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. Author manuscript; available in PMC: 2017 Oct 24.
Published in final edited form as: Nat Hum Behav. 2017 Apr 24;1:0069. doi: 10.1038/s41562-017-0069

Table 2.

Summary of tract-tracing study results in non-human animals, demonstrating anatomical connections between cortical visceromotor and primary interoceptive sensory regions, as well as between cortical and non-cortical visceromotor regions.

Primary Interoceptive Cortex Visceromotor Regions Subcortical and Brainstem Visceromotor Structures
To dpIns/dmIns To vaIns To sgACC (BA 25) To pACC (BA 24, 32) To aMCC (BA 24) To Amygdala To other subcortical and brainstem regionsa
From dpIns/dmIns - Case A, Fig 1155 Not evidentb Case 1, Fig 5157 Case B, Fig 3158 Case 2, Fig 3154
Case BB-B, Fig 160
Hypothalamus (rat)159
PAG: not observed160
PBN (rat)161,162
V. Striatum 163
NTS (rat)162
From vaInsc Case C, Fig 4155
Case A, Fig 1158
- Case OM20, Fig 8164 Case 1, Fig 5157 Case 2, Fig 6157
Case A, Fig 1158
Case A, Fig 1158
Case 103, Fig 3165
Fig 2, Table 2166
Hypothalamus43
PAG160
PBN (rat)161
V. striatum167
NTS (rat)162
From sgACC (BA 25) Not evidentd Case M707168 - Case 1, Fig 5157
Fig 2A169
Case 3, Fig 7157
Fig 3A169
Case 103, Fig 3165
Fig 5156
Hypothalamus154,170,171
PAG160,171
PBN 171
Striatum 171
NTS (rat)172,173
From pACC (BA 24, 32) Not evidentd Case M776168 Fig 1169 - Case 3, Fig 7157
Fig 3A169
Case 103, Fig 3165
Fig 5156
Hypothalamus43,
PAG160
PBN (cat)174
V. striatum (cat)174
NTS (rat)173
From aMCC (BA 24) Case C, Fig 4155 Case A, Fig 1155 Case 3, Fig 4175 Case 1, Fig 5157
Fig 2A169
- Case 103, Fig 3165
Fig 5156
Hypothalamus43
PAG160
PBN: not present176
V. striatum177
NTS (rat)172
From Amygdala Case C, Fig 4155
Lateral basal nucleus; Case 5, Fig 6154
Case A, Fig 1155
Case 4, Fig 5154
Fig 6156 Fig 13169 Fig 6156 - Hypothalamus43,
PAG160
PBN178
V. striatum179
NTS178

Note. Connectivity evidence is in monkeys unless otherwise indicated (e.g., rats, cats). Some connections from dpIns/dmIns to the NTS are unclear due to ambiguity in how Saper (1982)162 reported subregions of the insula.

a

We did not assess for projections from subcortical and brainstem regions to cortical regions because we only wanted to determine if the cortical regions support visceromotor control.

b

Connection from dpIns/dmIns to sgACC not evident in several monkey studies that have the potential to show them (e.g., 158,169,180182).

c

The medial portion of the vaIns exhibits connectivity with subcortical and brainstem regions, but not the lateral portion of the vaIns43,183.

d

Connection from sgACC to dpIns/dmIns and from pACC to dpIns/dmIns not evident in several monkey studies that have the potential to show them (e.g., 155,168,180,181), although weak, direct connectivity is evident in a recent tractography study in humans (Ghaziri, et al., 2015184, Figure 5). Moreover, connections between sgACC, pACC, and dpIns have been observed in intrinsic functional connectivity analyses in humans (e.g., Fig. 6 of 185). The discrepancy between human findings and the tract tracing studies in monkeys failing to show connectivity might reflect an expansion of Brodmann area (BA) 24 anterior and ventral to the corpus callosum in humans relative to monkeys and/or the presence of connections between BAs 25/32 and the posterior insula in humans that do not exist in monkeys (Evrard, H. personal communication, December 27, 2015).

BA = Brodmann area; aMCC = anterior midcingulate cortex; dmIns = dorsal mid insula; dpIns = dorsal posterior insula; NTS = nucleus of the solitary tract; PAG = periaqueductal gray; PBN = parabrachial nucleus; pACC = pregenual anterior cingulate cortex; sgACC = subgenual anterior cingulate cortex; V. striatum = ventral striatum.