Co-activation patterns distinguish cortical modules, their connectivity and functional differentiation

“…physiology of correlations in the absence of a structured task remains somewhat elusive, it seems plausible that “resting” state, as a mixture of various cognitive processes, may likewise sample the repertoire of operations brain networks can perform (Buckner and Vincent, 2007; Smith et al., 2009).”

“Other approaches such as psychophysiological interactions or Granger causalitymapping are used to identify regions, showing a context-dependent change in their coupling with a particular seed region (Friston et al., 1997; Goebel et al., 2003) under the assumptions of the employed model.”

“…likely conveying meaningful functional relationships between brain regions are illustrated by the fact that they were reported to widely correspond with both task-state networks (Biswal et al., 1995; Smith et al., 2009) and structural connectivity (Greicius et al., 2009; Hagmann et al., 2008).”

“In addition, a necessary step toward understanding cortical organization is to relate connectivity-defined modules and functional differentiations to microstructural maps of the human cerebral cortex (Amunts et al., 2007; Eickhoff et al., 2005; Eickhoff et al., 2008b; Zilles et al., 2002).”

“Other approaches such as psychophysiological interactions or Granger causalitymapping are used to identify regions, showing a context-dependent change in their coupling with a particular seed region (Friston et al., 1997; Goebel et al., 2003) under the assumptions of the employed model.”

“On these, tractography algorithms may be employed to derive information about the most likely fiber tract direction from particular seed or the course and strength of pathways connecting two different regions of the cortex (Behrens et al., 2003; Ramnani et al., 2004).”

“…likely conveying meaningful functional relationships between brain regions are illustrated by the fact that they were reported to widely correspond with both task-state networks (Biswal et al., 1995; Smith et al., 2009) and structural connectivity (Greicius et al., 2009; Hagmann et al., 2008).”

“…described co-activation parcellation of the seed region, ALE scores for the MACM analysis of this cluster were compared to a null-distribution that reflects a random spatial association between experiments, but regards the within-experiment distribution of foci as fixed (Eickhoff et al., 2009).”

“It thus summarizes any analysis that assesses correlations among brain signals without the explicit modeling of the underlying networks (as in effective connectivity), ranging from spike train correlations (Nuding et al., 2009) to EEG coherence (Gross et al., 2001) and various fMRI basedmethods.”

“Based on these observations and functional activation studies, the SMA has been implicated in executive aspects of motor control, e.g. movement initiation (Chouinard and Paus, 2006; Cunnington et al., 2002; Eickhoff et al., 2008a; Picard and Strick, 1996).”

“Together with previous computational modeling (Honey et al., 2007), this supports the notion of a good congruency between anatomical and functional connectivities.”

“Regions that co-activated with both clusters delineated in the medial premotor cortex by the co-activation based parcellation were identified using a minimum-statistic conjunction by computing the intersection of the thresholded ALE-maps (Caspers et al., 2010; Kurth et al., 2010).”

“Regions that co-activated with both clusters delineated in the medial premotor cortex by the co-activation based parcellation were identified using a minimum-statistic conjunction by computing the intersection of the thresholded ALE-maps (Caspers et al., 2010; Kurth et al., 2010).”

“Fax: + E-mail address: S.Eickhoff@fz-juelich.de (S.B. Eickho 1053-8119/$ – see front matter © 2011 Elsevier Inc. Al doi:10.1016/j.neuroimage.2011.05.021 a b s t r a c t a r t i c l e i n f o”

“Supporting this view, connectivity tracing in non-human primates revealed that the preSMA receives afferences from the inferior parietal lobule (Luppino et al., 1993) and the prefrontal cortex.”

“Approaches investigating cortical sub-specialization capitalized on this type of inter-regional connectivity, such as for the successful parcellation of the premotor cortex (Kim et al., 2010), insular cortex (Cauda et al., 2011), and thalamus (Zhang et al., 2008).”

“In the widely used Dynamic Causal Modeling (DCM) and Structural Equation Modeling (SEM) approaches, this is assessed by taking an explicit network perspective and modeling the interactions between regions as a function of the experimental context (Buchel and Friston, 1997; Stephan et al., 2007).”

“The latter correlate regional BOLD signal changes in order to quantify the degree of functional connectivity between them or identify regions that correlate with a particular seed (Hampson et al., 2002; Ramnani et al., 2004; Xiong et al., 1999).”

“Following earlier reports that cortical modules may be defined by differences in anatomical connectivity (Johansen-Berg et al., 2004; Klein et al., 2007; Tomassini et al., 2007) or resting state correlations (Kim et al.”

