatial memory, non-social olfactory memory, and habituation, and can retain social memory if intruders are painted with a non-social odor.

    “Importantly, the OT KO mice display no deficits in spatial memory, non-social olfactory memory, and habituation, and can retain a social memory if the novel intruders are painted with a non-social odor (Ferguson et al 2000; author’s unpublished data).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Ferguson et al 2000 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Both OT and AVP are released in the brain and bind to receptors there to affect social behaviors.

    “In addition, both peptides are released in the brain and bind to receptors there to affect social behaviors (Buijs et al 1983; Alonso et al 1986; Loup et al 1991).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Buijs et al 1983 source

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    • cites Alonso et al 1986 source

      Not checked in the free run.

    • cites Loup et al 1991 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Mice lacking the OT gene have impaired social memory that can be recovered by OT treatment before the initial social encounter, showing OT at the time of the salient event is necessary and sufficient for social memory.

    “Mice lacking the OT gene have impaired social memory that can be recovered by OT treatment prior to the initial social encounter, demonstrating that the presence of OT at the time of the salient event is necessary and sufficient for social memory (Ferguson et al 2000; Ferguson et al 2001).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Ferguson et al 2000 source

      Not checked in the free run.

    • cites Ferguson et al 2001 source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Interpretationcitation

    OT release induced by vaginocervical and nipple stimulation during human sex acts analogously to labor and nursing to induce bonding between sexual partners.

    “This has contributed to the theory that the OT release induced by vaginocervical stimulation and nipple stimulation during human sex, acts analogous to labor and nursing, serves to induce bonding between sexual partners (Young et al 2005); and furthermore, that pair bonding is mediated by neural systems that were elaborated in evolution from circuits originally adapted for maternal behavior (Ross”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Young et al 2005 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Exogenous OT attenuates many immediate behavioral effects of drugs of abuse, including reducing drug consumption, with some effects localized to the NAC.

    “Exogenous OT attenuates many of the immediate behavioral effects of drugs of abuse, including reducing drug consumption, and some of the attenuating effects of OT are localized to the NAC (Kovacs et al 1990; Sarnyai et al 1990; Sarnyai et al 1991; Qi et al 2009; Carson et al 2010a; Baracz et al 2012).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Kovacs et al 1990 source

      Not checked in the free run.

    • cites Sarnyai et al 1990 source

      Not checked in the free run.

    • cites Sarnyai et al 1991 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    OT is released by MDMA and is associated with the prosocial effects of this drug in humans and rats.

    “With respect to drugs of abuse, OT is best known for being released by 3,4-methylenedioxymethamphetamine (MDMA, Ecstasy), and is associated with the prosocial effects of this drug in both humans and rats (Wolff et al 2006; Thompson et al 2007; Dumont et al 2009; Thompson et al 2009).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Wolff et al 2006 source

      Not checked in the free run.

    • cites Thompson et al 2007 source

      Not checked in the free run.

    • cites Dumont et al 2009 source

      Not checked in the free run.

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

    Premisecitation

    OT is released peripherally during labor and helps evoke uterine contractions; release is also induced by vaginocervical stimulation and nipple stimulation, which induces milk letdown.

    “OT is released peripherally during labor and helps to evoke uterine contractions (Burbach et al 2005). The release of OT is also induced by vaginocervical stimulation, either from birth or from copulation. OT is also released peripherally in response to nipple stimulation, which, in the context of nursing, induces the letdown of milk from the mammary glands (Christensson et al 1989).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Burbach et al 2005

      Not checked in the free run.

    • cites Christensson et al 1989 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    OT and AVP facilitate social memory, while OT and AVP antagonists interfere with it.

    “OT and AVP facilitate this kind of social memory, while OT and AVP antagonists interfere with it.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Dantzer et al 1987 source

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    In partner addiction, encounters with the partner become more frequent and the relationship may continue despite negative emotions or consequences, understood as dependence-induced escalation of consumption.

