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

In the paper

“In prairie voles, pair bonding between mating partners is prevented if DA receptors in the NAC shell are non-specifically blocked (Wang et al 1999; Aragona et al 2003).”

What the sources say

• Not checked yetcites Wang et al 1999 source
• Partly supported

  Experiment 4a reports that haloperidol injected into the nucleus accumbens blocked mating-induced partner preferences in male prairie voles; Experiment 1a identifies haloperidol as a nonselective dopamine antagonist. The paper does not identify the Experiment 4a blockade specifically as a shell effect: its discussion locates apomorphine injections in the shell but says a core–shell distinction remains under investigation. No page number is available in our copy. (LLM, conf=0.84)

  cites Aragona et al 2003 sourceExperimental — accuracy not yet measured

In the paper

“In humans, a genetic variant of the OTR gene has been shown to correlate with pair bonding behavior in women (Walum et al 2012).”

What the sources say

• Supported

  The Abstract Results report that an OXTR variant, rs7632287, was associated with traits reflecting pair-bonding in women. The Discussion likewise describes an association between OXTR variation and behaviors reflecting pair-bonding in women. (LLM, conf=0.94)

  cites Walum et al 2012 source

In the paper

“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).”

In the paper

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

In the paper

“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).”

What the sources say

• Not checked yetcites Deroche-Gamonet et al 2004 source
• Not checked yetcites Wolffgramm and Heyne 1995 source
• Not checked yetcites Vanderschuren and Everitt 2004 source

In the paper

“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).”

In the paper

“a wide range of paradigms have been developed that model individual features of addiction behavior (Sanchis-Segura and Spanagel 2006).”

What the sources say

• Not checked yetcites Sanchis-Segura and Spanagel 2006 source

In the paper

“A deep and systematic concordance exists between the brain regions and neurochemicals involved in both addiction and social attachment.”

In the paper

“The strong overlap between characteristics of substance dependence and characteristics of social attachment suggests that attachment behavior may draw on the same psychological constructs, and perhaps the same biological substrates, as dependence on drugs of abuse.”

In the paper

“commonalities between compulsive disorders and substance addiction have been identified in terms of symptomatology, neurochemistry, and adaptations in brain function (Shaffer 1999; Holden 2001; Potenza 2006; Leeman and Potenza 2012).”

What the sources say

• Not checked yetcites Shaffer 1999 source
• Not checked yetcites Holden 2001 source
• Not checked yetcites Potenza 2006 source

In the paper

“However, when the diagnostic criteria for substance dependence are looked at side-by-side with related phenomena observable in normal human relationships, striking parallels emerge.”

In the paper

“Social attachment may be understood as a behavioral addiction, whereby the subject becomes addicted to another individual and the cues that predict social reward.”

In the paper

“DA release in the NAC is particularly important, and is correlated with human subjective ratings of drug reward and drug craving for many drugs of abuse (Volkow et al 1996a; Drevets et al 2001).”

What the sources say

• Not checked yetcites Volkow et al 1996a
• Not checked yetcites Drevets et al 2001 source

In the paper

“This suggests that, similar to drugs of abuse, one role of the mesolimbic DA pathway and of accumbal DA in particular is to generate the powerful motivational states that produce such dramatic maternal behaviors (reviewed in Numan 2007; for a comprehensive treatment of the subject, see Numan and Insel 2003).”

What the sources say

• Not checked yetcites Numan 2007 source
• Not checked yetcites Numan and Insel 2003

In the paper

“This same reduction in D2R is also seen in obesity, internet addiction, and trait impulsivity (Wang et al 2001; Buckholtz et al 2010; Kim et al 2011).”

What the sources say

• Not checked yetcites Wang et al 2001 source
• Not checked yetcites Buckholtz et al 2010 source
• Not checked yetcites Kim et al 2011 source

In the paper

“DA is released naturally in the NAC of maternal rats during interactions with pups (Hansen et al 1993).”

What the sources say

• Not checked yetcites Hansen et al 1993 source

In the paper

“Under normal conditions, a synergistic balance of D1R- and D2R-like activation exists in striatal regions (Walters et al 1987; LaHoste et al 1993; Gerfen et al 1995; Wise et al 1996; Hu and White 1997).”

What the sources say

• Not checked yetcites Walters et al 1987 source
• Not checked yetcites LaHoste et al 1993 source
• Not checked yetcites Gerfen et al 1995 source

In the paper

“Rat mothers even prefer young pups to cocaine, indicating the power of their motivation toward maternal care (Mattson et al 2001).”