“Other approaches such as psychophysiological interactions or Granger causalitymapping are used to identify regions, showing a context-dependent change in their coupling with a particular seed region (Friston et al., 1997; Goebel et al., 2003) under the assumptions of the employed model.”

“Evidence from primate research indicated that microstructure and connectivity of the cortex are the main determinants of its functional segregation (Luppino et al., 1991; Matelli et al., 1991).”

“Analysis of the behavioral domain and paradigm class meta-data (Laird et al., 2009a) moreover allowed the delineation of functional characteristics for the ensuing cortical modules.”

“Based on these observations and functional activation studies, the SMA has been implicated in executive aspects of motor control, e.g. movement initiation (Chouinard and Paus, 2006; Cunnington et al., 2002; Eickhoff et al., 2008a; Picard and Strick, 1996).”

“Evidence from primate research indicated that microstructure and connectivity of the cortex are the main determinants of its functional segregation (Luppino et al., 1991; Matelli et al., 1991).”

“By comparing the anatomical connectivity profiles of the individual seed voxels with each other, these can then be grouped into distinct clusters of homogeneous connectivity (Anwander et al., 2007; Johansen-Berg et al., 2004; Klein et al., 2007).”

“While the underlying physiology of correlations in the absence of a structured task remains somewhat elusive, it seems plausible that “resting” state, as a mixture of various cognitive processes, may likewise sample the repertoire of operations brain networks can perform (Buckner and Vincent, 2007; Smith et al., 2009).”

“Approaches investigating cortical sub-specialization capitalized on this type of inter-regional connectivity, such as for the successful parcellation of the premotor cortex (Kim et al., 2010), insular cortex (Cauda et al., 2011), and thalamus (Zhang et al., 2008).”

“Among those, effective connectivity analyses, such as dynamic causal modeling (Friston et al., 2003) or structural equation modeling (Buchel and Friston, 1997), allow the investigation of taskdependent influences among cortical areas (Grefkes et al., 2008).”

“DTI, resting state analyses, and MACM each focus on a different aspects of connectivity, and we therefore predict that in spite of their good convergence, each capture different features of cortical organization (Eickhoff et al., 2010; Honey et al., 2007; Ramnani et al., 2004).”

“Based on these observations and functional activation studies, the SMA has been implicated in executive aspects of motor control, e.g. movement initiation (Chouinard and Paus, 2006; Cunnington et al., 2002; Eickhoff et al., 2008a; Picard and Strick, 1996).”

“While many hypotheses derived in particular from primate work (Bremmer et al., 2001) and lesion mapping studies (Riecker et al., 2005) could be explicitly tested using this approach, neuroimaging is intrinsically less well suited to delineate the organization of a particular brain region.”

“…earlier reports that cortical modules may be defined by differences in anatomical connectivity (Johansen-Berg et al., 2004; Klein et al., 2007; Tomassini et al., 2007) or resting state correlations (Kim et al., 2010) we demonstrated that SMA and pre-SMA may be distinguished by amodel-free…”

“and Luppino, 2001; Vogt et al., 2007).”

“, 2007) or resting state correlations (Kim et al., 2010) we demonstrated that SMA and pre-SMA may be distinguished by amodel-free analysis of co-activation patterns across activation studies.”

“In addition, a necessary step toward understanding cortical organization is to relate connectivity-defined modules and functional differentiations to microstructural maps of the human cerebral cortex (Amunts et al., 2007; Eickhoff et al., 2005; Eickhoff et al., 2008b; Zilles et al., 2002).”

“On these, tractography algorithms may be employed to derive information about the most likely fiber tract direction from particular seed or the course and strength of pathways connecting two different regions of the cortex (Behrens et al., 2003; Ramnani et al., 2004).”

“In addition, a necessary step toward understanding cortical organization is to relate connectivity-defined modules and functional differentiations to microstructural maps of the human cerebral cortex (Amunts et al., 2007; Eickhoff et al., 2005; Eickhoff et al., 2008b; Zilles et al., 2002).”

“In the widely used Dynamic Causal Modeling (DCM) and Structural Equation Modeling (SEM) approaches, this is assessed by taking an explicit network perspective and modeling the interactions between regions as a function of the experimental context (Buchel and Friston, 1997; Stephan et al., 2007).”

“Following earlier reports that cortical modules may be defined by differences in anatomical connectivity (Johansen-Berg et al., 2004; Klein et al., 2007; Tomassini et al., 2007) or resting state correlations (Kim et al., 2010) we demonstrated that SMA and pre-SMA may be distinguished by amodel-free…”

“, a functional differentiation between two regions (Reddy and Kanwisher, 2006) or a dichotomy between the neural correlates of two processes (Charron and Koechlin, 2010).”

