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. 2023 Aug 30;17:1225758. doi: 10.3389/fncel.2023.1225758

TABLE 1.

Overview of the input-output connectivity patterns and the chemoarchitecture of the BNSTALG.

Rat
(Rattus norvegicus)
Mouse
(Mus musculus)
Macaque
(Macaca mulatta)
Human
(Homo sapiens)
BNST subnucleus alBNST (Allen et al., 1983; Swanson et al., 1983; Chronwall et al., 1985; Fallon and Leslie, 1986; Gray and Magnuson, 1987, 1992; Ju et al., 1989; Poulin et al., 2009) Inline graphic
ovBNST (Allen et al., 1983; Swanson et al., 1983; Chronwall et al., 1985; Fallon and Leslie, 1986; Gray and Magnuson, 1987, 1992; Ju et al., 1989; Poulin et al., 2009) Inline graphic
juBNST (Allen et al., 1983; Swanson et al., 1983; Chronwall et al., 1985; Fallon and Leslie, 1986; Gray and Magnuson, 1987, 1992; Ju et al., 1989; Poulin et al., 2009) Inline graphic
amBNST (Allen et al., 1983; Swanson et al., 1983; Chronwall et al., 1985; Fallon and Leslie, 1986; Gray and Magnuson, 1987, 1992; Ju et al., 1989; Poulin et al., 2009) Inline graphic
alBNST (Nguyen et al., 2016; Ortiz-Juza et al., 2021) Inline graphic
ovBNST (Nguyen et al., 2016; Smith et al., 2016; Ortiz-Juza et al., 2021) Inline graphic
juBNST (Nguyen et al., 2016) Inline graphic
amBNST (Pleil et al., 2012; Nguyen et al., 2016; Ortiz-Juza et al., 2021) Inline graphic
BNSTlp (deCampo and Fudge, 2013; Fudge et al., 2022) Inline graphic
BNSTld (deCampo and Fudge, 2013; Fudge et al., 2022) Inline graphic
BNSTlj (deCampo and Fudge, 2013; Fudge et al., 2022) Inline graphic
BNSTma
BNSTc (Walter et al., 1991) Inline graphic
BNSTl (Adrian et al., 1983; Walter et al., 1991) Inline graphic
BNSTlj BNSTm (Walter et al., 1991) Inline graphic
Input Cerebral Cortex:
– mPFC (Radley et al., 2009)
– Insula (Dong et al., 2001a)
– Infralimbic Cortex (Vertes, 2004)
– Prelimbic Cortex (Vertes, 2004)
Amygdala:
– CeA (Myers et al., 2014)
– MeA (Dong et al., 2001b)
– BLA (Krettek and Price, 1978) Hippocampus:
– Subiculum (Dong et al., 2001a)
Hypothalamus:
– PVN (Berk and Finkelstein, 1982)
– MPN (Myers et al., 2014)
– VMH (Myers et al., 2014)
– mPOA (Myers et al., 2014)
Thalamus:
– PVT (Dong et al., 2001a)
Midbrain/Hindbrain:
– PBN (Dong et al., 2001a)
– LC (Dong et al., 2001a)
– VTA (Dong et al., 2001a; Hasue and Shammah-Lagnado, 2002)
– NST (Dong et al., 2001a)
– DR (Azmitia and Segal, 1978; Petit et al., 1995)
– LS (Myers et al., 2014)
Basal ganglia:
– NAc (Myers et al., 2014)
– SNc (Hasue and Shammah-Lagnado, 2002)
Cerebral Cortex:
– Insula (Sun et al., 2023)
– Prelimbic cortex (Sun et al., 2023)
– Anterior cingulate cortex (Sun et al., 2023)
– Piriform cortex (Sun et al., 2023)
– Entorhinal cortex (Sun et al., 2023)
– Perirhinal cortex (Sun et al., 2023)
Amygdala:
– CeA (Sun et al., 2023)
– MeA (Sun et al., 2023)
