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

In the paper

“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).”

What the sources say

• Not checked yetcites Sanchis-Segura and Spanagel 2006 source

In the paper

“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).”

What the sources say

• Not checked yetcites Swanson 1982 source
• Not checked yetcites Di Chiara and Imperato 1988 source
• Not checked yetcites Koob and Bloom 1988 source

In the paper

“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.”

What the sources say

• Not checked yetcites de Wit and Stewart 1981 source

In the paper

“Nonetheless, rodents will not self-administer selective D1R agonists, which, by themselves, induce conditioned place aversion (Woolverton et al 1984; Hoffman and Beninger 1988).”

What the sources say

• Not checked yetcites Woolverton et al 1984 source
• Not checked yetcites Hoffman and Beninger 1988

In the paper

“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).”

What the sources say

• Not checked yetcites Woolverton et al 1984 source
• Not checked yetcites Sanger 1986
• Not checked yetcites Woolverton 1986 source

In the paper

“Most rodents are not spontaneously maternal, and naïve females will often attack or ignore pups (Wiesner and Sheard 1933).”

What the sources say

• Not checked yetcites Wiesner and Sheard 1933

In the paper

“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).”

What the sources say

• Not checked yetcites Nakajima 1986
• Not checked yetcites Nakajima and Mckenzie 1986 source
• Not checked yetcites Beninger et al 1987 source

In the paper

“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).”

What the sources say

• Not checked yetcites Wiesner and Sheard 1933

In the paper

“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).”

What the sources say

• Not checked yetcites Nissen 1930 source
• Not checked yetcites Wilsoncroft 1969 source

In the paper

“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”

What the sources say

• Not checked yetcites Gaffori and Le Moal 1979 source
• Not checked yetcites Hansen et al 1991a
• Not checked yetcites Hansen et al 1991b

In the paper

“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).”

What the sources say

• Not checked yetcites Mogenson 1987
• Not checked yetcites Blackburn et al 1992 source
• Not checked yetcites Ikemoto and Panksepp 1999 source

In the paper

“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).”

What the sources say

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

In the paper

“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).”

What the sources say

• Not checked yetcites Ljungberg et al 1992 source
• Not checked yetcites Henry and White 1995 source
• Not checked yetcites Self et al 1996 source

In the paper

“D2R also seems to be necessary for maternal behavior, though the site of action has not been determined (Silva et al 2001).”

What the sources say

• Not checked yetcites Silva et al 2001 source

In the paper

“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).”

What the sources say

• Not checked yetcites Gaffori and Le Moal 1979 source
• Not checked yetcites Hansen et al 1991a
• Not checked yetcites Hansen et al 1991b

In the paper

“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.”

What the sources say

• Not checked yetcites Self et al 1996 source

In the paper

“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.)”

What the sources say

• Not checked yetcites Aragona and Wang 2009 source

In the paper

“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).”

What the sources say

• Not checked yetcites Wang et al 1999 source

In the paper

“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).”

What the sources say

• Not checked yetcites Thomas and Birney 1979 source
• Not checked yetcites Getz et al 1981 source
• Not checked yetcites Ahern et al 2011 source

In the paper

“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).”

What the sources say

• Not checked yetcites Williams et al 1992 source
• Not checked yetcites Winslow et al 1993 source
• Not checked yetcites Insel et al 1995 source

In the paper

“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).”

What the sources say

• Not checked yetcites Aragona et al 2003 source

In the paper

“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.”

What the sources say

• Not checked yetcites Beninger et al 1989 source

In the paper

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

What the sources say

• Not checked yetcites Carter and Roberts 1997

In the paper

“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).”

What the sources say

• Not checked yetcites Carter et al 1995 source

In the paper

“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).”

What the sources say

• Not checked yetcites Carter et al 1995 source

In the paper

“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).”

What the sources say

• Not checked yetcites Curtis and Wang 2007 source

In the paper

“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).”

What the sources say

• Not checked yetcites Wang et al 1999 source
• Not checked yetcites Gingrich et al 2000 source
• 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

“Sexually naïve male prairie voles are highly social and show very little aggressive behavior toward novel conspecifics (Insel et al 1995).”

What the sources say

• Not checked yetcites Insel et al 1995 source

In the paper

“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.”

What the sources say

• Not checked yetcites Aragona et al 2006 source

In the paper

“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).”