What the sources say

• Not checked yetcites Mattson et al 2001 source

In the paper

“Adult attachments between mating partners are relatively rare in mammals, occurring in only 3-5% of mammalian species (Orians 1969; Kleiman 1977).”

What the sources say

• Not checked yetcites Orians 1969 source
• Not checked yetcites Kleiman 1977 source

In the paper

“PET studies show that D2R availability is decreased after long-term exposure to many different drugs of abuse (Volkow et al 1996b; Volkow et al 1997; Wang et al 1997a; Volkow et al 2001; Fehr et al 2008; reviewed in Volkow et al 2009).”

What the sources say

• Not checked yetcites Volkow et al 1996b
• Not checked yetcites Volkow et al 1997 source
• Not checked yetcites Wang et al 1997a

In the paper

“animal studies have shown that DA is released in this pathway in response to a wide variety of rewards, including sex, food, water, and intracranial self-stimulation (Cooper and Breese 1975; Hernandez and Hoebel 1988; Damsma et al 1992; Yoshida et al 1992; Young et al 1992).”

What the sources say

• Not checked yetcites Cooper and Breese 1975
• Not checked yetcites Hernandez and Hoebel 1988 source
• Not checked yetcites Damsma et al 1992 source

In the paper

“These two classes of DA receptor also seem to have distinct roles in the response to drugs of abuse.”

In the paper

“DA in the mesolimbic pathway also has a role in maternal behavior.”

In the paper

“The same chronic cocaine exposure is associated with decreases in tonic D2R signaling, and these decreases are involved in drug withdrawal, compulsive intake, and reinstatement of drug-seeking behavior (Nestler et al 1990; Self et al 1996; Perez et al 2011; Grieder et al 2012).”

What the sources say

• Not checked yetcites Nestler et al 1990 source
• Not checked yetcites Self et al 1996 source
• Not checked yetcites Perez et al 2011 source

In the paper

“Furthermore, if cocaine is administered with D2R-like antagonists, it loses its reinforcing effects (Woolverton and Virus 1989; Bachtell et al 2005; Claytor et al 2006; Peng et al 2009).”

What the sources say

• Not checked yetcites Woolverton and Virus 1989 source
• Not checked yetcites Bachtell et al 2005 source
• Not checked yetcites Claytor et al 2006 source

In the paper

“This body of evidence suggests that a disruption in the balance of D1R- and D2R-like pathways in favor of D1R may underlie the behavioral changes seen in addiction and other impulse control disorders.”

In the paper

“When D1R is blocked with antagonist in the NAC shell of pair-bonded males, selective aggression toward an unfamiliar female is greatly reduced and affiliative behaviors are increased (Aragona et al 2006).”

What the sources say

• Not checked yetcites Aragona et al 2006 source

In the paper

“As D1R is upregulated during pair bonding and D2R is stable, this plastic change represents an alteration in the balance of D1R/D2R signaling in the striatum in favor of D1R, similar to what is seen in human PET studies of drug addiction (Volkow et al 2009).”

What the sources say

• Not checked yetcites Volkow et al 2009 source

In the paper

“There is strong overlap between these findings and the literature on drug addiction.”

In the paper

“DA also plays a prominent role in pair bond maintenance.”

In the paper

“These data suggest that the plastic change in D1R that occurs during pair bonding is causative in the development of selective aggression behavior, which helps to prevent the formation of a new pair bond with a new potential mate. (These topics are reviewed in Aragona and Wang 2009.)”

What the sources say

• Not checked yetcites Aragona and Wang 2009 source

In the paper

“Furthermore, non-specific activation of DA receptors in the NAC shell is sufficient to induce pair bonding, even if no mating occurs.”

In the paper

“Conversely, pharmacological activation of D1R prevents pair bond formation, with or without concurrent activation of D2R (Aragona et al 2006).”

What the sources say

• Not checked yetcites Aragona et al 2006 source

In the paper

“Like with drugs of abuse, mesolimbic DA is a major contributor to the formation of pair bonds in prairie voles, and particularly in the NAC shell region.”

In the paper

“In the context of pair bonding, the aversion induced by pure pharmacological D1R activation may associate with social cues, leading to a “conditioned partner aversion” that could explain the disruption of pair bonding.”

In the paper

“There is an exceptionally strong parallel between these plastic changes from pair bonding, and the plastic changes seen in drug addiction.”