– BLA (Vantrease et al., 2022; Sun et al., 2023)
Hippocampus:
– Subiculum (Sun et al., 2023)
Hypothalamus:
– POA (Sun et al., 2023)
– TMN (Lin et al., 2023)
Thalamus:
– Anteromedial (Sun et al., 2023)
Midbrain/Hindbrain:
– PAG (Sun et al., 2023)
– LS (Sun et al., 2023)
– DR (Sun et al., 2023)
Basal ganglia:
– pDMS (Lu et al., 2021)
Cerebral Cortex:
– OFC (Fox et al., 2010)
– ?
Amygdala: (Novotny, 1977)
– CeA (deCampo and Fudge, 2013; Oler et al., 2017)
– BLA (deCampo and Fudge, 2013)
– SLEA (Oler et al., 2017)
Hippocampus:
– Subiculum (Berry et al., 2022)
Hypothalamus:
– ?
Thalamus:
– ?
Midbrain/Hindbrain:
– VTA (Fudge and Haber, 2001)
– ?
Basal Ganglia:
– NAc (Kalin et al., 2005)
– ?
Cerebral Cortex:
– mPFC (Krüger et al., 2015; Torrisi et al., 2015)
– dlPFC (Krüger et al., 2015; Torrisi et al., 2015)
– vmPFC (Motzkin et al., 2015)
– OFC (Krüger et al., 2015)
– Precuneus (Torrisi et al., 2015)
– Insula (Flook et al., 2020)
Amygdala (Avery et al., 2014):
CeA (Krüger et al., 2015; Gungor and Paré, 2016; Shackman and Fox, 2016; Brinkmann et al., 2018; Hofmann and Straube, 2019, 2021)- BLA (Hofmann and Straube, 2019, 2021)
– SLEA (Hofmann and Straube, 2019)
Hippocampus (Avery et al., 2014):
– Subiculum (Berry et al., 2022)
Hypothalamus
– PVN (Krüger et al., 2015)
Thalamus (Avery et al., 2014):
– IL/CM midline (Torrisi et al., 2015)
Midbrain/hindbrain:
– DR (Torrisi et al., 2015)
– PAG (Torrisi et al., 2015)
– Habenula (Torrisi et al., 2015)
Basal ganglia:
– Caudate nucleus (Torrisi et al., 2015)
– Putamen (Avery et al., 2014)
– NAc (Torrisi et al., 2015)
*Note:
Human input-output is mostly based on magnetic resonance imaging (MRI) data. In particular, functional/resting state connectivity patterns.
Output Cerebral Cortex:
– Prelimbic cortex (Dabrowska et al., 2016) Inline graphic
Prelimbic cortex (Dabrowska et al., 2016) Inline graphic
– Habenula (Dabrowska et al., 2016)
Amygdala:
– CeA (Dong et al., 2001a; Dong and Swanson, 2004; Dabrowska et al., 2016) Inline graphic
– BLA (Dong et al., 2001a; Dabrowska et al., 2016; Leitermann et al., 2016) Inline graphic
– MeA (Dabrowska et al., 2016) Inline graphic
Hypothalamus:
– PVN (Dong et al., 2001a; Dabrowska et al., 2016) Inline graphic
– LH (Dong et al., 2001a) Inline graphic
Thalamus
– Subthalamic nucleus (Dong et al., 2001a)
– Midline nuclei (Dong et al., 2001a)
– PVT (Dabrowska et al., 2016; Brinkmann et al., 2018) Inline graphic
Midbrain/Hindbrain:
– PAG (Gray and Magnuson, 1987; Dong and Swanson, 2004, 2006; Dabrowska et al., 2016) Inline graphic
– VTA (Gray and Magnuson, 1987; Dong et al., 2001a; Dabrowska et al., 2016) Inline graphic
– PBN (Dong et al., 2001a; Brinkmann et al., 2018) Inline graphic
– Zona incerta (Dong et al., 2001a)
– DR (Dong et al., 2001a; Dabrowska et al., 2016) Inline graphic
