Salience network

The salience network (SN), also known anatomically as the midcingulo-insular network (M-CIN) or ventral attention network, is a large scale network of the human brain that is primarily composed of the anterior insula (AI) and dorsal anterior cingulate cortex (dACC). It is involved in detecting and filtering salient stimuli, as well as in recruiting relevant functional networks.[3][4] Together with its interconnected brain networks, the SN contributes to a variety of complex functions, including communication, social behavior, and self-awareness through the integration of sensory, emotional, and cognitive information.[5]

The salience network is theorized to mediate switching between the default mode network and central executive network.[1][2]

The network is detectable through independent component analysis of resting state fMRI images, as well as seed based functional connectivity analysis. The functional connectivity has been linked with structural connectivity through diffusion tensor imaging, which reveals white matter tracts between the AI and dACC.

Anatomy

The salience network is primarily anchored at the AI and dACC. The node in the AI corresponds with the dorsal-anterior division distinguished in meta-analyses of task-positive network related neuroimaging studies. The AI and dACC are linked via a white matter tract along the uncinate fasciculus. Other regions of the network may include the inferior parietal cortex, right temporoparietal junction, and lateral prefrontal cortex.[6]

The subcortical nodes have yet to be structurally linked to the AI and dACC, however both seed-based and resting-state studies have observed intrinsic connectivity of the cortical nodes, with subcortical nodes consisting of the sublenticular extended amygdala, the putamen, the ventral striatum, the dorsomedial thalamus, hypothalamus, and the substantia nigra/ventral tegmental area.[7] The salience network is also distinguished by distinct cellular components, including von Economo neurons in the AI/dACC.[7] Cortico-striatal-thalamic loop circuits contribute to the salience network.[4]

Function
Dorsal and ventral attention systems
Interaction between the ventral (salience) and dorsal attention networks enables dynamic control of attention in relation to top-down goals and bottom-up sensory stimulation.[8]

While the function of the salience network is not exactly known, it has been implicated in the detection and integration of emotional and sensory stimuli,[9] as well as in modulating the switch between the internally directed cognition of the default mode network and the externally directed cognition of the central executive network.[7] Evidence that the salience network mediates a switch between the DMN and CEN comes from Granger causality analysis and studies utilizing transcranial magnetic stimulation.[10] The timing of electrophysiological responses during the oddball task is consistent with interaction, as after the initial mismatch negativity response is transmitted "bottom-up" from sensory regions, a "top-down" signal localized to the AI and dACC occurs before a widespread evoked potential that corresponds to attentional shifting.[3] It has also been hypothesized that the AI receives multimodal sensory input and the ACC and the associated dorsomedial prefrontal cortex sends motor output.[3]

Clinical significance

Dysfunction in the salience network have been observed in various psychiatric disorders, including anxiety disorders, post-traumatic stress disorder, schizophrenia, frontotemporal dementia, and Alzheimer's disease. The AI node of the salience network has been observed to be hyperactive in anxiety disorders, which is thought to reflect predictions of aversive bodily states leading to worrisome thoughts and anxious behaviors. In schizophrenia, both structural and functional abnormalities have been observed, thought to reflect excessive salience being ascribed to internally generated stimuli.[11] In individuals with autism, the relative salience of social stimuli, such as face, eyes, and gaze, may be diminished, leading to poor social skills.[5]

Nomenclature

The cingulo-opercular network (CO) has generally been equated with the salience network, but it may represent a distinct but adjacent network[12] or a part of the SN.[13] The CO may involve more dorsal areas, while the SN involves more ventral and rostral areas of the anterior insula and medial frontal cortex containing von Economo neurons.[12] The CO is sometimes also referred to as the cingulo-insular network.[12]

The ventral attention network (VAN), also known as the ventral frontoparietal network (VFN) or ventral attention system (VAS), has also been equated with the SN. The VAN is commonly defined as a right-hemisphere-dominant network involving the temporoparietal junction and the ventral frontal cortex that responds to unexpected salient stimuli.[14][15] Some have defined it as a larger, bilateral network that is a combination of the SN and CO,[16] while others have described it as a part of the salience network involving the more dorsal anterior insular cortex.[17]

In 2019, Uddin et al. proposed that midcingulo-insular network (M-CIN) be used as a standard anatomical name for the network that includes the SN, CO, and VAN.[6]