In the paper

“Blockade of D1R neither enhances nor prevents pair bonding (Wang et al 1999; Aragona et al 2006; Curtis et al 2006).”

What the sources say

• Not checked yetcites Wang et al 1999 source
• Not checked yetcites Aragona et al 2006 source
• Not checked yetcites Curtis et al 2006 source

In the paper

“However, when males have cohabitated with females for two weeks, they develop intense selective aggression toward male and female strangers and not toward their female partners (Insel et al 1995; Wang et al 1997b).”

What the sources say

• Not checked yetcites Insel et al 1995 source
• Not checked yetcites Wang et al 1997b

In the paper

“However, despite these non-specific effects, the two DA receptor types seem to play opposite roles in pair bonding (Fig. 1).”

In the paper

“Mating has been shown to cause DA release in the NAC in rodents (Damsma et al 1992; Gingrich et al 2000).”

What the sources say

• Not checked yetcites Damsma et al 1992 source
• Not checked yetcites Gingrich et al 2000 source

In the paper

“Over the course of 2 weeks of cohabitation, while selective aggression behavior is forming, the mesolimbic DA system in male prairie voles undergoes plastic changes (Aragona et al 2006).”

What the sources say

• Not checked yetcites Aragona et al 2006 source

In the paper

“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).”

What the sources say

• Not checked yetcites Getz and Hofmann 1986 source
• Not checked yetcites Ross and Young 2009

In the paper

“More directly, administering amphetamines to prairie voles daily for 3 days causes an upregulation of D1R in the NAC that mimics the upregulation seen in pair bonding (Liu et al 2010).”

What the sources say

• Not checked yetcites Liu et al 2010 source

In the paper

“The endogenous opiate system is well-known to modulate, in part, the rewarding and reinforcing effects of food, water, sex, intracranial self-stimulation, and other rewards (Broekkamp and Phillips 1979; Turkish and Cooper 1983; West et al 1983; Agmo and Berenfeld 1990; Yeomans and Gray 1996; Gerrits et al 2003).”

What the sources say

• Not checked yetcites Broekkamp and Phillips 1979 source
• Not checked yetcites Turkish and Cooper 1983 source
• Not checked yetcites West et al 1983 source

In the paper

“KOR is also involved in addiction-related processes.”

In the paper

“Similarly, the rewarding or reinforcing value of many drugs of abuse can be blocked by MOR antagonists injected into the NAC, the VP, or the VTA (Hiroi and White 1993; Skoubis and Maidment 2003; Soderman and Unterwald 2008).”

What the sources say

• Not checked yetcites Hiroi and White 1993 source
• Not checked yetcites Skoubis and Maidment 2003 source
• Not checked yetcites Soderman and Unterwald 2008 source

In the paper

“While opiate drugs are themselves strongly addictive, the role of the endogenous opioid system in addiction is not limited to these, as this system plays a strong role in the development of addiction to every class of drug (Altshuler et al 1980; Karras and Kane 1980; De Vry et al 1989; Kuzmin et al 1997; Ismayilova and Shoaib 2010; for a review see van Ree et al, 1999).”

What the sources say

• Not checked yetcites Altshuler et al 1980 source
• Not checked yetcites Karras and Kane 1980 source
• Not checked yetcites De Vry et al 1989 source

In the paper

“During withdrawal, this up-regulation and subsequent activation of KOR may contribute to negative affect and thus promote relapse (Przewlocka et al 1997; Simonin et al 1998; Walker and Koob 2008).”

What the sources say

• Not checked yetcites Przewlocka et al 1997 source
• Not checked yetcites Simonin et al 1998 source
• Not checked yetcites Walker and Koob 2008 source

In the paper

“Opioids are involved in every stage of the addiction process, including initiation, maintenance, withdrawal, and relapse (Gerrits et al 2003).”

What the sources say

• Not checked yetcites Gerrits et al 2003 source

In the paper

“Based on animal studies, the role of opioids in the positive reinforcing effects of natural rewards and drugs of abuse has largely been attributed to MOR in the NAC, VP, and VTA, though some part is also played by DOR (van Ree and de Wied 1980; Shippenberg et al 1987; Hubner and Koob 1990; Olmstead and Franklin 1997a; Corrigall et al 2000; Pecina and Berridge 2000; Van Ree et al 2000; Gerrits et”

What the sources say

• Not checked yetcites van Ree and de Wied 1980 source
• Not checked yetcites Shippenberg et al 1987 source
• Not checked yetcites Hubner and Koob 1990 source

In the paper

“Human studies have largely corroborated the link between MOR and positive reinforcement from natural rewards and drugs of abuse.”

