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• W3196607911 abstract "•Multiple large-scale cortical networks converged in the thalamus•Neurocognitive associated hub existed in the anterior and medial region•Control-processing hub localized in the intermediate thalamus•Sensorimotor network was located around the lateral pulvinar nucleus The thalamus is critical for the brain's integrative hub functions; however, the localization and characterization of the different thalamic hubs remain unclear. Using a voxel-level network measure called functional connectivity overlap ratio (FCOR), we examined the thalamus' association with large-scale resting-state networks (RSNs) to elucidate its connector hub roles. Connections to the core-neurocognitive networks were localized in the anterior and medial parts, such as the anteroventral and mediodorsal nuclei areas. Regions functionally connected to the sensorimotor network were distinctively located around the lateral pulvinar nucleus but to a limited extent. Prominent connector hubs include the anteroventral, ventral lateral, and mediodorsal nuclei with functional connections to multiple RSNs. These findings suggest that the thalamus, with extensive connections to most of the RSNs, is well placed as a critical integrative functional hub and could play an important role for functional integration facilitating brain functions associated with primary processing and higher cognition. The thalamus is critical for the brain's integrative hub functions; however, the localization and characterization of the different thalamic hubs remain unclear. Using a voxel-level network measure called functional connectivity overlap ratio (FCOR), we examined the thalamus' association with large-scale resting-state networks (RSNs) to elucidate its connector hub roles. Connections to the core-neurocognitive networks were localized in the anterior and medial parts, such as the anteroventral and mediodorsal nuclei areas. Regions functionally connected to the sensorimotor network were distinctively located around the lateral pulvinar nucleus but to a limited extent. Prominent connector hubs include the anteroventral, ventral lateral, and mediodorsal nuclei with functional connections to multiple RSNs. These findings suggest that the thalamus, with extensive connections to most of the RSNs, is well placed as a critical integrative functional hub and could play an important role for functional integration facilitating brain functions associated with primary processing and higher cognition. The thalamus is a nuclear complex consisting of dozens of nuclei located in the diencephalon. The thalamus has been traditionally considered as a relay station in the flow of various sensory signals. More recent evidence, however, has shown that the thalamus has roles in connecting sensory and cognitive processing (Sherman, 2016Sherman S.M. Thalamus plays a central role in ongoing cortical functioning.Nat. Neurosci. 2016; 19: 533-541Crossref PubMed Scopus (352) Google Scholar; Wolff et al., 2020Wolff M. Morceau S. Martin-Cortecero J. Folkard R. Groh A. A thalamic bridge from sensory perception to cognition.Neurosci. Biobehav. Rev. 2020; 120: 222-235Crossref PubMed Scopus (12) Google Scholar). Based on animal studies, Sherman categorized thalamic relays into two types based on their inputs: first-order relays, which receive subcortical driver input, and higher-order relays, which receive input from layer 5 of the cortex and participate in cortico-thalamocortical circuits (Sherman, 2016Sherman S.M. Thalamus plays a central role in ongoing cortical functioning.Nat. Neurosci. 2016; 19: 533-541Crossref PubMed Scopus (352) Google Scholar). The presence of these higher-order relays involved in transthalamic corticocortical communications suggests that the thalamus continues to participate in the processing of information within cortical hierarchies. Citing the role of the thalamic reticular nucleus and modulator inputs to the thalamus, Wolff and colleagues further proposed that the thalamus as a whole, including both first-order and higher-order nuclei, may be critical for integrating environmental signals in cognitive processes and thus serving as a bridge linking sensory perception and cognition (Wolff et al., 2020Wolff M. Morceau S. Martin-Cortecero J. Folkard R. Groh A. A thalamic bridge from sensory perception to cognition.Neurosci. Biobehav. Rev. 2020; 120: 222-235Crossref PubMed Scopus (12) Google Scholar). Although cortical circuitries associated with cognition have been well studied, the role of the thalamus in cognitive processes is just beginning to be elucidated (Halassa and Kastner, 2017Halassa M.M. Kastner S. Thalamic functions in distributed cognitive control.Nat. Neurosci. 