See also

References
  1. Sridharan, D.; Levitin, D. J.; Menon, V. (22 August 2008). "A critical role for the right fronto-insular cortex in switching between central-executive and default-mode networks". Proceedings of the National Academy of Sciences. 105 (34): 12569–12574. Bibcode:2008PNAS..10512569S. doi:10.1073/pnas.0800005105. PMC 2527952. PMID 18723676.
  2. Nekovarova, Tereza; Fajnerova, Iveta; Horacek, Jiri; Spaniel, Filip (30 May 2014). "Bridging disparate symptoms of schizophrenia: a triple network dysfunction theory". Frontiers in Behavioral Neuroscience. 8: 171. doi:10.3389/fnbeh.2014.00171. PMC 4038855. PMID 24910597.
  3. Menon, V; Uddin, LQ (June 2010). "Saliency, switching, attention and control: a network model of insula function". Brain Structure & Function. 214 (5–6): 655–67. doi:10.1007/s00429-010-0262-0. PMC 2899886. PMID 20512370.
  4. Peters, SK; Dunlop, K; Downar, J (2016). "Cortico-Striatal-Thalamic Loop Circuits of the Salience Network: A Central Pathway in Psychiatric Disease and Treatment". Frontiers in Systems Neuroscience. 10: 104. doi:10.3389/fnsys.2016.00104. PMC 5187454. PMID 28082874.
  5. Menon V. (2015) Salience Network. In: Arthur W. Toga, editor. Brain Mapping: An Encyclopedic Reference, vol. 2, pp. 597-611. Academic Press: Elsevier. https://med.stanford.edu/content/dam/sm/scsnl/documents/Menon_Salience_Network_15.pdf
  6. Uddin, Lucina Q.; Yeo, B. T. Thomas; Spreng, R. Nathan (2019-11-01). "Towards a Universal Taxonomy of Macro-scale Functional Human Brain Networks". Brain Topography. 32 (6): 926–942. doi:10.1007/s10548-019-00744-6. ISSN 1573-6792. PMC 7325607. PMID 31707621.
  7. Menon, V; Toga, A (2015). Salience Network. Elsevier. pp. 597–611. ISBN 978-0-12-397316-0.
  8. Vossel, S; Geng, JJ; Fink, GR (April 2014). "Dorsal and ventral attention systems: distinct neural circuits but collaborative roles". The Neuroscientist. 20 (2): 150–9. doi:10.1177/1073858413494269. PMC 4107817. PMID 23835449.
  9. Downar, J.; Crawley, A. P.; Mikulis, D. J.; Davis, K. D. (2000). "A multimodal cortical network for the detection of changes in the sensory environment". Nature Neuroscience. 3 (3): 277–283. doi:10.1038/72991. PMID 10700261. S2CID 8807081.
  10. Uddin, Lucina Q. (19 November 2014). "Salience processing and insular cortical function and dysfunction". Nature Reviews Neuroscience. 16 (1): 55–61. doi:10.1038/nrn3857. PMID 25406711. S2CID 7786680.
  11. Menon, V (October 2011). "Large-scale brain networks and psychopathology: a unifying triple network model". Trends in Cognitive Sciences. 15 (10): 483–506. doi:10.1016/j.tics.2011.08.003. PMID 21908230. S2CID 26653572.
  12. Gratton, Caterina; Sun, Haoxin; Petersen, Steven E. (2018). "Control networks and hubs". Psychophysiology. 55 (3): e13032. doi:10.1111/psyp.13032. ISSN 1469-8986. PMC 5811327. PMID 29193146.
  13. Sestieri, Carlo; Corbetta, Maurizio; Spadone, Sara; Romani, Gian Luca; Shulman, Gordon L. (March 2014). "Domain-general signals in the cingulo-opercular network for visuospatial attention and episodic memory". Journal of Cognitive Neuroscience. 26 (3): 551–568. doi:10.1162/jocn_a_00504. ISSN 0898-929X. PMC 3947512. PMID 24144246.
  14. Fox, M.D.; Corbetta, M.; Snyder, A.Z.; Vincent, J.L.; Raichle, M.E. (2006). "Spontaneous neuronal activity distinguishes human dorsal and ventral attention systems". PNAS. 103 (26): 10046–10051. Bibcode:2006PNAS..10310046F. doi:10.1073/pnas.0604187103. PMC 1480402. PMID 16788060.
  15. Farrant, Kristafor; Uddin, Lucina Q. (2015-02-12). "Asymmetric development of dorsal and ventral attention networks in the human brain". Developmental Cognitive Neuroscience. 12: 165–174. doi:10.1016/j.dcn.2015.02.001. ISSN 1878-9293. PMC 4396619. PMID 25797238.
  16. Webb, Taylor W.; Igelström, Kajsa M.; Schurger, Aaron; Graziano, Michael S. A. (2016-11-29). "Cortical networks involved in visual awareness independent of visual attention - Supporting Information" (PDF). Proceedings of the National Academy of Sciences. 113 (48): 13923–13928. doi:10.1073/pnas.1611505113. ISSN 0027-8424. PMC 5137756. PMID 27849616.
  17. Touroutoglou, Alexandra; Bliss-Moreau, Eliza; Zhang, Jiahe; Mantini, Dante; Vanduffel, Wim; Dickerson, Bradford C.; Barrett, Lisa Feldman (2016-05-15). "A Ventral Salience Network in the Macaque Brain". NeuroImage. 132: 190–197. doi:10.1016/j.neuroimage.2016.02.029. ISSN 1053-8119. PMC 4851897. PMID 26899785.
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