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. 2022 Oct 21;16:994504. doi: 10.3389/fnbeh.2022.994504

TABLE 6.

Summary by brain areas of the regions most commonly involved in the expression the different social emotions across species.

Brain area Neural changes in socioemotional contexts (↗ increase/↘ decrease) Other associations–including non-imaging studies
ACC Romantic Love: in humans (Bartels and Zeki, 2000)
Social and motivational reward: in macaques (Chang et al., 2013)
Jealousy: in in male titi monkeys (Maninger et al., 2017b) (left ACC) and women (Steis et al., 2021)
Grief: in women (Gündel et al., 2003)
and Social Pain: in humans (Eisenberger et al., 2003)
Distress vocalizations
(Smith, 1945; MacLean and Newman, 1988; Eisenberger, 2012)
Empathy-Consolation
(Burkett et al., 2016)
Amygdala Romantic Love: in humans (Bartels and Zeki, 2000; Acevedo et al., 2012) at early stages but also in the longer term
Long-term Bonding: in male titi monkeys (Bales et al., 2007)
Jealousy: in male macaques, dogs and men; in male titi monkeys (Rilling et al., 2004; Takahashi et al., 2006; Maninger et al., 2017b; Cook et al., 2018)
Loneliness: in humans (Lam et al., 2021)
Fear
(Nesse, 1990)
Untrustworthiness of others
(Winston et al., 2002)
BNST Mother infant attachment
(Insel and Young, 2001)
Globus pallidus Jealousy: ↗ in women (Steis et al., 2021)
Hippocampus Romantic love and Feeling of friendship: in humans (Bartels and Zeki, 2004; Güroğlu et al., 2008; Acevedo et al., 2012)
Loneliness: in humans (Lam et al., 2021)
Insula Romanic and maternal love: in humans (Bartels and Zeki, 2000, 2004) (medial insula)
Feeling of Friendship: in humans (Acevedo et al., 2012)
Jealousy: in male macaques, men and women (Rilling et al., 2004; Takahashi et al., 2006; Steis et al., 2021) (bilateral)
Romantic rejection and acceptance: in humans (Hsu et al., 2020; van der Watt et al., 2021)
Grief: in women (Gündel et al., 2003; O’Connor et al., 2008)
and Social Pain: in humans (Eisenberger, 2012; Rotge et al., 2015) (anterior insula) and Loneliness: in humans (Lam et al., 2021)
Distress vocalizations
(Eisenberger, 2012)
LS Short-term Pair-bonding: lateral septum (D1R binding) (Hostetler et al., 2017) and Long-term Pair-bonding: in male titi monkeys (Bales et al., 2007)
Jealousy: in male titi monkeys (right LS) (Maninger et al., 2017b)
Separation: in male titi monkeys (Hinde et al., 2016)
MPOA Mother infant attachment
(Insel and Young, 2001)
NAcc Romantic Love*: in humans (Acevedo et al., 2012), but
in human in medial accumbens (Xu et al., 2011, 2012) and in titi monkeys in long-term bonds (Bales et al., 2007)
*Conflicting directions
Feeling of Friendship: in humans (Acevedo et al., 2012)
Grief: in women (O’Connor et al., 2008)
Separation
In voles: ↘ of striatal oxytocin signaling in the nucleus accumbens shell (Bosch et al., 2016)
OFC Romantic love*: in human (Acevedo et al., 2012) and in the mid-OFC (Xu et al., 2011)
and in human in medial OFC and negative correlation with happiness (Xu et al., 2011, 2012)
*Conflicting directions
Maternal Love: in human (Bartels and Zeki, 2004; Noriuchi et al., 2008)
Feeling of Friendship: in human (Acevedo et al., 2012) and macaques (Azzi et al., 2012; Watson and Platt, 2012; Brent et al., 2014a)
Untrustworthiness of others
(Winston et al., 2002)
PAG Maternal Love: in humans (Bartels and Zeki, 2004; Noriuchi et al., 2008)
Romantic Love and Feeling of Friendship: in humans (Acevedo et al., 2012)
Maternal behaviors
(Miranda-Paiva et al., 2003)
Distress vocalizations
(Panksepp, 1998; Eisenberger, 2012)
PCC Romantic Love: in humans (Bartels and Zeki, 2000)
Jealousy: in ↗ male titi monkeys (Maninger et al., 2017b) and women (Steis et al., 2021)
Grief: in women (Gündel et al., 2003)
Happiness, Fear, sadness, and other unpleasant stimuli
(Maddock, 1999)
Partner choice
(Maninger et al., 2017a; Yokoyama et al., 2017)
PFC Feeling of Friendship: in humans (Güroğlu et al., 2008)–ventro-medial PFC
Jealousy: ↗ in women (Steis et al., 2021)–medial PFC
Separation: ↗ in young macaques in the right dorsolateral PFC and in the left dorsolateral PFC (Rilling et al., 2001)
and Social Pain: in humans (Eisenberger et al., 2003; Rotge et al., 2015) (rostro ventral PFC)
Romantic rejection and acceptance: in humans (Hsu et al., 2020; van der Watt et al., 2021) and Loneliness: in humans (Lam et al., 2021)
PVN Separation (short-term): ↗ in male titi monkeys (Hinde et al., 2016)
PCL Love: ↗ in humans (Acevedo et al., 2012)
STS Jealousy: in male macaques (right) and women (posterior) (Rilling et al., 2004; Takahashi et al., 2006) Untrustworthiness of others
(Winston et al., 2002)
Gaze following in macaques
(Roy et al., 2014)
VP Attachment
↘ in titi monkey pair bonding (Bales et al., 2007)
VTA Romantic Love: ↗ in humans (Bartels and Zeki, 2004; Fisher et al., 2005; Acevedo et al., 2012) Onset of maternal behavior
(Pedersen et al., 1994; Numan and Young, 2016)
Mother infant attachment
(Insel and Young, 2001)
SN Love: ↗ in humans (Bartels and Zeki., 2004; Acevedo et al., 2012)
Jealousy: ↗ in women (Steis et al., 2021)

*Emphasize results going in conflicting directions.