2017; 20: 1669-1679Crossref PubMed Scopus (191) Google Scholar). Using neuroimaging, several studies have examined both the anatomical and functional organization of the thalamus in humans. Recent investigations using resting-state functional magnetic resonance imaging (fMRI) have also indicated that the thalamus and basal ganglia connect to several cortical functional networks and contribute to multimodal cognitive functions (Bell and Shine, 2016Bell P.T. Shine J.M. Subcortical contributions to large-scale network communication.Neurosci. Biobehav. Rev. 2016; 71: 313-322Crossref PubMed Scopus (79) Google Scholar; Greene et al., 2020Greene D.J. Marek S. Gordon E.M. Siegel J.S. Gratton C. Laumann T.O. Gilmore A.W. Berg J.J. Nguyen A.L. Dierker D. et al.Integrative and network-specific connectivity of the basal ganglia and thalamus defined in individuals.Neuron. 2020; 105: 742-758.e6Abstract Full Text Full Text PDF PubMed Scopus (68) Google Scholar). Graph-theoretic network analysis revealed that several thalamic subdivisions have network properties capable of integrating information across multiple cortical functional networks (Hwang et al., 2017Hwang K. Bertolero M.A. Liu W.B. D’Esposito M. The human thalamus is an integrative hub for functional brain networks.J. Neurosci. 2017; 37: 5594-5607Crossref PubMed Scopus (221) Google Scholar). With widespread structural and functional connectivity to the cerebral cortex, the thalamus is well positioned to mediate the interactions between distributed, large-scale cortical functional networks, which may be associated with higher brain functions (Shine, 2020Shine J.M. The thalamus integrates the macrosystems of the brain to facilitate complex, adaptive brain network dynamics.Prog. Neurobiol. 2020; 101951Google Scholar). To elucidate the thalamus' integrative functional role, it is necessary to fully understand the thalamus' association with large-scale functional brain networks, which can be considered the brain's functional modules. Considering that thalamic dysfunction is at the core of numerous psychiatric pathologies with varying clinical manifestations such as major depressive disorder, obsessive compulsive disorder, bipolar disorder, and schizophrenia, among others (Parnaudeau et al., 2018Parnaudeau S. Bolkan S.S. Kellendonk C. The mediodorsal thalamus: an essential partner of the prefrontal cortex for cognition.Biol. Psychiatry. 2018; 83: 648-656Abstract Full Text Full Text PDF PubMed Scopus (105) Google Scholar), understanding this functional association would be extremely valuable to characterize these dysfunctions. The goal of this paper is to examine this association comprehensively, identify multi-network integrative hub locations in the thalamus, and clarify the connectivity properties of the identified hub regions. To achieve this, we used a voxel-level functional connectivity (FC) measure called functional connectivity overlap ratio (FCOR) that can be used to quantify the spatial extent of a voxel's connection to several well-known large-scale cortical functional networks (Bagarinao et al., 2020Bagarinao E. Watanabe H. Maesawa S. Mori D. Hara K. Kawabata K. Ohdake R. Masuda M. Ogura A. Kato T. et al.Identifying the brain’s connector hubs at the voxel level using functional connectivity overlap ratio.Neuroimage. 