“, a functional differentiation between two regions (Reddy and Kanwisher, 2006) or a dichotomy between the neural correlates of two processes (Charron and Koechlin, 2010).”

“…that diffusion-based tractography does not provide information about anatomical connectivity sensu stricto, i.e. axonal connectivity as revealed by tracer studies in macaques (Pons and Kaas, 1985), but can merely assess the presence and strength of macroanatomical fiber bundles between regions.”

“We then assessed whether the two original regions could be recovered in a model-free analysis from this merged, i.e., single, VOI based on similarities between co-activation patterns of the individual seed voxels across neuroimaging experiments.”

“Based on these observations and functional activation studies, the SMA has been implicated in executive aspects of motor control, e.g. movement initiation (Chouinard and Paus, 2006; Cunnington et al., 2002; Eickhoff et al., 2008a; Picard and Strick, 1996).”

“Sets of voxels that feature similar brain-wide coactivation profiles were identified by hierarchical cluster analysis (Eickhoff et al., 2007; Timm, 2002).”

“Based on these observations and functional activation studies, the SMA has been implicated in executive aspects of motor control, e.g. movement initiation (Chouinard and Paus, 2006; Cunnington et al., 2002; Eickhoff et al., 2008a; Picard and Strick, 1996).”

“It thus summarizes any analysis that assesses correlations among brain signals without the explicit modeling of the underlying networks (as in effective connectivity), ranging from spike train correlations (Nuding et al., 2009) to EEG coherence (Gross et al., 2001) and various fMRI basedmethods.”

“…brain region or provide more comprehensive maps of the cerebral cortex (Tzourio-Mazoyer et al. 2002; Desikan et al. 2006; Cohen, Fair, et al. 2008; Cohen, Lombardo, et al. 2008; Eickhoff et al. 2011; Wang et al. 2012; Fan et al. 2014; Wig et al. 2014; Laumann Timothy et al. 2015; Figure 4.”

“Differences between conditions were tested by first performing separate ALE analyses for each condition and computing the voxel-wise difference between the ensuing ALE maps (cf. Eickhoff et al., 2011b).”

“MACM is based on assessing the brain-wise co-activation patterns of a seed region across a large number of data based neuroimaging results (Eickhoff et al., 2011; Laird et al., 2009b).”

“This procedure should effectively balance inference on reliable areas of activation without generating a bias toward false positives (Laird et al., 2005; Eickhoff et al., 2011).”

“The third and last step distinguished between random and ‘true’ convergence by comparing the ensuing ALE scores against an empirical null distribution reflecting a random spatial association between the experiments’ MA maps (Eickhoff et al. 2012).”

“In recent years, several openaccess frameworks have been developed to pool data from thousands of neuroimaging studies (Eickhoff et al., 2011; Poldrack and Yarkoni, 2016).”

“Such meta-analytic measures of task-related coactivation have been explored using “seed-based” analysis (identifying which other regions are coactivated with an arbitrary region or “seed”) (15, 16) and by independent component analysis (ICA) (identifying a set of components or systems of regions that are consistently…”

“groups, keeping the sample size of the two original data sets (Eickhoff et al., 2011; Rottschy et al., 2012).”

“05, cluster-level corrected for multiple comparisons (Eickhoff et al., 2011).”

“In light of this, connectivity-based parcellation informed by MACM might serve as a useful tool to generate hypotheses for targeted experiments assessing structural, connectional, or functional properties and their disturbances [Eickhoff et al., 2011].”

“For these contrasts, a subtraction analysis was run to identify distinct areas of activation (Eickhoff et al., 2011), by repeating the following process 5,000 times: (i) all the results that contributed to either of the two contrasts being examined, were pooled and randomly split into two groups that were the same siz…”

“, 2015), meta-analytic connectivity techniques (Eickhoff et al., 2011; Power et al., 2011), dictionary learning (Varoquaux et al.”

“Keywords: action, connectivity, database, fMRI, prefrontal”

“The technique of ALE subtraction analysis is new (Eickhoff et al., 2011), and the effect of unbalanced groups on subtraction analysis results has not yet been determined.”

“…as gyro-66 sulcal anatomy ( Desikan et al. , 2006; Klein and Tourville , 2012), post-mortem in vitro receptor autoradiography for cyto-architecture ( Eickhoff et al. , 2008; Fischl et al. , 2008), anatomical connectivity ( Roca et al. , 2010) with diffusion imaging, or functional features with…”

“…such as diffusion tractography (Johansen-Berg et al., 2004; Tomassini et al., 2007; Beckmann et al., 2009), resting-state fMRI (Cohen et al., 2008; Kelly et al., 2010; Kim et al., 2010; Nelson et al., 2010; Cauda et al., 2011), and coactivations (Wager et al., 2009; Eickhoff et al., 2011).”

