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

TABLE 2.

Summary of the brain areas involved in the expression of “love” across species.

Species (Sex) How to elicit it (Imaging method) Neural changes (↗increase/↘ decrease) References
Love and attachment maternal and romantic
Titi monkey–Plecturocebus cupreus
(Males)
Males involved in long-term pair bond as compared to newly paired males or lone males
(PET scan)
Pair-bonding
↘ nucleus accumbens, ventral pallidum, medial preoptic area, medial amygdala, and the supraoptic nucleus of the hypothalamus, lateral septum
Bales et al., 2007
Before and 48 h after pair-bonding
(PET scan)
↘ right nucleus accumbens and ventral pallidum
Titi monkey–Plecturocebus cupreus
(Males)
Newly paired males (1 week) and long-term paired (4 month) as compared to non-paired males
(PET scan)
nucleus accumbens, ventral pallidum, caudate, putamen
(“motivational areas”)
Maninger et al., 2017a
Titi monkey–Plecturocebus cupreus
(Males)
Newly paired males (4–8 weeks after pair formation) compared to unpaired males
(PET scan)
↗ lateral septum (D1R binding) Hostetler et al., 2017
Sprague Dawley Rats–Rattus norvegicus
(Females)
Pup stimulation via suckling
(fMRI)
Mother-infant attachment
nucleus accumbens, caudate nucleus, putamen, prefrontal cortex
(“reward pathway”)
Ferris et al., 2005
Humans–Homo sapiens
(Men and Women)
Deeply involved adults scan presented to the picture of the loved one compared to the picture of a friend
(fMRI)
Romantic love
↗ medial insula, anterior cingulate cortex, caudate nucleus and the putamen
↘ amygdala, posterior cingulate gyrus,
right prefrontal, parietal and middle temporal cortices
Bartels and Zeki, 2000
Humans–Homo sapiens Mothers presented to pictures of their own compared to a picture of acquainted children, of their best friend and of acquainted adults
(fMRI)
Specific to Maternal love
↗ periaqueductal central gray, lateral orbitofrontal cortex
Common to Romantic and maternal love
↗ medial insula, anterior cingulate cortex; caudate nucleus, putamen, the globus pallidus, ventral tegmental area, substantia nigra
↘ amygdala, posterior cingulate cortex, middle prefrontal, inferior parietal, middle temporal cortices, temporal poles, parietotemporal junction, mesial prefrontal cortex
Bartels and Zeki, 2004
Humans–Homo sapiens
(Men and Women)
Participants intensely in love watching picture of their beloved partner compared to acquaintance.
(fMRI)
↗ ventral tegmental area, caudate nucleus
But also for longer-term relationships:
mid-insular cortex, right anterior and posterior cingulate cortex
Fisher et al., 2005
Humans–Homo sapiens Mother watching videoclips of their child compared to other children.
(fMRI)
Maternal love:
↗ periaqueductal central gray orbitofrontal cortex, anterior insula, and dorsal and ventrolateral parts of putamen.
Noriuchi et al., 2008
Humans–Homo sapiens
(Men and Women)
Comparison of long-term couples to long-term friendship or familiar neutral
(fMRI)
nucleus accumbens, ventral tegmental area, caudate nucleus, cerebellum, putamen, mid-insula, posterior hippocampus, left, globus pallidus, substantia nigra, raphe nucleus, thalamus, anterior cingulate cortex, posterior cingulate cortex, periaqueductal gray, hypothalamus
+ increase of nucleus accumbens activation over the years
Acevedo et al., 2012
Humans–Homo sapiens
(Men and Women)
Newly married couple resented to a picture of partner or neutral acquaintance (highly familiar) at two time points: around the weeding date and approximately 11 months after the wedding ↗ substantia nigra, left paracentral lobule (PCL); ventral tegmental area, insular cortex, periaqueductal gray, posterior hippocampus, occipital cortex, Septum/fornix region
↘ anterior temporal gyrus, inferior frontal gyrus, Superior temporal gyrus/angular gyrus
Acevedo et al., 2020
Humans Homo sapiens
(Men)
Pair-bonded male volunteers presented to pictures of their bond or another women
(fMRI)
nucleus accumbens Scheele et al., 2013
Humans–Homo sapiens
(Men and Women)
Participants involved in a romantic relationship and very intensely in love, presented to pictures of their partner or a familiar acquaintance
(fMRI)
ventral tegmental area, caudate nucleus (tail), cerebellum, middle orbitofrontal cortex
↘ right amygdala, right accumbens, right medial orbitofrontal
Xu et al., 2011

Names in bold are brain areas regularly found in neural changes related to emotion.