– Reticular area (Dong et al., 2001a)
– Lateral Tegmental nucleus (Dong et al., 2001a)
– Red Nucleus (Dabrowska et al., 2016) Inline graphic
– Nucleus of the solitary tract
– Substantia innominata (Dong et al., 2001a; Brinkmann et al., 2018) Inline graphic
Basal Ganglia:
– NAc (Dabrowska et al., 2016) Inline graphic
– Globus Pallidus (Dong et al., 2001a)
– Septal Nucleus (Ju et al., 1989)
– SNc (Dong et al., 2001a; Dabrowska et al., 2016) Inline graphic
Cerebral cortex:
– Prelimbic cortex (Dabrowska et al., 2016)
Amygdala:
– MeA (Dabrowska et al., 2016; Bruzsik et al., 2021) Inline graphic
– CeA (Dabrowska et al., 2016; Bruzsik et al., 2021) Inline graphic
– BLA (Dabrowska et al., 2016; Bruzsik et al., 2021) Inline graphic
Thalamus:
– Parasubthalamic nucleus (Lin et al., 2023) Inline graphic
Hypothalamus:
– PVN (Barbier et al., 2021) Inline graphic
– DMH (Barbier et al., 2021)
– ARH (Barbier et al., 2021)
– LH (Barbier et al., 2021; Bruzsik et al., 2021) Inline graphic
Midbrain/Hindbrain:
– VTA (Kudo et al., 2012; Fellinger et al., 2021; Miura et al., 2022) Inline graphic
– PAG (Dabrowska et al., 2016; Bruzsik et al., 2021) Inline graphic
– PBN (Dabrowska et al., 2016) PBN (Dabrowska et al., 2016)
– DR (Dabrowska et al., 2016; Bruzsik et al., 2021) Inline graphic
– Red Nucleus (Dabrowska et al., 2016) Inline graphic
– Nucleus of the solitary tract (Dabrowska et al., 2016)
– Substantia innominata (Bruzsik et al., 2021) Inline graphic
– TMN (Bruzsik et al., 2021) Inline graphic
– Locus Coeruleus (Bruzsik et al., 2021) Inline graphic
Basal Ganglia:
– Dorsal striatum (Smith et al., 2016; Lu et al., 2021) Inline graphic
– NAc (Dabrowska et al., 2016; Xiao et al., 2021) Inline graphic
– SNc (Smith et al., 2016; Bruzsik et al., 2021) Inline graphic
Cerebral Cortex:
– OFC (Fox et al., 2018)
– ?
Amygdala: (Novotny, 1977)
– SLEA (Oler et al., 2017)
– CeA
Hypothalamus:
– ?
Thalamus:
– ?
Midbrain/hindbrain
– VTA (Fudge et al., 2017)
Basal Ganglia:
– Ventromedial striatum (Fudge et al., 2017)
– NAc (Kalin et al., 2005)
Cerebral Cortex:
– mPFC, dlPFC (Torrisi et al., 2015)
– Precuneus (Torrisi et al., 2015)
Amygdala: (Avery et al., 2014; Klumpers et al., 2017)
– CeA (Gungor and Paré, 2016; Shackman and Fox, 2016; Brinkmann et al., 2018; Hofmann and Straube, 2019)
– SLEA (Hofmann and Straube, 2019)
Hippocampus (Avery et al., 2014)
Hypothalamus:
– PVN (Krüger et al., 2015; Hofmann and Straube, 2019)
Thalamus (Avery et al., 2014):
– CM midline (Torrisi et al., 2015)
Midbrain/Hindbrain:
– DR (Torrisi et al., 2015)
– PAG (Torrisi et al., 2015)
– Habenula (Torrisi et al., 2015)
Basal Ganglia:
– Caudate nucleus (Torrisi et al., 2015)
– Putamen (Avery et al., 2014)
– NAc (Torrisi et al., 2015)
Behavioural function and dysfunction – Stress induced nociception (Tran et al., 2012)


