Sexually dimorphic vasopressin cells modulate social investigation and communication in sex-specific ways

“First, we tested mice on an EPM to test for anxiety-related behavior (Lister, 1987).”

“Although central AVP has been repeatedly implicated in the generation of anxiety states (Ebner et al., 2002; Bielsky et al., 2004; Raggenbass, 2008), we did not observe changes in anxiety-like behavior of lesioned males or females in a non-social anxiogenic environment (Hogg, 1996).”

“Introduction Social behavior and communication show profound sex differences in many species (Darwin, 1871; Bradbury and Vehrencamp, 1998), and in humans, dysfunction in social behavior and communication is prominent in chronic, debilitating, and pervasive psychopathologies (Insel, 2010) that show sex differences in p…”

“Estrous state was verified by color, swelling, and expanded size of vaginal opening (Caligioni, 2009).”

“While several studies have directly tested the social function of sexually-dimorphic cell populations (Yang et al., 2013; Scott et al., 2015; Unger et al., 2015; Wei et al., 2018), only one has found a strong sex difference in social behavior tied to an anatomic sexual dimorphism, in this case a female-biased one (Sco…”

“For example, AVP has been implicated in aggression, pair bonding, maternal behavior, and communication across vertebrates (Goodson and Bass, 2001; Albers, 2012).”

“Lesioning AVP cells also removed all other neuroactive substances co-expressed by these cells, for example, galanin (Miller et al., 1993), which also has been implicated in control of social behavior (Wu et al., 2014).”

“USV, UM, and social investigation were recorded in an acrylic three-chamber apparatus (Crawley, 2007; Arakawa et al., 2008a; Moy et al., 2009; Harvard Apparatus; dimensions: 20.3 42 22 cm).”

“Although central AVP has been repeatedly implicated in the generation of anxiety states (Ebner et al., 2002; Bielsky et al., 2004; Raggenbass, 2008), we did not observe changes in anxiety-like behavior of lesioned males or females in a non-social anxiogenic environment (Hogg, 1996).”

“In rats, where AVP release in the septum, one of the most prominent projection areas of BNST and MeA AVP cells (De Vries and Buijs, 1983; De Vries and Panzica, 2006), correlates positively with intermale aggression in a resident-intruder test, a behavioral response could be blocked by…”

“Given the metabolic costs of urination (Gosling et al., 2000) and the increased risks associated with attraction of aggressive competitors and predators toward urine marks (Desjardins et al., 1973; January/February 2019, 6(1) e0415-18.”

“These mice were generated using a bacterial artificial chromosome (BAC) that expressed codon-improved Cre recombinase (Shimshek et al., 2002) under the transcriptional control of the AVP promoter (AVP-iCre mice).”

“All subject males attacked the intruder male stimulus by the second encounter, and all subordinate stimulus males displayed submissive behavior, defined as defensive postures (e.g., on-back), fleeing, and non-social exploring (Koolhaas et al., 2013).”

“This is somewhat surprising because AVP in the LS, an important target of the BNST AVP cells (De Vries and Panzica, 2006), controls anxious states.”

“For example, injections of AVP or V1aR antagonists, or V1aR knockdown within LS all support the idea that AVP is anxiogenic at this site (Landgraf et al., 1995; Liebsch et al., 1996; Beiderbeck et al., 2007).”

“In addition, injecting specific V1a receptor agonists or boosting V1a receptor expression in target areas of AVP cells in the BNST and MeA promotes affiliation in voles (Wang et al., 1994; Liu et al., 2001; Pitkow et al., 2001; Lim and Young, 2004; Lim et al., 2004) and social recognition and active social behaviors i…”

“For example, AVP has been implicated in aggression, pair bonding, maternal behavior, and communication across vertebrates (Goodson and Bass, 2001; Albers, 2012).”

“In addition, injecting specific V1a receptor agonists or boosting V1a receptor expression in target areas of AVP cells in the BNST and MeA promotes affiliation in voles (Wang et al., 1994; Liu et al., 2001; Pitkow et al., 2001; Lim and Young, 2004; Lim et al., 2004) and social recognition and active social behaviors i…”

“To directly test the hypothesis that AVP cells in the BNST modulate social behavior, we injected an adeno-associated virus (AAV) with a Cre-dependent, genetically modified executioner caspase-3 complex (Yang et al., 2013; Unger et al., 2015) into the BNST of adult AVP-iCre and AVP-iCre– (Mieda et al.”