    “Second, encounters with the partner become more frequent, and the relationship may continue despite negative emotions or consequences; this can be understood as a dependence-induced escalation of consumption of the object of addiction.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    In partner addiction, encounters with new potential mates cause novelty-induced DA release, but predominant D1R signaling promotes rejection, aggressive territory defense, and decreased probability of a second pair bond.

    “Finally, encounters with new potential mates continue to cause novelty-induced DA release, but now a predominance of D1R signaling promotes rejection, aggressive responses to defend the territory (including the mate), and a decrease in the probability that a second pair bond will form.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    OT in humans is higher during early romantic relationships, correlates with couples' interactive reciprocity, and predicts which couples stay together after 6 months.

    “OT in humans is also higher during early romantic relationships, is correlated with couples’ interactive reciprocity, and predicts which couples will stay together after 6 months (Schneiderman et al 2012).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Schneiderman et al 2012 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    AVP plays a role in paternal care and pair bonding in male prairie voles.

    “AVP also plays a role in paternal care and pair bonding in male prairie voles (Fig. 3d-f).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    In substance addiction, sensory information concerns the drug's taste, smell, unique experience, and context; in partner addiction, it is primarily social.

    “In substance addiction, this applies to the sensory modalities appropriate for the drug: the taste and smell; the particular experience unique to the drug; and the context in which the drug is taken. With partner addiction, this information is primarily social: looks, touches, words, scents, the shape of the body and face, and possibly sexual experiences.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.

    Interpretationnone given

    Concurrent activation of D1R and D2R may represent a balance of positive and negative behavioral responses, with D1R enhancing positive incentive value of active or aggressive responses and D2R enhancing positive incentive value of passive, reward-related, or pro-social responses.

    “Concurrent activation of D1R and D2R may represent a balance of positive and negative behavioral responses – D1R enhancing the positive incentive value of active or aggressive responses, and D2R enhancing the positive incentive value of passive, reward-related, or pro-social responses.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationreasoning

    The balance of DA signaling is altered in favor of D1R, leading to a progressive decrease in reward and an increase in negative affect or aggressive responses.

    “The balance of DA signaling is altered in favor of D1R, leading to a progressive decrease in reward and an increase in negative affect or aggressive responses.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationreasoning

    Both AVP and OT are involved in social information processing and are thought to enhance the salience of social stimuli.

    “Both AVP and OT are involved in social information processing and are thought to act to enhance the salience of social stimuli.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationcitation

    OT released during withdrawal may mitigate withdrawal symptoms and decrease probability of relapse, which could explain consolation-seeking during break-ups and social support's positive outcomes in drug addiction.

    “OT released during the withdrawal period may act to mitigate withdrawal symptoms and decrease the probability of relapse, which could explain both consolation-seeking behavior during break-ups, and the strong ability of social support to promote positive outcomes in drug addiction (Wills and Cleary 1996; Measelle et al 2006).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Wills and Cleary 1996 source

      Not checked in the free run.

    • cites Measelle et al 2006 source

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Addiction is created by positive reinforcement and incentive salience from DA, reward from opioids, and in partner addiction enhanced salience of social cues by OT and AVP.

    “Thus, addiction is created by positive reinforcement and incentive salience from DA; by reward from opioids; and, in the case of partner addiction, by enhanced salience of social cues by OT and AVP.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    This potentiated CRF system is strongly activated during drug withdrawal and separation anxiety, resulting in a positive motivational state driving the subject toward the object of addiction.

    “This potentiated system is strongly activated during drug withdrawal and separation anxiety. This activation results in a positive motivational state driving the subject toward the object of addiction.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationnone given

    Upregulation of dynorphin and subsequent KOR activation during withdrawal promotes negative affect and drives maintenance behavior.

    “Upregulation of dynorphin and subsequent activation of KOR during withdrawal promotes negative affect and drives maintenance behavior.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Premisecitation

    OT is released in humans during hugging, touching, massage, nipple stimulation, and orgasm, and promotes increased eye gaze, trust, and attention to emotional cues.