What the sources say

• Not checked yetcites Yudofsky et al 1987 source
• Not checked yetcites Buckley 1999

In the paper

“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.”

What the sources say

• Not checked yetcites Beninger et al 1989 source

In the paper

“Prairie voles are socially monogamous rodents indigenous to most of the Midwest United States and Canada (Tamarin 1985).”

What the sources say

• Not checked yetcites Tamarin 1985

In the paper

“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).”

What the sources say

• Not checked yetcites Liu et al 2010 source

In the paper

“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).”

What the sources say

• Not checked yetcites Burns et al 1983 source
• Not checked yetcites Tsunoo et al 1986 source
• Not checked yetcites North et al 1987 source

In the paper

“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).”

What the sources say

• Not checked yetcites Aragona et al 2006 source
• Not checked yetcites Liu et al 2010 source

In the paper

“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).”

What the sources say

• Not checked yetcites Goeders et al 1984 source
• Not checked yetcites Olmstead and Franklin 1997b

In the paper

“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).”

What the sources say

• Not checked yetcites Hughes et al 1975 source
• Not checked yetcites Birdsall and Hulme 1976 source
• Not checked yetcites Goldstein et al 1979 source

In the paper

“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).”

What the sources say

• Not checked yetcites Khachaturian et al 1985 source
• Not checked yetcites Robson et al 1985 source
• Not checked yetcites Mansour et al 1987

In the paper

“The A118G variant has enhanced binding and signaling properties, but low expression of mRNA (Zhang et al 2005; Kroslak et al 2007).”

What the sources say

• Not checked yetcites Zhang et al 2005 source
• Not checked yetcites Kroslak et al 2007 source

In the paper

“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).”

What the sources say

• Not checked yetcites Lee et al 2011 source
• Not checked yetcites Koller et al 2012

In the paper

“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).”

What the sources say

• Not checked yetcites O’Malley et al 1992
• Not checked yetcites Volpicelli et al 1992 source
• Not checked yetcites Greenway et al 2010 source

In the paper

“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).”

What the sources say

• Not checked yetcites Pfeiffer et al 1986 source
• Not checked yetcites Shippenberg et al 1987 source
• Not checked yetcites Pecina and Berridge 2000 source

In the paper

“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).”

What the sources say

• Not checked yetcites Dichiara and Imperato 1988 source
• Not checked yetcites Johnson and North 1992 source
• Not checked yetcites Scavone et al 2011 source

In the paper

“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).”

What the sources say

• Not checked yetcites Oslin et al 2003 source

In the paper

“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).”

What the sources say

• Not checked yetcites Hanson et al 1988 source
• Not checked yetcites Sivam 1989 source
• Not checked yetcites Spanagel et al 1990 source

In the paper

“Mice that lack the MOR gene do not develop place preferences or withdrawal symptoms in response to morphine (Matthes et al 1996).”

What the sources say

• Not checked yetcites Matthes et al 1996 source

In the paper

“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).”

What the sources say

• Not checked yetcites Way et al 2009 source
• Not checked yetcites Troisi et al 2011a

In the paper

“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).”

What the sources say

• Not checked yetcites Panksepp et al 1980 source
• Not checked yetcites Kalin et al 1988 source
• Not checked yetcites Martel et al 1993 source

In the paper

“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).”

What the sources say

• Not checked yetcites Moles et al 2004 source

In the paper

“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).”

What the sources say

• Not checked yetcites Panksepp et al 1980 source

In the paper

“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.”

What the sources say

• Not checked yetcites Gerfen and Young 1988 source

In the paper

“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).”

What the sources say

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

In the paper

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

In the paper

“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.”

In the paper

“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).”

What the sources say

• Not checked yetcites Dunn and Berridge 1990 source
• Not checked yetcites Heinrichs et al 1994 source
• Not checked yetcites Menzaghi et al 1994 source

In the paper

“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).”

What the sources say

• Not checked yetcites Kendrick and Keverne 1989 source
• Not checked yetcites Keverne and Kendrick 1994 source
• Not checked yetcites Shayit et al 2003 source

In the paper

“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).”

What the sources say

• Not checked yetcites Miller and Baum 1987 source
• Not checked yetcites Mehrara and Baum 1990 source
• Not checked yetcites Ismail et al 2009 source

In the paper

“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).”