“, 2004) and social recognition and active social behaviors in rats (Dantzer et al., 1988; Veenema et al., 2012).”

“While several studies have directly tested the social function of sexually-dimorphic cell populations (Yang et al., 2013; Scott et al., 2015; Unger et al., 2015; Wei et al., 2018), only one has found a strong sex difference in social behavior tied to an anatomic sexual dimorphism, in this case a…”

“, 2015) into the BNST of adult AVP-iCre and AVP-iCre– (Mieda et al., 2015) male and female mice, which specifically deleted AVP cells in the area, and tested the effects of these deletions on social investigation, courtship ultrasonic vocalizations (USVs; Chabout et al.”

“While several studies have directly tested the social function of sexually-dimorphic cell populations (Yang et al., 2013; Scott et al., 2015; Unger et al., 2015; Wei et al., 2018), only one has found a strong sex difference in social behavior tied to an anatomic sexual dimorphism, in this case a female-biased one (Sco…”

“Recordings were then analyzed using a MATLAB (MathWorks, RRID:SCR_001622) plug-in that automates USV analysis (Van Segbroeck et al., 2017).”

“For example, male rats and mice have about two to three times as many AVP cells as females in these nuclei and the projections of these cells to areas such as lateral septum (LS) are denser as well (van Leeuwen et al., 1985; Rood and De Vries, 2011; Rood et al., 2013; Otero-Garcia et al., 2014).”

“…that one function of AVP cells in the BNST is to generate male-typical approach, investigation, and assessment of potential territorial competitors (Oldfield et al., 2015), which is consistent with the observation that knocking down AVT mRNA in the BNST of territorial birds reduces social contact…”

“USV, UM, and social investigation were recorded in an acrylic three-chamber apparatus (Crawley, 2007; Arakawa et al., 2008a; Moy et al., 2009; Harvard Apparatus; dimensions: 20.3 42 22 cm).”

“, 2008b), all aspects of mouse communication known to show pronounced sex differences (Crawley, 2012; Lehmann et al., 2013; Wöhr, 2014).”

“, 2015) male and female mice, which specifically deleted AVP cells in the area, and tested the effects of these deletions on social investigation, courtship ultrasonic vocalizations (USVs; Chabout et al., 2015), and territorial urine marking (UM; Arakawa et al.”

“16-mm outer diameter; Dow Corning Corporation) containing estradiol benzoate (E; diluted 1:1 with cholesterol) were implanted subcutaneously in the scapular region immediately following ovariectomy (GDX E; Bakker et al., 2002; Ström et al., 2012).”

“CD1(ICR) (Charles River Laboratories) mice were used as stimuli for behavioral testing and to provide male and female subjects with social experience because strain differences between subjects and stimulus mice increase social investigation (Gheusi et al., 1994).”

“In rats, where AVP release in the septum, one of the most prominent projection areas of BNST and MeA AVP cells (De Vries and Buijs, 1983; De Vries and Panzica, 2006), correlates positively with intermale aggression in a resident-intruder test, a behavioral response could be blocked by retro-dialysis of an AVP antagoni…”

“For example, injections of AVP or V1aR antagonists, or V1aR knockdown within LS all support the idea that AVP is anxiogenic at this site (Landgraf et al., 1995; Liebsch et al., 1996; Beiderbeck et al., 2007).”

“Animals were tracked by ANY-maze so that measurements of time spent in open and closed arms were recorded automatically whereas the number of risk assessment behaviors (stretch-attend posture, headdips) were manually scored from video (Cole and Rodgers, 1993).”

“This is consistent with the few studies examining male courtship vocalizations (in birds) that report no effect of central infusions of AVT or V1a antagonists on femaledirected singing (Goodson and Adkins-Regan, 1999; Goodson and Evans, 2004) or got mixed results from antisense knock-down of BNST-AVP, with one study f…”

“A particularly well-positioned circuit in this respect is the arginine vasopressin (AVP) innervation of the brain, which shows marked sex differences across many species, including humans (Goodson and Bass, 2001; De Vries and Panzica, 2006; de Vries, 2008).”