    “OT is released in humans during hugging, touching, massage, nipple stimulation, and orgasm (Carmichael et al 1987; Christensson et al 1989; Turner et al 1999; Light et al 2000; Light et al 2005), and promotes increased eye gaze, trust, and attention to emotional cues (Kosfeld et al 2005; Domes et al 2007; Guastella et al 2008; Andari et al 2010).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Carmichael et al 1987 source

      Not checked in the free run.

    • cites Christensson et al 1989 source

      Not checked in the free run.

    • cites Turner et al 1999 source

      Not checked in the free run.

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

    Interpretationnone given

    In the nascent phase of addiction, large amounts of sensory information are gathered about the object of addiction.

    “In the nascent phase of addiction, large amounts of sensory information are gathered about the object of addiction.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationreasoning

    The OT system, as an evolutionary elaboration of maternal circuitry, may promote nurturing behaviors and identification of the partner as an object of care.

    “The OT system, as an evolutionary elaboration of maternal circuitry, may promote nurturing behaviors and the identification of the partner as an object of care.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Premisecitation

    AVP induces pair bonding in male prairie voles when injected into the LS, and V1aR antagonist in the LS or VP prevents pair bonding.

    “AVP induces pair bonding in male prairie voles when injected into the LS, and V1aR antagonist in the LS or the ventral pallidum (VP) prevents pair bonding (Wang et al 1994; Lim and Young 2004).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Wang et al 1994 source

      Not checked in the free run.

    • cites Lim and Young 2004

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    AVP injected into the LS increases paternal behavior in male prairie voles, while V1aR antagonist decreases it.

    “AVP injected into the LS increases, while V1aR antagonist decreases, paternal behavior in male prairie voles (Wang et al 1994).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wang et al 1994 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Meadow voles are closely related to prairie voles but mate promiscuously without forming bonds between mating partners.

    “Meadow voles are a closely related vole species that mates promiscuously without forming bonds between mating partners (Madison 1980).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Madison 1980 source

      Not checked in the free run.

115 of 115

  • The authors state this as an assumption.

    The maternal attachment and pair bonding behaviors observed in animals are the evolutionary antecedents of human social attachments, which the paper refers to as love.

    “Our extrapolation from animal studies of maternal attachment and pair bonding to human love relies on the supposition that these behaviors are the evolutionary antecedents of human social attachments, which we refer to as love.”

  • The paper relies on this without support.

    These behaviors in humans and animals share common underlying neurobiological mechanisms.

    “While this may be debated, it is likely that these behaviors in humans and animals share common underlying neurobiological mechanisms.”

  • The paper relies on this without support.

    Findings from animal models of attachment, such as prairie voles and rodents, can be generalized to human attachment and love.

    “Human studies have shown the role of OT and OTR is largely conserved across species.”

  • The paper relies on this without support.

    The neurochemical systems underlying maternal bonding and pair bonding are the same systems that drugs of abuse subvert.

    “These data also provide evidence for the theory that social attachment systems governing maternal bonding and pair bonding to a mating partner are subverted by drugs of abuse to create addictions that are just as powerful as natural attachments.”
4 of 4

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?

  • This could initiate a feed-forward loop that progressively increases desire for the partner, similar to a drug addict's craving.

    “In this way, a feed-forward loop would be initialized, resulting in a progressive increase in desire for the partner, similar to a drug addict's increased craving for drug consumption (42).”

    From Oxytocin enhances brain reward system responses in men viewing the face of their female partner (2013) · cites this paper as [42]

    The Partner Addiction Hypothesis describes a positive feedback loop in both partner and substance addiction and says encounters with the partner become more frequent, paralleling escalating drug use. It does not say that desire or craving for the partner progressively increases; it describes decreasing reward as the loop continues. Page numbers are unavailable in our copy. (LLM, conf=0.82)

    Experimental — accuracy not yet measured

  • Oxytocin is a hypothalamic neuropeptide that acts as a complex neuromodulator capable of affecting a wide range of social and reproductive behaviors including pair bonding, maternal, affiliative and sexual behaviors.