In the paper

“Opioid antagonists effectively reverse the effects of opiate drugs in humans (Bradberry and Raebel 1981).”

What the sources say

• Not checked yetcites Bradberry and Raebel 1981 source

In the paper

“Rhesus macaques also have a functionally similar variant of the MOR gene, C77G, which also results in increased alcohol intake (Barr et al 2007).”

What the sources say

• Not checked yetcites Barr et al 2007 source

In the paper

“Furthermore, the changes in D1R signaling that occur during pair bonding act directly to decrease the reinforcing effects of amphetamines (Liu et al 2011).”

What the sources say

• Not checked yetcites Liu et al 2011 source

In the paper

“Humans with the analogous A118G polymorphism are susceptible to developing fearful attachment style in response to low maternal care (Troisi et al 2011b).”

What the sources say

• Not checked yetcites Troisi et al 2011b

In the paper

“Some evidence also suggests that these effects may be specific to the MOR, at least with respect to infant-mother attachment.”

In the paper

“Additionally, they localized this effect to the NAC shell, where a KOR antagonist (but not a MOR antagonist) abolished selective aggression.”

In the paper

“The authors first showed that a peripherally administered KOR antagonist, but not a MOR-preferential antagonist, prevents the expression of selective aggression in voles that have already formed pair bonds.”

In the paper

“The majority of mammalian species are promiscuous breeders that do not form selective partner preferences toward specific mating partners (Kleiman 1977).”

What the sources say

• Not checked yetcites Kleiman 1977 source

In the paper

“However, a MOR-selective antagonist administered into the caudate-putamen (CP), but not the NAC shell, prevents pair bonding without affecting sexual behavior.”

In the paper

“This hypothesis has subsequently been supported by work on social motivation in rats and Rhesus macaques (Panksepp et al 1985; Martel et al 1993; Martel et al 1995).”

What the sources say

• Not checked yetcites Panksepp et al 1985 source
• Not checked yetcites Martel et al 1993 source
• Not checked yetcites Martel et al 1995 source

In the paper

“This CRF system coordinates stress responding on several levels, including behavior, autonomic response, and the HPA axis.”

What the sources say

• Not checked yetcites Bale and Vale 2004 source

In the paper

“These experiments demonstrate that MOR and the CP are necessary for pair bond formation in this species, and the roles of both seem to be conserved in humans.”

In the paper

“As such, KOR may be a mechanism for maintenance of drug use (for a review of KOR in addiction, see Bruijnzeel 2009).”

What the sources say

• Not checked yetcites Bruijnzeel 2009 source

In the paper

“These and other observations led to the opioid hypothesis of social attachment, which posits that opioid receptors mediate both social reward and social motivation (Herman and Panksepp 1978).”

What the sources say

• Not checked yetcites Herman and Panksepp 1978 source

In the paper

“Conversely, long-term exposure to opioid antagonists decreases maternal competence and motivation, suggesting that mothers adapt their behavior based on reduced social reward (Martel et al 1993; Martel et al 1995).”

What the sources say

• Not checked yetcites Martel et al 1993 source
• Not checked yetcites Martel et al 1995 source

In the paper

“This suggests that the two receptor systems are acting in a coordinated fashion in the striatum to modulate different aspects of pair bonding (Resendez et al 2012).”

What the sources say

• Not checked yetcites Resendez et al 2012 source

In the paper

“Low-dose morphine also reduces social contact in guinea pigs and rats (Herman and Panksepp 1978; Panksepp et al 1979).”

What the sources say

• Not checked yetcites Herman and Panksepp 1978 source
• Not checked yetcites Panksepp et al 1979 source

In the paper

“Studies in Rhesus macaques show similar effects of acute administration of agonists and antagonists on maternal care (Kalin et al 1995).”

What the sources say

• Not checked yetcites Kalin et al 1995 source

In the paper

“suggesting that mothers adapt their behavior based on reduced social reward”

In the paper

“Additionally, the same polymorphisms of the MOR gene that are associated with increased risk for alcoholism in humans and Rhesus macaques are also related to maternal behavior in both species (Barr et al 2007; Barr et al 2008; Higham et al 2011; Troisi et al 2011b).”

What the sources say

• Not checked yetcites Barr et al 2007 source
• Not checked yetcites Barr et al 2008 source
• Not checked yetcites Higham et al 2011 source

In the paper

“The formation of these non-social preferences is prevented by peripheral administration of non-selective opioid antagonists.”