2020; 222: 117241Crossref PubMed Scopus (8) Google Scholar). In our previous report, we successfully obtained voxel-level FCOR measurement to the whole-brain FC to identify cortical connector hubs. Here, we focused on the thalamus' connectivity to the cortex and investigated the extent of each thalamic voxel's connection to the various functional networks, which can provide a more in-depth understanding of the voxel's functional role. The distribution of the thalamic connector hubs was then identified using voxels with extensive connections with not just one but multiple networks (Figure 1). Using the general classification of cortical networks into default mode (e.g. the dorsal and ventral default mode networks), control (e.g., the salience and executive control networks), and processing networks (e.g., the sensorimotor and auditory networks), we further categorized the thalamic connector hub voxels into control-default, cross-control, and control-processing hubs (Bagarinao et al., 2020Bagarinao E. Watanabe H. Maesawa S. Mori D. Hara K. Kawabata K. Ohdake R. Masuda M. Ogura A. Kato T. et al.Identifying the brain’s connector hubs at the voxel level using functional connectivity overlap ratio.Neuroimage. 2020; 222: 117241Crossref PubMed Scopus (8) Google Scholar; Gordon et al., 2018Gordon E.M. Lynch C.J. Gratton C. Laumann T.O. Gilmore A.W. Greene D.J. Ortega M. Nguyen A.L. Schlaggar B.L. Petersen S.E. et al.Three distinct sets of connector hubs integrate human brain function.Cell Rep. 2018; 24: 1687-1695.e4Abstract Full Text Full Text PDF PubMed Scopus (47) Google Scholar). We also parcellated the thalamus into functional subcomponents using FCOR values to identify regions primarily associated with primary processing from those involved in higher cognitive processes. In the analysis, two sets of resting-state network (RSN) templates were employed: one set from Shirer et al. (Shirer et al., 2012Shirer W.R. Ryali S. Rykhlevskaia E. Menon V. Greicius M.D. Decoding subject-driven cognitive states with whole-brain connectivity patterns.Cereb. Cortex. 2012; 22: 158-165Crossref PubMed Scopus (1173) Google Scholar) with 13 RSNs and the other from Gordon, et al. (Gordon et al., 2016Gordon E.M. Laumann T.O. Adeyemo B. Huckins J.F. Kelley W.M. Petersen S.E. Generation and evaluation of a cortical area parcellation from resting-state correlations.Cereb. Cortex. 2016; 26: 288-303Crossref PubMed Scopus (599) Google Scholar) with 10 RSNs (Figure S1). For each RSN template, we calculated FCOR values for all voxels in the thalamus and all participants. The mean values across participants for each RSN were then computed. The results for both the Shirer and Gordon atlases are shown in Figure 2 after applying a threshold value of 0.1 or 10% and in Figure S2A for other threshold values (15% and 20%). Results for FCOR values computed at different false discovery rates (see STAR Methods) are shown in Figure S2B. Of all the RSNs examined, the thalamus showed functional connections to all except the visuospatial (dorsal attention) network with the anterior salience network having the most extensive connections with the thalamus and the language network the least. This finding is consistent for both the Shirer and Gordon atlases and for surviving voxels of seed-based functional connectivity analysis results shown in Figure S2A. For the Shirer atlas (Figure 2A), voxels with significantly higher FCOR values to the core neurocognitive networks (Menon, 2011Menon V. Large-scale brain networks and psychopathology: a unifying triple network model.Trends Cogn. Sci. 2011; 15: 483-506Abstract Full Text Full Text PDF PubMed Scopus (1934) Google Scholar; Seeley et al., 2007Seeley W.W. Menon V. Schatzberg A.F. Keller J. Glover G.H. Kenna H. Reiss A.L. Greicius M.D. Dissociable intrinsic connectivity networks for salience processing and executive control.J. Neurosci. 2007; 27: 2349-2356Crossref PubMed Scopus (4656) Google Scholar), such as dorsal default mode network (dDMN), ventral default mode network (vDMN), precuneus network (Prec), left executive control network (LECN), right executive control network (RECN), anterior salience network (aSal), and posterior salience network (pSal), were predominantly located in the thalamus' medial part. In addition, voxels with higher FCOR values to the LECN and RECN were more localized in the anterior portion. Voxels with stronger FCOR to both anterior salience network (aSal) and posterior salience network (pSal) extensively covered the thalamus except the dorsal lateral part. Even at the 20% threshold, a larger number of voxels with significant connections to aSal and Prec still survived. Voxels associated with the language network were located mostly in the anterior medial thalamus but were very limited in extent, even at a threshold value of 10%. These core-neurocognitive-network