    “oxytocin is a highly influential neuropeptide known to significantly impact a variety of social and reproductive behaviors including pair bonding, as well as maternal, affiliative and sexual behaviors (see Burkett and Young, 2012, Campbell, 2010, Donaldson and Young, 2008, Lee et al., 2009, Meyer-Lindenberg et al., 2011).”

    From Oxytocin, motivation and the role of dopamine (2014) · cites this paper as Burkett and Young, 2012

  • Oxytocin's ability to influence behavioral responses to social stimuli is at least partially mediated by its capacity to increase the salience of social cues.

    “Several groups of researchers have posited that oxytocin’s ability to influence behavioral responses to social stimuli is at least partially mediated by its capacity to increase the salience of social cues (Averbeck, 2010, Bartz et al., 2011, Burkett and Young, 2012, Gordon et al., 2011, Shamay-Tsoory et al., 2009).”

    From Oxytocin, motivation and the role of dopamine (2014) · cites this paper as Burkett and Young, 2012

Cites

Flagged first, then by citations

6 of the 407 references couldn’t be matched to a known paper.

Research works printed but not linked: 3

  1. Freeman SM, Young LJ (2012) Oxytocin, vasopressin, and the evolution of monogamy in mammals. In: Choleris E, Pfaff DW (eds) Oxytocin and vasopressin. Cambridge University Press, Cambridge
  2. Nakajima S (1986) Suppression of operant responding in the rat by dopamine D1-receptor blockade with SCH-23390. Physiol Psychol 14:111–114
  3. Plato Rowe CJ (1986) Phaedrus. Aris & Phillips, Warminster, Distributed in the USA and Canada by Humanities Press

Cited by

Sorted by citations

Related

Authors
Burkett, J. P., & Young, L. J.
Affiliations
Emory University (US)
Corresponding author
Young, Larry J.
ORCID
Venue
Psychopharmacology
Publisher
Springer Science and Business Media LLC
Published
2012-08-11
Volume
224
Issue
1
Pages
1-26
ISSN
  • 0033-3158 (ISSN-L)
  • 1432-2072
DOI
10.1007/s00213-012-2794-x
PMC
PMC3469771
OpenAlex
W2063959832
Semantic Scholar
f92f687ad0567900bb3e400905560e1c167e6bdf
Cited by
307
References
407
Citations by year
2026: 9 · 2025: 11 · 2024: 20 · 2023: 35 · 2022: 14 · 2021: 15 · 2020: 60 · 2019: 14 · 2018: 14 · 2017: 24 · 2016: 34 · 2015: 16 · 2014: 24 · 2013: 14 · 2012: 3
FWCI
2.58
Type
journal article
Language
en
Open access
Licence
www.springer.com/tdm (tdm)
Locations
  • Psychopharmacology (publishedVersion)
  • PubMed (publishedVersion)
  • Europe PMC (PubMed Central) (acceptedVersion)
  • PubMed Central (acceptedVersion)
Funders
  • National Institutes of Health (P51OD011132)
  • National Institute of Mental Health (MH-64692, K02MH064692)
  • National Center for Research Resources (P51RR000165)
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 › Social Psychology › Attachment and Relationship Dynamics
Keywords
social attachment, love, addiction, drug addiction, mesolimbic dopamine system, opioids, corticotropin-releasing factor, oxytocin, vasopressin, social reward, neurobiology of attachment
MeSH terms
Animals, Brain, Dopamine, Humans, Love, Object Attachment, Pair Bond, Parent-Child Relations, Reward, Social Behavior, Behavior, Addictive, Substance-Related Disorders

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 2026-10-08.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-08.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

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