What the sources say

• Not checked yetcites Burkett et al 2011 source

In the paper

“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.”

In the paper

“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).”

What the sources say

• Not checked yetcites Panksepp et al 1978

In the paper

“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).”

What the sources say

• Not checked yetcites Panksepp et al 1978
• Not checked yetcites Warnick et al 2005 source

In the paper

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

In the paper

“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.”

In the paper

“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).”

What the sources say

• Not checked yetcites Bartels and Zeki 2000 source
• Not checked yetcites Aron et al 2005 source
• Not checked yetcites Acevedo et al 2012 source

In the paper

“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).”

What the sources say

• Not checked yetcites DeVries et al 2002 source
• Not checked yetcites Lim et al 2007 source

In the paper

“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).”

What the sources say

• Not checked yetcites Sarnyai et al 1995 source
• Not checked yetcites Tucci et al 2003 source
• Not checked yetcites Knapp et al 2004 source

In the paper

“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).”

What the sources say

• Not checked yetcites DeVries et al 1996 source

In the paper

“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).”

What the sources say

• Not checked yetcites Pliakas et al 2001 source
• Not checked yetcites Kreibich et al 2009 source

In the paper

“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.”

In the paper

“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).”

What the sources say

• Not checked yetcites Gainer and Wray 1994
• Not checked yetcites Burbach et al 2005

In the paper

“However, a CRF-R2 agonist also blocked excessive alcohol consumption in dependent rats, suggesting opposite roles for these two receptors (Valdez et al 2004).”

What the sources say

• Not checked yetcites Valdez et al 2004 source

In the paper

“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).”

What the sources say

• Not checked yetcites Chen et al 2012b

In the paper

“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).”

What the sources say

• Not checked yetcites Grippo et al 2007a
• Not checked yetcites Grippo et al 2007b

In the paper

“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).”

What the sources say

• Not checked yetcites Lim et al 2005 source
• Not checked yetcites Lim et al 2006 source

In the paper

“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).”

What the sources say

• Not checked yetcites Wang et al 2005 source
• Not checked yetcites Chen et al 2012a
• Not checked yetcites Ungless et al 2003 source

In the paper

“This permanent social loss can be traumatic and cause both deterioration of physical health and susceptibility to depression (Prigerson et al 1997; Ott 2003).”

What the sources say

• Not checked yetcites Prigerson et al 1997 source
• Not checked yetcites Ott 2003 source

In the paper

“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.”

In the paper

“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).”

What the sources say

• Not checked yetcites Bosch et al 2009 source

In the paper

“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).”

What the sources say

• Not checked yetcites Ho et al 2001
• Not checked yetcites Takahashi et al 2001 source
• Not checked yetcites Valdez et al 2004 source

In the paper

“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).”

What the sources say

• Not checked yetcites Archer 1974
• Not checked yetcites van Kesteren et al 1992 source

In the paper

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

In the paper

“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).”

What the sources say

• Not checked yetcites Kovacs et al 1981 source
• Not checked yetcites Szabo et al 1988
• Not checked yetcites Carson et al 2010a

In the paper

“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).”

What the sources say

• Not checked yetcites Olazabal and Young 2006a
• Not checked yetcites Ross et al 2009b
• Not checked yetcites Keebaugh and Young 2011 source

In the paper

“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).”

What the sources say

• Not checked yetcites Liu and Wang 2003 source

In the paper

“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).”

What the sources say

• Not checked yetcites Bielsky et al 2004
• Not checked yetcites Bielsky et al 2005 source
• Not checked yetcites Lukas et al 2011 source

In the paper

“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).”

What the sources say

• Not checked yetcites Gainer and Wray 1994

In the paper

“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”

What the sources say

• Not checked yetcites Gainer and Wray 1994
• Not checked yetcites Burbach et al 2005
• Not checked yetcites Caldwell et al 2008 source

In the paper

“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).”

What the sources say

• Not checked yetcites Pedersen and Prange 1979 source
• Not checked yetcites Fahrbach et al 1985 source
• Not checked yetcites van Leengoed et al 1987 source

In the paper

“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).”

What the sources say

• Not checked yetcites Ferguson et al 2000 source

In the paper

“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).”

What the sources say

• Not checked yetcites Buijs et al 1983 source
• Not checked yetcites Alonso et al 1986 source
• Not checked yetcites Loup et al 1991 source