associated voxels were predominantly related to the anteroventral (AV), lateral posterior (LP), ventral anterior (VA), and ventral lateral (VL) nuclei as well as mediodorsal medial magnocellular (MDm) and mediodorsal lateral parvocellular (MDl) nuclei as defined in the AAL3 atlas (Figure 3A). Voxels with higher FCOR values to sensory processing networks such as the sensorimotor, auditory, and visual networks were mainly located in the dorsal lateral part of the thalamus. The sensorimotor network-associated voxels were localized around pulvinar lateral (PuL), ventral posterolateral (VPL), intralaminar (IL), and pulvinar anterior (PuA); however, the spatial extent was limited. Voxels with higher FCOR values to the auditory network were located across several nuclei such as IL, MDm, MDl, medial geniculate (MGN), PuA, and PuL, while voxels connected to the primary visual and higher visual networks were located in the dorsal lateral thalamus, around lateral geniculate (LGN), MGN, and PuA. No voxels with connections to the visuospatial (dorsal attention) network survived at 10% threshold FCOR value. The FCOR profile of the thalamus using the Gordon atlas also showed similar distribution. Voxels with significantly higher FCOR values to the core neurocognitive networks (default, fronto-parietal, and cingulo-parietal) of this atlas were similarly located in the medial thalamus, and the distribution of voxels connected to the salience network was also widespread and located in the anterior and medial part of the thalamus (Figure 2B). Stronger FCOR values to the cingulo-parietal and salience networks were also observed with more voxels surviving even at 20% threshold for connections in these networks compared to other networks. Connections to the two sensorimotor networks (hand and mouth related) were located around PuL, PuA, IL, and VPL (Figure 3B). Voxels with higher FCOR values to the visual network were located around LGN, MGN, and PuA, whereas that to the auditory network involved several thalamic nuclei. Like the visuospatial network in the Shirer atlas, no voxels with FCOR values to the dorsal and ventral attention network in the Gordon atlas survived even at 10% threshold. To identify connector hubs in the thalamus, we binarized the FCOR maps shown in Figure 2 by assigning a value of 1 to voxels with FCOR value exceeding the threshold (FCOR = 0.1) and 0 to the rest. The binarized maps of all RSNs were then combined. The resulting maps are shown in Figure 4 for both the Shirer (Figure 4A) and Gordon (Figure 4B) atlases. Voxels showing a higher RSN count represent connector hubs, which could represent potential sites for functional integration. From the figure, these voxels were mostly located in the anterior and medial thalamus, particularly in AV, LP, MDm, and MDl, which exhibited prominent connections with not just one but multiple RSNs for both Shirer and Gordon atlases. Spider plots of these regions indicating their functional connections to the different RSNs are shown in Figure 5 and in the Figure S3.Figure 5Spider plots of FCOR values for subregions with the 4 highest number of connected resting-state networksShow full caption(A–B) Spider plots show the occupancy ratio of the 4 thalamic subregions including the AV, LP, MDm, and MDl with the highest number of overlapped RSNs for the (A) Shirer and (B) Gordon atlases. See also Figure S3.View Large Image Figure ViewerDownload Hi-res image Download (PPT) (A–B) Spider plots show the occupancy ratio of the 4 thalamic subregions including the AV, LP, MDm, and MDl with the highest number of overlapped RSNs for the (A) Shirer and (B) Gordon atlases. See also Figure S3. We further classified connector hub regions in the thalamus into control-default, cross-control, and control-processing connector hubs shown in Figure 6 (Bagarinao et al., 2020Bagarinao E. Watanabe H. Maesawa S. Mori D. Hara K. Kawabata K. Ohdake R. Masuda M. Ogura A. Kato T. et al.Identifying the brain’s connector hubs at the voxel level using functional connectivity overlap ratio.Neuroimage. 