“In rats, a V1a antagonist reduces social play in males while increasing them in females when injected intracerebroventricularly, but has the exact opposite effect when injected into the septum (Veenema et al., 2013).”

“, 2015), which is consistent with the observation that knocking down AVT mRNA in the BNST of territorial birds reduces social contact with other males (Kelly and Goodson, 2013b), and that overexpression of V1a receptors (Landgraf et al., 2003) or AVP injections (Koolhaas et al.”

“USV syllable types were identified from a subset of males (iCre– n 6; iCre n 7) using criterion previously described: short, composite, downward, upward, 1 frequency jump, modulated, multiple frequency jumps, u-shape, flat, chevron (Hanson and Hurley, 2012).”

“Indeed, long-term pharmacological reduction of V1a receptor activity in the LS, a key target of AVP cells in the BNST (De Vries and Panzica, 2006), produces different changes in behavior than acute receptor blockade (Liebsch et al., 1996; Everts and Koolhaas, 1999).”

“For example, the density of AVP fibers in BNST and MeA projection areas and c-Fos activation in AVP cells in the BNST correlate negatively with aggression in male mice and rats (Compaan et al., 1993; Everts et al., 1997; Ebner et al., 2000; Beiderbeck et al., 2007; Veenema et al., 2010) but positively with prosocial b…”

“In addition, injecting specific V1a receptor agonists or boosting V1a receptor expression in target areas of AVP cells in the BNST and MeA promotes affiliation in voles (Wang et al., 1994; Liu et al., 2001; Pitkow et al., 2001; Lim and Young, 2004; Lim et al., 2004) and social recognition and active social behaviors i…”

“Removal of these cells in males did not alter copulatory behavior, which is consistent with the absence of copulatory deficits in whole-animal knock-outs of V1a or V1b receptors (Wersinger et al., 2004, 2007).”

“…fibers in BNST and MeA projection areas and c-Fos activation in AVP cells in the BNST correlate negatively with aggression in male mice and rats (Compaan et al., 1993; Everts et al., 1997; Ebner et al., 2000; Beiderbeck et al., 2007; Veenema et al., 2010) but positively with prosocial behavior…”

“Although AVP has been implicated in female rat sexual behavior, the pattern of prior results is opposite to our findings: in rats, AVP reduced, whereas V1a receptor antagonists increased, female sexual behavior in rats (Södersten et al., 1983; Pedersen and Boccia, 2006).”

“Removal of these cells in males did not alter copulatory behavior, which is consistent with the absence of copulatory deficits in whole-animal knock-outs of V1a or V1b receptors (Wersinger et al., 2004, 2007).”

“, 2004) and social recognition and active social behaviors in rats (Dantzer et al., 1988; Veenema et al., 2012).”

“For example, the density of AVP fibers in BNST and MeA projection areas and c-Fos activation in AVP cells in the BNST correlate negatively with aggression in male mice and rats (Compaan et al., 1993; Everts et al., 1997; Ebner et al., 2000; Beiderbeck et al., 2007; Veenema et al., 2010) but positively with prosocial b…”

“Odor discrimination We used a habituation-discrimination procedure on a subset of subjects to test whether they could distinguish between social odors (Baum and Keverne, 2002).”

“In hamsters, hypothalamic injections of vasopressin stimulate aggression in males, while reducing it in females (Terranova et al., 2016).”

“…1998), it may simply mean that, as with aggressive behavior, AVP cells in the BNST are not critically involved in anxiety-like behaviors, and that AVP derived from other sources, such as the PVN or the amygdala (Rood et al., 2013), also drive AVP’s anxiogenic action in the septum and elsewhere.”

“16-mm outer diameter; Dow Corning Corporation) containing estradiol benzoate (E; diluted 1:1 with cholesterol) were implanted subcutaneously in the scapular region immediately following ovariectomy (GDX E; Bakker et al., 2002; Ström et al., 2012).”