“That is, RSFC and MACM (but not dMRI) may be driven by indirect connections.”

“In addition, algorithms have been provided for meta-analytic contrast analyses using fixed-effects [Laird et al., 2005] and random-effects [Eickhoff et al., 2011] models.”

“Hence, pooling across the close spatial neighborhood has become the dominant approach in MACM analysis (Cauda et al., 2011; Eickhoff et al., 2011).”

“It has to be further emphasised that our approach with definition of seed and target areas is different from the connectivity-based parcellationwhich aims at parcellation of a cortical region based on its connectivity pattern to every other voxel in the brain (Anwander et al., 2007; Behrens et al., 2003a; Eickhoff et…”

“organization from neural circuits through large-scale cortical networks relate to behavior (18-20), the age-dependent changes that underpin hierarchical shifts across”

“All experiments contributing to either analysis were then pooled and randomly divided into two groups of the same size as the two original sets of experiments defined by activation in the first or second cluster (Eickhoff et al., 2011).”

“The observed discrepancies might have resulted from different causes, such as systematic or unsystematic measurement errors and noise or conceptual differences between anatomical and FC [Eickhoff et al., 2011; Eickhoff et al., 2010].”

“Contrasts and conjunctions (i.e., overlaps) between ALE meta-analyses were all done using ten thousand permutations to derive p-values (Eickhoff et al., 2011).”

“Second, hierarchical clustering is an approach in which clusters are formed by first linking the seeds that were more similar to each other and then performing a complete linkage of the clusters (Eickhoff et al. 2011).”

“GingerALE 2.1 also includes a more streamlined interface and a revision of the subtraction analysis [40] originally developed by Laird et al. [37] that has been substantially improved using the new statistical framework developed by Eickhoff et al. [38].”

“j o u r n a l h o m e p a g e : w w w . e l s e v i e r . c o m / l o c a t e / y n i m g (Johansen-Berg et al., 2004), resting state functional connectivity (Zhang and Li, 2012), and meta-analytic connectivity modeling (Cauda et al., 2012; Eickhoff et al., 2011).”

“…co-activation networks, but also functional brain networks derived from meta-analytic connectivity modeling (Laird et al., 2009; Eickhoff et al., 2010; Robinson et al., 2010; Zald et al., 2012) and connectivity-based parcellation (Eickhoff et al., 2011; Bzdok et al., 2013; Cieslik et al., 2013).”

“Conjunction and contrast analyses were computed to examine overlapping and distinct brain regions for the two ALE maps that included both active and passive tasks for symbolic and nonsymbolic number processing (Eickhoff et al., 2011).”

“A subtraction analysis [Eickhoff et al., 2011] between the original and the flipped foci was then performed to compute the voxel-wise difference between the two ALE maps, respectively.”

“The SMA and pre-SMA division has also been seen by others [52–54].”

“These analyses were computed using the voxel-wise minimum value of the input ALE images such that only significant clusters in the individual analyses were included [Eickhoff et al., 2011].”

“Permutations at the contrast level are used to correct for variability among studies (Eickhoff et al., 2011).”

“In light of the as-yet limited number of studies combining MACM and resting state functional connectivity analyses (Cieslik et al., 2013; Eickhoff et al., 2011; Jakobs et al., 2012; Reetz et al., 2012; Rottschy et al., in press), the above results obtained for the V5 region that was shown to be…”

“05; Eickhoff et al., 2011; Nickl-Jockschat et al., 2012).”

“Contrast Analysis To determine whether regions showed significantly different levels of coactivation with the mOFC versus lOFC, we performed a contrast analysis that computed the voxel-wise difference between ALE scores for the 2 sets of coactivation foci (Eickhoff et al. 2011).”

“Analyses of activation peaks were performed using activation likelihood estimation via GingerALE software developed by the Human Brain Mapping Project (ALE) [Eickhoff et al., 2009, 2011, 2012; Turkeltaub et al., 2012].”

“Eickhoff and colleagues (2011) applied connectivity-based parcellation to BrainMap data for the same brain regions (SMA and preSMA) and obtained virtually identical borders (Eickhoff et al. 2011).”

“…of different methods should be less susceptible to drawbacks inherent in each individual method (MACM, resting-state analysis), the conjunction analyses should moreover represent a conservative approach to FC [Eickhoff et al., 2011; Jakobs et al., 2012; Reetz et al., 2012; Smith et al., 2009].”