2020; 222: 117241Crossref PubMed Scopus (8) Google Scholar; Gordon et al., 2018Gordon E.M. Lynch C.J. Gratton C. Laumann T.O. Gilmore A.W. Greene D.J. Ortega M. Nguyen A.L. Schlaggar B.L. Petersen S.E. et al.Three distinct sets of connector hubs integrate human brain function.Cell Rep. 2018; 24: 1687-1695.e4Abstract Full Text Full Text PDF PubMed Scopus (47) Google Scholar). Control-default connector hubs link control and default mode networks, whereas cross-control hubs link different control networks (see STAR Methods). The control-default and cross-control networks existed in the medial and anterior part of the thalamus. Subregions with the highest mean values of both control-default and cross-control were AV, LP, MDl, and MDm. The control-default and cross-control regions were localized in similar regions, and the peak location of the control-default was in the medial part. On the other hand, the control-processing connector hubs, which link control and primary processing networks, were located in the intermediate part of the thalamus. Intriguingly, all types of connector hubs (control default, cross control, and control processing) converged in the intermedial part of the thalamus (Figure 6D). Subregions with the highest number of RSN connections were MDl, MDm, LP, and PuL. The subregions of LP, MDl, and MDm were associated with both control-default and control-processing hubs, whereas AV and VA were mainly connected to cognitive hubs. The lateral nuclei of LGN, MGN, and PuA were mainly involved in processing networks. Using FCOR values as voxel features, clustering analysis results are shown in Figure 7. The k value with the highest mean silhouette was 2, followed by 3 in both the Shirer and the Gordon atlases. Using k = 2, the thalamus was divided into core-neurocognitive thalamus and sensory processing thalamus but with the auditory network-related regions split into an anterior division belonging to the neurocognitive cluster and a posterior division to the sensory processing cluster. Both the Shirer and Gordon atlases provided consistent parcellation. For the 3-cluster parcellation, the thalamus was divided into a core-neurocognitive cluster, sensory processing cluster, and a third cluster involving both cognitive and sensory processing networks. In the Shirer atlas, the third highest mean silhouette was using 4 clusters, followed by 8 clusters (Figures S4 and S5). For 4-cluster parcellation, the two clusters were still predominantly associated with primary processing and neurocognitive networks, while the remaining two were non-specific with components from auditory and salience networks. For the 8-cluster parcellation, the core-neurocognitive associated cluster was further split into two clusters associated with the default mode and executive control networks and the sensory processing cluster into SMN and visual networks. In the Gordon atlas, the k value with the third highest mean silhouette was 5, followed by 8 (Figures S4 and S5). The resulting clusters using the Gordon templates remained relatively consistent with that of the Shirer atlas. We examined the functional connections of the thalamus to known RSNs using a FC measure called FCOR. Regions strongly connected to the core-neurocognitive networks that include the default mode, salience, and executive control networks were localized predominantly in the anterior and medial thalamus, such as the anteroventral and mediodorsal nuclei areas. Specifically, regions connected to the salience network covered a more widespread thalamic area, regions connected to the default mode networks were mainly located in the medial thalamus, and to the executive control networks were predominantly in the anterior part. No regions with link to the visuospatial (dorsal attention) network were detected in the thalamus. The sensorimotor network regions were located around the lateral pulvinar nucleus, but the spatial extent was limited. The auditory network regions were localized in the more anterior part than the sensorimotor network and across multiple nuclei. The visual network-related regions were in the dorsal lateral thalamus around the lateral geniculate nucleus. The anteroventral, ventral lateral, and mediodorsal nuclei were prominent as connector hubs with connections to multiple RSNs. Both control-default and cross-control connector hubs were concentrated in these regions, and all types of hubs converged in the intermediate part. Our findings showed that the thalamus is extensively connected to almost all RSNs with the posterior regions mainly associated with primary processing networks while a larger subregion significantly involved in networks associated with cognitive functions. Recent studies have investigated the hub properties of the thalamus (Greene et al., 2020Greene D.J. Marek S. Gordon E.M. Siegel J.S. Gratton C. Laumann T.O. Gilmore A.W. Berg J.J. Nguyen A.L. Dierker D. et al.Integrative and network-specific connectivity of the basal ganglia and thalamus defined in individuals.Neuron. 