“Given the metabolic costs of urination (Gosling et al., 2000) and the increased risks associated with attraction of aggressive competitors and predators toward urine marks (Desjardins et al., 1973; January/February 2019, 6(1) e0415-18.”

“, 2008b), all aspects of mouse communication known to show pronounced sex differences (Crawley, 2012; Lehmann et al., 2013; Wöhr, 2014).”

“…of AVT or V1a antagonists on femaledirected singing (Goodson and Adkins-Regan, 1999; Goodson and Evans, 2004) or got mixed results from antisense knock-down of BNST-AVP, with one study finding no effect (Kelly et al., 2011) and another observing decreased singing (Kelly and Goodson, 2013b).”

“Although central AVP has been repeatedly implicated in the generation of anxiety states (Ebner et al., 2002; Bielsky et al., 2004; Raggenbass, 2008), we did not observe changes in anxiety-like behavior of lesioned males or females in a non-social anxiogenic environment (Hogg, 1996).”

“AVP cells in the medial amygdala (MeA) and bed nucleus of the stria terminalis (BNST) contribute to the most pronounced sex differences in AVP innervation in brain (De Vries and Boyle, 1998).”

“Although central AVP has been repeatedly implicated in the generation of anxiety states (Ebner et al., 2002; Bielsky et al., 2004; Raggenbass, 2008), we did not observe changes in anxiety-like behavior of lesioned males or females in a non-social anxiogenic environment (Hogg, 1996).”

“For example, the density of AVP fibers in BNST and MeA projection areas and c-Fos activation in AVP cells in the BNST correlate negatively with aggression in male mice and rats (Compaan et al., 1993; Everts et al., 1997; Ebner et al., 2000; Beiderbeck et al., 2007; Veenema et al., 2010) but positively with prosocial b…”

“, 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“In mice, a genetic knock-out of the vasopressin V1a receptor gene reduces anxiety-related behaviors in males, but not in females (Bielsky et al., 2004, 2005).”

“…(in birds) that report no effect of central infusions of AVT or V1a antagonists on femaledirected singing (Goodson and Adkins-Regan, 1999; Goodson and Evans, 2004) or got mixed results from antisense knock-down of BNST-AVP, with one study finding no effect (Kelly et al., 2011) and…”

“UM Following testing, the substrate sheet was allowed to dry for 1 h and then sprayed with ninhydrin fixative (LCNIN-16; Tritech Forensics Inc.) to visualize urine marks (Meyer, 1957; Lehmann et al., 2013).”

“As opposite-sex sexual experience and attaining competitive status (“social dominance”) promote male and female communicative behaviors (Lumley et al., 1999; Roullet et al., 2011), mice received social experience over five consecutive days (sexual encounters on days 1 and 4, aggressive encounters…”

“In addition, injecting specific V1a receptor agonists or boosting V1a receptor expression in target areas of AVP cells in the BNST and MeA promotes affiliation in voles (Wang et al., 1994; Liu et al., 2001; Pitkow et al., 2001; Lim and Young, 2004; Lim et al., 2004) and social recognition and active social behaviors i…”

“USV, UM, and social investigation were recorded in an acrylic three-chamber apparatus (Crawley, 2007; Arakawa et al., 2008a; Moy et al., 2009; Harvard Apparatus; dimensions: 20.3 42 22 cm).”

“, 2010) but positively with prosocial behavior (Goodson et al., 2009; Ho et al., 2010).”

“Although UM strongly depends on chemosensory processing (Maruniak et al., 1986; Labov and Wysocki, 1989), the increase in UM by lesioned males was not driven primarily by female urine cues because marking to female urine was not increased, suggesting that other cues, chemosensory as well as…”

“16-mm outer diameter; Dow Corning Corporation) were filled with crystalline T (Sigma) and inserted subcutaneously between the scapulae after gonadectomy; this procedure leads to physiologic levels of T (Barkley and Goldman, 1977; Matochik et al., 1994).”

“16-mm outer diameter; Dow Corning Corporation) were filled with crystalline T (Sigma) and inserted subcutaneously between the scapulae after gonadectomy; this procedure leads to physiologic levels of T (Barkley and Goldman, 1977; Matochik et al., 1994).”