“All experiments contributing to either analysis were then pooled and randomly divided into 2 groups of the same size as the 2 original sets of experiments defined by activation in the first or second cluster (Eickhoff et al. 2011).”

“To discover clusters with a higher likelihood of activation by affective touch compared with discriminative touch, a contrast between the affective and discriminative ALE maps was performed [Eickhoff et al., 2011].”

“In addition to examining convergence across all empathy-related studies, we performed several contrasts to test for statistical differences across convergence (Eickhoff et al., 2011).”

“This was achieved through an agglomerative approach in which clusters initially formed by individual regions that are subsequently merged according to their similarity using standardized Euclidean distances and Ward's incremental sum of squares method (Eickhoff et al., 2011; Timm, 2002).”

“To explore the BD associated with the FC network, we added to the ALE analysis a MACM. r 1656 r Importantly, the MACM allows meta-analyses, which instead of pooling studies that share a common experimental design, look for global coactivation patterns across a diverse range of tasks, thus responding to the question “f…”

“…connected regions using connectivity determined by stimulation based TMS/PET (Fox et al., 1997, 2006; Paus et al., 1997; Laird et al., 2008) and by meta-analytic connectivity modeling (MACM) (Fox et al., 1998; Robinson et al., 2010; Cauda et al., 2011; Eickhoff et al., 2011; Torta and Cauda, 2011).”

“The SPM generated from these groups are subsequently subtracted from one another to yield ALE difference maps, with the difference statistics averaged across 10000 iterations to obtain the null distribution ALE SPM used to determine the likelihood of the foci observed in the review.(24) Although Britton et al.”

“Significance was established using a binomial test [P< 0.001; Eickhoff et al., 2011; Laird et al., 2009].”

“, Set A: 13 early bilingual studies, 27 late bilingual studies) can be corrected by this approach (Eickhoff et al., 2011): Simulated data is created by pooling the foci data sets and randomly dividing them into two groups of the same size as the original data sets (e.”

“For each comparison, a conjunction image was created, using the voxel-wise minimum value of the included ALE images, to display the similarity between datasets (Eickhoff et al., 2011).”

“A recent meta-analysis on pre-SMA and SMA, including coordinates very similar to the regions found in our study, suggested that the pre-SMA is more involved in complex or ‘cognitive’ aspects of motor inhibition, for example (successful) motor inhibition (Eickhoff et al., 2011).”

“Clusters of overlap between MACM and RS-connectivity analyses for the left vp-MTG (A), left STG (B), and left dp-MTG (C) seed region.”

“The limited number of studies should not exert a strong influence on the results (Eickhoff et al., 2009, 2011).”

“A comparison (subtraction) analysis of two ALE results was conducted using the algorithm in Eickhoff et al. (2011).”

“This is achieved through an agglomerative approach in which clusters initially formed by individual regions are subsequently merged according to their similarity using standardized Euclidean distances and Ward's incremental sum of squares method (Eickhoff et al., 2011; Timm and Kruse, 2002).”

“Notably, there are still no methods to correct for ALE difference maps for multiple comparisons (Eickhoff et al., 2011), and given the small number of papers in the sub-sample analyses (8–24 experiments), an uncorrected threshold of p < 0.”

“Functional magnetic resonance imaging (fMRI) We performed high resolution resting state fMRI to further characterize the neurofunctional segments identified by CBP. Twenty-three healthy individuals (21 righthanded, 7 males, 16 females, M 6 SD 5 21.17 6 1.44)1 were 1For both resting state and DTI, we carried out analys…”

“MACM assesses brain-wise co-activation patterns of an anatomical region across a large number of databased neuroimaging results [Eickhoff et al., 2011; Laird et al., 2009a].”

“We then submitted the obtained thresholded ALE maps from the three individual meta-analyses to a meta-analytic contrast analysis (Eickhoff et al., 2011), which produced the conjunction of the Stop > Go & No-Think > Think contrasts.”

“Identification of contiguous areas of cortex that exhibit similar network connectivity can be used to define a functional parcellation (Raichle et al., 2001; Sporns et al., 2005, Smith et al., 2013, Kim et al., 2010; Eickhoff et al., 2011; Eickhoff et al., 2015).”

“05 corrected for multiple comparisons using FDR method) [Eickhoff et al., 2011].”

“Connectivity-based parcellation approaches have been applied to diffusion tractography [Jbabdi et al., 2009], resting state functional connectivity [Kim et al., 2010], and MACM [Eickhoff et al., 2011] to define a region’s functional anatomy.”