2020; 105: 742-758.e6Abstract Full Text Full Text PDF PubMed Scopus (68) Google Scholar; Hwang et al., 2017Hwang K. Bertolero M.A. Liu W.B. D’Esposito M. The human thalamus is an integrative hub for functional brain networks.J. Neurosci. 2017; 37: 5594-5607Crossref PubMed Scopus (221) Google Scholar). Multiple thalamic subdivisions have been shown to display network properties that could integrate multimodal information across diverse cortical functional networks (Hwang et al., 2017Hwang K. Bertolero M.A. Liu W.B. D’Esposito M. The human thalamus is an integrative hub for functional brain networks.J. Neurosci. 2017; 37: 5594-5607Crossref PubMed Scopus (221) Google Scholar). Some regions were identified as network specific, whereas others were characterized as multi-network integration zones (Greene et al., 2020Greene D.J. Marek S. Gordon E.M. Siegel J.S. Gratton C. Laumann T.O. Gilmore A.W. Berg J.J. Nguyen A.L. Dierker D. et al.Integrative and network-specific connectivity of the basal ganglia and thalamus defined in individuals.Neuron. 2020; 105: 742-758.e6Abstract Full Text Full Text PDF PubMed Scopus (68) Google Scholar). In this study, we further categorized the divergent functional integrative hub regions in the thalamus with anatomical subcomponents. In addition, we also classified connector hubs into different categories such as control-processing, control-default, and cross-control and thalamic voxels into clusters, which appeared to be arranged into a topographical motif (Bagarinao et al., 2020Bagarinao E. Watanabe H. Maesawa S. Mori D. Hara K. Kawabata K. Ohdake R. Masuda M. Ogura A. Kato T. et al.Identifying the brain’s connector hubs at the voxel level using functional connectivity overlap ratio.Neuroimage. 2020; 222: 117241Crossref PubMed Scopus (8) Google Scholar; Power et al., 2011Power J.D. Cohen A.L. Nelson S.M. Wig G.S. Barnes K.A. Church J.A. Vogel A.C. Laumann T.O. Miezin F.M. Schlaggar B.L. et al.Functional network organization of the human brain.Neuron. 2011; 72: 665-678Abstract Full Text Full Text PDF PubMed Scopus (2342) Google Scholar) with a cluster associated with the core neurocognitive networks at one end and a cluster associated with primary processing networks at the other end. This provides a more extensive characterization of the functional connectivity profile of the thalamus. To achieve all of these, we used the same FCOR metric. Approaches to identify connector hubs commonly used the network metric called participation coefficient (Guimerà and Nunes Amaral, 2005Guimerà R. Nunes Amaral L.A. Functional cartography of complex metabolic networks.Nature. 2005; 433: 895-900Crossref PubMed Scopus (2422) Google Scholar; Rubinov and Sporns, 2010Rubinov M. Sporns O. Complex network measures of brain connectivity: uses and interpretations.Neuroimage. 2010; 52: 1059-1069Crossref PubMed Scopus (6443) Google Scholar). Estimating this metric at the voxel-level resolution with nodes reaching hundreds of thousands would be computationally challenging, particularly in terms of memory requirement. There is also the intermediate step of identifying community membership of each node, which requires an additional clustering step. Thus, to minimize the needed computation, an initial brain parcellation would be necessary to reduce nodes to a few hundred, limiting the spatial resolution of identified connector hubs to the size of the parcellation. Identifying connector hubs in the thalamus requires resolution at the voxel level given the overall size of the thalamus as well as its dense and extensive connectivity to the cortex. This problem can be properly addressed by using FCOR, which can be used to identify regions with high between-network connectivity at the voxel level. Moreover, FCOR can also be independently computed at each voxel so that only FCOR