In the paper

“For instance, in chicks, a reduction in DVs was only observed when a MOR-selective agonist was used (Warnick et al 2005).”

What the sources say

• Not checked yetcites Warnick et al 2005 source

In the paper

“and the roles of both seem to be conserved in humans”

In the paper

“In turn, increased stress and withdrawal symptoms increase drug craving, generating a powerful motivation to continue use or to relapse from abstinence (Hershon 1977; Cooney et al 1997; Sinha et al 2000).”

What the sources say

• Not checked yetcites Hershon 1977 source
• Not checked yetcites Cooney et al 1997 source
• Not checked yetcites Sinha et al 2000 source

In the paper

“CRF is also involved in the formation of pair bonds.”

In the paper

“These data also suggest that the ability of CRF-R1 antagonists to block excessive consumption is related to the ability of these drugs to block the aversive aspects of withdrawal (Koob and Zorrilla 2010).”

What the sources say

• Not checked yetcites Koob and Zorrilla 2010

In the paper

“In the wild, when one member of a mating pair is lost, the surviving member typically will not take on a new partner for the duration of his or her life (Getz et al 1981).”

What the sources say

• Not checked yetcites Getz et al 1981 source

In the paper

“Nonselective CRF antagonists, as well as CRF-R1 antagonists, selectively block excessive consumption of several drugs of abuse in dependent rats, but not in non-dependent rats (Valdez et al 2004; Funk et al 2006; Funk et al 2007).”

What the sources say

• Not checked yetcites Valdez et al 2004 source
• Not checked yetcites Funk et al 2006 source
• Not checked yetcites Funk et al 2007 source

In the paper

“This research shows that CRF receptors, particularly in the amygdala, are important in mediating the motivational effects of withdrawal from drugs.”

In the paper

“In drug addiction, CRF is primarily involved in withdrawal from drugs of abuse (Koob and Kreek 2007; Koob 2008).”

What the sources say

• Not checked yetcites Koob and Kreek 2007 source
• Not checked yetcites Koob 2008 source

In the paper

“CRF in the NAC shell can also enhance the positive motivational value and salience of incentive cues (Pecina et al 2006; Kreibich et al 2009).”

What the sources say

• Not checked yetcites Pecina et al 2006 source
• Not checked yetcites Kreibich et al 2009 source

In the paper

“Furthermore, it may be that CRF release during separation from the partner creates an aversive, stressful state that motivates the prairie vole to return to the partner, which in turn maintains the pair bond (Bosch et al 2009).”

What the sources say

• Not checked yetcites Bosch et al 2009 source

In the paper

“CRF production in, and release from, the amygdala is greatly potentiated during withdrawal from a variety of drugs of abuse, as well as during chronic stress (Merlo Pich et al 1995; Rodriguez de Fonseca et al 1997; Richter and Weiss 1999; Stout et al 2000; Zorrilla et al 2001; Olive et al 2002; Funk et al 2006; George et al 2007).”

What the sources say

• Not checked yetcites Merlo Pich et al 1995 source
• Not checked yetcites Rodriguez de Fonseca et al 1997 source
• Not checked yetcites Richter and Weiss 1999

In the paper

“Some of the stress-related effects of CRF in the NAC shell may be DA-dependent; recall that DA is also modulated by acute and chronic stressors (Miczek et al 2011; Wang and Tsien 2011; Chen et al 2012b).”

What the sources say

• Not checked yetcites Miczek et al 2011 source
• Not checked yetcites Wang and Tsien 2011 source
• Not checked yetcites Chen et al 2012b

In the paper

“This literature also suggests that the role of CRF in pair bonding is principally focused on the NAC shell.”

In the paper

“Specifically, males separated from their partner display robust increases in immobility and hanging in the forced swim test and tail suspension tests, respectively.”

In the paper

“However, the research discussed above strongly implicates the extended amygdala as the source of CRF-containing neurons mediating both drug dependence and relapse.”

In the paper

“The facilitative effects of acute stressors on male pair bonding are directly analogous to the facilitative effects of acute stressors on drug-taking behavior, a study which, interestingly, was also performed in male rodents (Miczek et al 2011).”

What the sources say

• Not checked yetcites Miczek et al 2011 source

In the paper

“This depressive-like behavior is reversed by CRF-R1 or CRF-R2 antagonists injected into the cerebral ventricle.”