“, 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“Although AVP has been implicated in female rat sexual behavior, the pattern of prior results is opposite to our findings: in rats, AVP reduced, whereas V1a receptor antagonists increased, female sexual behavior in rats (Södersten et al., 1983; Pedersen and Boccia, 2006).”

“However, the anatomy of AVP projections suggests that AVP control of social behavior is complex and, in most cases, the anatomic substrate of AVP’s control of social behavior is unclear (Kelly and Goodson, 2013a; Ludwig and Stern, 2015; Dumais and Veenema, 2016).”

“, 2010) but positively with prosocial behavior (Goodson et al., 2009; Ho et al., 2010).”

“, 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“Most other studies supporting a role for AVP/AVT in vocalizations have not identified the relevant neuroanatomical source of the peptide or its locus of action (Wilczynski et al., 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al.”

“2019 eNeuro.org Roberts et al., 2001), UM should be strongly regulated so that signaling occurs in only specific social contexts.”

“UM Following testing, the substrate sheet was allowed to dry for 1 h and then sprayed with ninhydrin fixative (LCNIN-16; Tritech Forensics Inc.) to visualize urine marks (Meyer, 1957; Lehmann et al., 2013).”

“Lesioning AVP cells also removed all other neuroactive substances co-expressed by these cells, for example, galanin (Miller et al., 1993), which also has been implicated in control of social behavior (Wu et al.”

“, 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“However, since large lesions of BNST have no effect on anxiety-like behavior in the EPM task either (Treit et al., 1998), it may simply mean that, as with aggressive behavior, AVP cells in the BNST are not critically involved in anxiety-like behaviors, and that AVP derived from other sources, such…”

“To induce sexual receptivity, stimulus females were injected subcutaneously with 0.1 ml of progesterone (500 g dissolved in sesame oil, Sigma) 4 h preceding sexual experience, urine collection, and behavioral testing (Veyrac et al., 2011).”

“This system activates an apoptotic signaling cascade, cleaving multiple structural and regulatory proteins critical for cell survival and maintenance (Yang et al., 2013) and thereby inducing far less inflammation than other lesion approaches (Morgan et al., 2014).”

“…which is consistent with the observation that knocking down AVT mRNA in the BNST of territorial birds reduces social contact with other males (Kelly and Goodson, 2013b), and that overexpression of V1a receptors (Landgraf et al., 2003) or AVP injections (Koolhaas et al., 1991) in the LS, a…”

“…source of the peptide or its locus of action (Wilczynski et al., 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“USV, UM, and social investigation were recorded in an acrylic three-chamber apparatus (Crawley, 2007; Arakawa et al., 2008a; Moy et al., 2009; Harvard Apparatus; dimensions: 20.3 42 22 cm).”

“For all subjects, 500 nl of AAV-flex-taCasp3-TEVp was delivered bilaterally to the BNST (coordinates: AP –0.01 mm; ML 0.75 mm; DV 4.8 mm; Paxinos and Franklin, 2012) at a rate of 100 nl/min using a 5- l Hamilton syringe with a 30-gauge beveled needle mounted on a stereotaxic injector.”

“Urine marks that were larger than 6 cm(2) and directed toward corners were counted as eliminative “pools” and were counted separately (Bishop and Chevins, 1987).”

“…which is consistent with the observation that knocking down AVT mRNA in the BNST of territorial birds reduces social contact with other males (Kelly and Goodson, 2013b), and that overexpression of V1a receptors (Landgraf et al., 2003) or AVP injections (Koolhaas et al., 1991) in the LS, a…”

“Although UM strongly depends on chemosensory processing (Maruniak et al., 1986; Labov and Wysocki, 1989), the increase in UM by lesioned males was not driven primarily by female urine cues because marking to female urine was not increased, suggesting that other cues, chemosensory as well as…”

“, 2017) or tended to focus on stress-related or aggressive vocalizations across development (Winslow and Insel, 1993; Bleickardt et al., 2009; Iijima et al., 2014; Lukas and Wöhr, 2015; Paul et al., 2016; Freeman et al., 2018).”