values at specific regions of interest need to be computed as we have demonstrated in this study. Therefore, it enables voxel-level mapping of thalamic hub regions, even though the thalamus is relatively a small structure. We have shown that the anterior subregions of AV and LP and medial parts of MDm and MDl, regions critical for cognitive processing and global amnesia (Mair et al., 2015Mair R.G. Miller R.L.A. Wormwood B.A. Francoeur M.J. Onos K.D. Gibson B.M. The neurobiology of thalamic amnesia: contributions of medial thalamus and prefrontal cortex to delayed conditional discrimination.Neurosci. Biobehav. Rev. 2015; 54: 161-174Crossref PubMed Scopus (20) Google Scholar), were highly connected to multiple RSNs predominantly associated with cognitive processing. The anterior nucleus is a crucial component of the hippocampal system for episodic memory (Child and Benarroch, 2013Child N.D. Benarroch E.E. Anterior nucleus of the thalamus: functional organization and clinical implications.Neurology. 2013; 81: 1869-1876Crossref PubMed Scopus (137) Google Scholar), and damages of this region manifest memory and language impairment (Nishio et al., 2011Nishio Y. Hashimoto M. Ishii K. Mori E. Neuroanatomy of a neurobehavioral disturbance in the left anterior thalamic infarction.J. Neurol. Neurosurg. Psychiatry. 2011; 82: 1195-1200Crossref PubMed Scopus (39) Google Scholar). The mediodorsal thalamus is critical for long-term memory and several cognitive functions (Pergola et al., 2018Pergola G. Danet L. Pitel A.L. Carlesimo G.A. Segobin S. Pariente J. Suchan B. Mitchell A.S. Barbeau E.J. The regulatory role of the human mediodorsal thalamus.Trends Cogn. Sci. 2018; 22: 1011-1025Abstract Full Text Full Text PDF PubMed Scopus (69) Google Scholar) and closely interacts with the prefrontal cortex in line with multiple cognitive tasks such as working memory, attentional control, and cognitive flexibility (Mitchell, 2015Mitchell A.S. The mediodorsal thalamus as a higher order thalamic relay nucleus important for learning and decision-making.Neurosci. Biobehav. Rev. 2015; 54: 76-88Crossref PubMed Scopus (152) Google Scholar; Mitchell and Chakraborty, 2013Mitchell A.S. Chakraborty S. What does the mediodorsal thalamus do?.Front. Syst. Neurosci. 2013; 7: 1-19Crossref PubMed Scopus (161) Google Scholar; Parnaudeau et al., 2013Parnaudeau S. O’Neill P.K. Bolkan S.S. Ward R.D. Abbas A.I. Roth B.L. Balsam P.D. Gordon J.A. Kellendonk C. Inhibition of mediodorsal thalamus disrupts thalamofrontal connectivity and cognition.Neuron. 2013; 77: 1151-1162Abstract Full Text Full Text PDF PubMed Scopus (229) Google Scholar, Parnaudeau et al., 2018Parnaudeau S. Bolkan S.S. Kellendonk C. The mediodorsal thalamus: an essential partner of the prefrontal cortex for cognition.Biol. Psychiatry. 2018; 83: 648-656Abstract Full Text Full Text PDF PubMed Scopus (105) Google Scholar; Rikhye et al., 2018Rikhye R.V. Wimmer R.D. Halassa M.M. Toward an integrative theory of thalamic function.Annu. Rev. Neurosci. 2018; 41: 163-183Crossref PubMed Scopus (58) Google Scholar; Schmitt et al., 2017Schmitt L.I. Wimmer R.D. Nakajima M. Happ M. Mofakham S. Halassa M.M. Thalamic amplification of cortical connectivity sustains attentional control.Nature. 2017; 545: 219-223Crossref PubMed Scopus (279) Google Scholar). These functions are mostly associated with the default mode and control networks. Thus, our findings showing that cross-control and control-default connector hubs concentrated in the medial and anterior thalamus are consistent with these functional roles, supporting the idea that connector hubs in these regions are critical for cognitive functions. The core neurocognitive networks, including the default mode, salience, and executive control networks, are predominantly connected to the anterior and medial thalamus, such as the anteroventral and mediodorsal nuclei areas, with some networks (salience from both Shirer and Gordon atl" @default.
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• W3196607911 title "Bridging large-scale cortical networks: Integrative and function-specific hubs in the thalamus" @default.
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• W3196607911 doi "https://doi.org/10.1016/j.isci.2021.103106" @default.
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