In the paper

“Monogamous prairie voles have significantly greater OTR density in the prefrontal cortex (the cortical component of the mesolimbic DA pathway), as CP, and NAS than do non-monogamous vole species (Insel and Shapiro 1992).”

What the sources say

• Not checked yetcites Insel and Shapiro 1992 source

In the paper

“In general, OT appears to play a modulatory role in many aspects of drug addiction.”

In the paper

“This suggests the possibility that the baseline activity of the endogenous OT system in these paradigms is not sufficient to affect drug-related behaviors.”

In the paper

“Nonetheless, there is a relative lack of studies showing that OT antagonists alone affect these processes (but see Kovacs et al 1987).”

What the sources say

• Not checked yetcites Kovacs et al 1987

In the paper

“This exogenous OT acts in the NAC to reduce both formation and expression of tolerance to some of the behavioral and physiological effects of drugs of abuse (Krivoy et al 1974; Van Ree and De Wied 1977; Kovacs et al 1981; Kovacs and Vanree 1985; Ibragimov et al 1987; Kovacs and Telegdy 1987; Szabo et al 1989; Sarnyai et al 1992).”

What the sources say

• Not checked yetcites Krivoy et al 1974 source
• Not checked yetcites Van Ree and De Wied 1977 source
• Not checked yetcites Kovacs et al 1981 source

In the paper

“In many of these studies, the effects of exogenously administered OT and OT agonists were reversed with OT antagonists in the brain, showing the specificity of the results to central OT.”

In the paper

“OTR activation in the NAC and prefrontal cortex, but not the CP, is necessary for the formation of pair bonds in female prairie voles (Young et al 2001).”

What the sources say

• Not checked yetcites Young et al 2001 source

In the paper

“Taken together, these data demonstrate a role for OT and AVP in the selective processing of social aspects of memory (for reviews, see Bielsky and Young 2004; Wacker and Ludwig 2012).”

What the sources say

• Not checked yetcites Bielsky and Young 2004
• Not checked yetcites Wacker and Ludwig 2012 source

In the paper

“The dual central and peripheral role for OT in the onset of maternal behavior, and particularly in the formation of mother-offspring attachments, suggests that coordinated peripheral and central OT release during labor and nursing act to induce bonding between mother and infant (Ross and Young 2009; Feldman 2012).”

What the sources say

• Not checked yetcites Ross and Young 2009
• Not checked yetcites Feldman 2012 source

In the paper

“and furthermore, that pair bonding is mediated by neural systems that were elaborated in evolution from circuits originally adapted for maternal behavior (Ross et al 2009a).”

What the sources say

• Not checked yetcites Ross et al 2009a

In the paper

“Early expression of maternal behavior in virgin rodents is correlated with OTR density in the NAC shell, the lateral septum (LS) and parts of the extended amygdala, all regions implicated in pair bonding and in the regulation of motivated behaviors (Champagne et al 2001; Sheehan et al 2004; Olazabal and Young 2006b; Modi and Young 2011).”

What the sources say

• Not checked yetcites Champagne et al 2001 source
• Not checked yetcites Sheehan et al 2004 source
• Not checked yetcites Olazabal and Young 2006b

In the paper

“The onset of maternal behavior can be prevented by injection of an OTR antagonist into the VTA (Pedersen et al 1994).”

What the sources say

• Not checked yetcites Pedersen et al 1994 source

In the paper

“However, OT does not appear to be necessary for the maintenance of maternal behavior in experienced females (Fahrbach et al 1985).”

What the sources say

• Not checked yetcites Fahrbach et al 1985 source

In the paper

“Central OT is also sufficient to induce selective mother-offspring bonds in sheep (Kendrick et al 1987).”

What the sources say

• Not checked yetcites Kendrick et al 1987 source

In the paper

“The convergence of evidence from the fields of attachment and addiction reveals an inexorable series of parallels (see Fig. 4, Table 2).”

In the paper

“These studies lend support to the hypothesis that OT release during human romantic activities serves to induce the formation of long-term bonds (Young et al 2005; Feldman 2012).”

What the sources say

• Not checked yetcites Young et al 2005 source
• Not checked yetcites Feldman 2012 source

In the paper

“First, the early, euphoric excitement that comes with new relationships subsides, and this euphoria is gradually replaced by a more subdued sense of contentment. This first effect can be understood as tolerance to the addictive partner; tolerance to opioids may contribute to this effect as well.”

In the paper

“Simultaneously, CRF stress circuitry is primed for maintenance through the up-regulation of CRF peptide in the extended amygdala.”