– Autonomic stress response (modulating ACTH release) (Dong et al., 2001a)
– Contextual, cued, and unpredictable threat processing (Walker et al., 2009; Urien and Bauer, 2022)
– (Conditioned) anxiety-like behaviour (Sahuque et al., 2006; Sink et al., 2013; Sherman et al., 2022)
– Modulation of behavioural despair (Pezük et al., 2006)
– Modulating stress-induced changes in sensorimotor gating (Rajbhandari and Bakshi, 2020)
– Modulating locomotor, orofacial and ingestive behaviour (Dong et al., 2001a)
– PTSD (Rodríguez-Sierra et al., 2016)
– Chronic Stress (Normandeau et al., 2018)
– Alcohol/drugs-motivated behaviour (Epping-Jordan et al., 1998; Eiler et al., 2003)
– Anxiety like behaviour during withdrawal (Huang et al., 2010)
(Acute) pain related behaviors (Myers et al., 2014; Fang et al., 2023)

– Arousal (Ju et al., 1989)
– Conditioned anxiety/ contextual fear/ sustained anxiety (Sullivan et al., 2004; Vantrease et al., 2022)
– Sustained anxiety/threat anticipation/ threat processing (Fellinger et al., 2021)
– Pavlovian fear and reward behaviour/learning (Bruzsik et al., 2021; Kaouane et al., 2021)
– Regulation of food intake (Wang et al., 2019)
– Alcohol- and drug-motivated behaviour (Silberman et al., 2013)
– Anxiety related responses (freezing behaviour) (Kalin et al., 2005)

– Threat anticipation/evaluation of threatening stimuli (Hoffman et al., 2007)
– Behavioural inhibition (Fox et al., 2010)
– Social/affiliative behaviour: aggression, mating behaviour, maternal care, social interaction (Jacobs et al., 2023)
– Sustained anxiety/threat anticipa
tion/threat processing (Davis et al., 2010; Alvarez et al., 2011; Grupe et al., 2012; Somerville et al., 2013; Herrmann et al., 2016; Brinkmann et al., 2018; Camilo et al., 2022)
– Phasic threat processing (Gungor and Paré, 2016; Shackman and Fox, 2016; Klumpers et al., 2017; Brinkmann et al., 2018; Koller et al., 2019)
– Anxious temperament (Fox et al., 2018)
– PTSD (Ahearn et al., 2007; Brinkmann et al., 2017; Awasthi et al., 2020; Feola et al., 2023)
– Generalized Anxiety disorder (Avery et al., 2016; Buff et al., 2017; Brinkmann et al., 2018)
– Social Anxiety (Avery et al., 2016; Clauss, 2019; Clauss et al., 2019)
– Affective disorders (Fox et al., 2010)

alBNST, anterolateral BNST; amBNST, anteromedial BNST; BNSTc, central BNST; BNSTl, lateral BNST; BNSTld, laterodorsal BNST; BNSTlj, lateral juxtacapsular BNST; BNSTlp, lateral posterior BNST; BNSTma, medial anterior BNST; BNSTm, medial BNST; juBNST, juxtacapsular BNST; ovBNST, oval BNST; BLA, basolateral amygdala; CeA, central amygdala; CM, centromedian nucleus; dlPFC, dorsolateral prefrontal cortex; DR, dorsal raphe; LC, locus coeruleus; LS, lateral septum; MeA, medial amygdala; mPFC, medial prefrontal cortex; MPN, median preoptic nucleus; mPOA, median preoptic area; NAc, nucleus accumbens; NST, solitary nucleus; PBN, parabrachial nucleus; PVN, paraventricular nucleus; SLEA, sublenticular extended amygdala; SNc, substantia nigra pars compacta; VMH, ventromedial hypothalamus; VTA, ventral tegmental area; ACTH, adrenocorticotropic hormone; CRF, corticotropin releasing factor; DYN, dynorphin; ENK, enkephalin; GABA, gamma-aminobutyric acid; NPY, neuropeptide Y; NT, neurotensin; PKC, protein kinase C type delta; PTSD, post-traumatic stress disorder; SOM, somatostatin.

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