“Gonadectomy, group housing, and social defeat of our subordinates reduce offensive aggression in mice, while GDX T male urine provides subjects with a male urinary cue that elicits offensive aggression (Beeman, 1947; Connor and Winston, 1972; Van Loo et al., 2001).”

“…a sex-specific manner, but it is not clear if they project to the VMH. Ablation of BNST AVP neurons in males increases urine marking in the presence of a female, but does not alter ultra-sonic vocalizations, copulatory behavior, or investigation of the females or their odor (Rigney et al., 2019).”

“(A) BNST AVP cell ablations in males reduce male-male social investigation [redrawn from (106)].”

“Among those are PACAP, oxytocin (OT), substance P (SP), neurokinin B (Tac2), vasopressin (Avp), cholecystokinin (CCK), and nociceptin (NOC) (Malsbury and McKay, 1987; Walter et al., 1991; Poulin et al., 2009; Kudo et al., 2014; Ahrens et al., 2018; Giardino et al., 2018; Zelikowsky et al., 2018; Kovner et al., 2019; R…”

“As we previously reported, BNST AVP cell deletion generates deficits in social communication in males only (Rigney et al., 2019).”

“As reported in the literature, machine learning could be applied in industry (Bertolini et al. 2021; Gourisaria et al. 2021), medical (Garg and Mago 2021), social investigation (Rigney et al. 2019), transportation (Abramowski 2008; Geng et al. 2020), and education.”

“Consequently, pharmacological and genetic manipulations of this pathway often impact social and affective behaviours in a sex-specific manner (Albers, 2015; Bredewold & Veenema, 2018; Rigney et al., 2019; Whylings et al., 2020).”

“This does not appear to be the case as we find that the ratio of cells expressing opsins in males and females is similar to the normal sex - biased ratio of AVP - expressing cells in BNST of these mice in an earlier study ( ~ 50% more AVP cells in males than in females) (27).”

“The BST (an SBN node) receives direct axonal projections from the olfactory bulbs in mammals and is well known for playing a role in social behavior and communication (Scalia and Winans, 1975; Newman, 1999; Been and Petrulis, 2010; Lebow and Chen, 2019; Rigney et al., 2019).”

“Neuropeptides CRF, OT, and AVP that contribute to anxiety-like and social behaviors, and are altered by ELS, are known to differ between sexes (Duque-Wilckens et al., 2018; Miller, Vician, Clifton, & Dorsa, 1989; Rigney et al., 2019; Rood et al., 2013; Uchida et al., 2019).”

“BST VP is involved in nonreproductive prosocial behaviors in prairie voles (Kelly et al., 2018b), sociosexual behavior in mice (Ho et al., 2010), social investigation in mice (Rigney et al., 2019), and maternal behavior in California mice (Bester-Meredith and Marler, 2012).”

“The serotonin (Grieb et al., 2021), oxytocin (Olazabal and Young, 2006), vasopressin (Rigney et al., 2019), endocannabinoid (Papilloud et al., 2018) and the opioid system (Achterberg et al., 2019) are also essential neuropeptide and neurotransmitter systems in the nucleus accumbens that may display…”

“Beyondthisperipheralparsingofmale/female cues, the processing of sex-specific information also occurs in higher order brain regions, such as the MeA and BNST (Bergan et al., 2014; Li et al., 2017; Rigney et al., 2019).”

“Rigney et al. (2019) found that BNST AVP cells influence female sexual behaviour and BNST AVP knockdown reduced male sex behaviour, and AVP is involved in the establishment of social bonds and could therefore mediate attachment between same sex sexual preferences (Keverne & Curley, 2004).”

“This role must come from a sexually dimorphic distribution and/or innervation of downstream targets 30,62 both of which we observed with Kiss1 BNST neurons.”

“These steroid-dependent AVP populations play crucial roles in sex-specific regulation of social and emotional behaviors, including social investigation, aggression, and anxiety (Kelley and Goodson, 2013; Rigney et al., 2019; Rigney et al., 2024; Veenema et al., 2013; Whylings et al., 2021).”

“Moreover, sex differences in OT/VP signaling further shape the function of these circuits (Lukas and Neumann, 2014; Dumais and Veenema, 2016; Rigney et al., 2019).”