Phase-encoded fMRI tracks down brainstorms of natural language processing with subsecond precision.

Natural language processing unfolds information overtime as spatially separated, multimodal, and interconnected neural processes. Existing noninvasive subtraction-based neuroimaging techniques cannot simultaneously achieve the spatial and temporal resolutions required to visualize ongoing information flows across the whole brain. Here we have developed rapid phase-encoded designs to fully exploit the temporal information latent in functional magnetic resonance imaging data, as well as overcoming scanner noise and head-motion challenges during overt language tasks. We captured real-time information flows as coherent hemodynamic waves traveling over the cortical surface during listening, reading aloud, reciting, and oral cross-language interpreting tasks. We were able to observe the timing, location, direction, and surge of traveling waves in all language tasks, which were visualized as "brainstorms" on brain "weather" maps. The paths of hemodynamic traveling waves provide direct evidence for dual-stream models of the visual and auditory systems as well as logistics models for crossmodal and cross-language processing. Specifically, we have tracked down the step-by-step processing of written or spoken sentences first being received and processed by the visual or auditory streams, carried across language and domain-general cognitive regions, and finally delivered as overt speeches monitored through the auditory cortex, which gives a complete picture of information flows across the brain during natural language functioning. PRACTITIONER POINTS: Phase-encoded fMRI enables simultaneous imaging of high spatial and temporal resolution, capturing continuous spatiotemporal dynamics of the entire brain during real-time overt natural language tasks. Spatiotemporal traveling wave patterns provide direct evidence for constructing comprehensive and explicit models of human information processing. This study unlocks the potential of applying rapid phase-encoded fMRI to indirectly track the underlying neural information flows of sequential sensory, motor, and high-order cognitive processes.

The human cerebellum has almost 80% of the surface area of the neocortex.

The surface of the human cerebellar cortex is much more tightly folded than the cerebral cortex. It was computationally reconstructed for the first time to the level of all individual folia from multicontrast high-resolution postmortem MRI scans. Its total shrinkage-corrected surface area (1,590 cm2) was larger than expected or previously reported, equal to 78% of the total surface area of the human neocortex. The unfolded and flattened surface comprised a narrow strip 10 cm wide but almost 1 m long. By applying the same methods to the neocortex and cerebellum of the macaque monkey, we found that its cerebellum was relatively much smaller, approximately 33% of the total surface area of its neocortex. This suggests a prominent role for the cerebellum in the evolution of distinctively human behaviors and cognition.

The ventral striatum dissociates information expectation, reward anticipation, and reward receipt.

Do dopaminergic reward structures represent the expected utility of information similarly to a reward? Optimal experimental design models from Bayesian decision theory and statistics have proposed a theoretical framework for quantifying the expected value of information that might result from a query. In particular, this formulation quantifies the value of information before the answer to that query is known, in situations where payoffs are unknown and the goal is purely epistemic: That is, to increase knowledge about the state of the world. Whether and how such a theoretical quantity is represented in the brain is unknown. Here we use an event-related functional MRI (fMRI) task design to disentangle information expectation, information revelation and categorization outcome anticipation, and response-contingent reward processing in a visual probabilistic categorization task. We identify a neural signature corresponding to the expectation of information, involving the left lateral ventral striatum. Moreover, we show a temporal dissociation in the activation of different reward-related regions, including the nucleus accumbens, medial prefrontal cortex, and orbitofrontal cortex, during information expectation versus reward-related processing.

Inferior Occipital Gyrus Is Organized along Common Gradients of Spatial and Face-Part Selectivity.

The ventral visual stream of the human brain is subdivided into patches with categorical stimulus preferences, like faces or scenes. However, the functional organization within these areas is less clear. Here, we used functional magnetic resonance imaging and vertex-wise tuning models to independently probe spatial and face-part preferences in the inferior occipital gyrus (IOG) of healthy adult males and females. The majority of responses were well explained by Gaussian population tuning curves for both retinotopic location and the preferred relative position within a face. Parameter maps revealed a common gradient of spatial and face-part selectivity, with the width of tuning curves drastically increasing from posterior to anterior IOG. Tuning peaks clustered more idiosyncratically but were also correlated across maps of visual and face space. Preferences for the upper visual field went along with significantly increased coverage of the upper half of the face, matching recently discovered biases in human perception. Our findings reveal a broad range of neural face-part selectivity in IOG, ranging from narrow to "holistic." IOG is functionally organized along this gradient, which in turn is correlated with retinotopy.SIGNIFICANCE STATEMENT Brain imaging has revealed a lot about the large-scale organization of the human brain and visual system. For example, occipital cortex contains map-like representations of the visual field, while neurons in ventral areas cluster into patches with categorical preferences, like faces or scenes. Much less is known about the functional organization within these areas. Here, we focused on a well established face-preferring area-the inferior occipital gyrus (IOG). A novel neuroimaging paradigm allowed us to map the retinotopic and face-part tuning of many recording sites in IOG independently. We found a steep posterior-anterior gradient of decreasing face-part selectivity, which correlated with retinotopy. This suggests the functional role of ventral areas is not uniform and may follow retinotopic "protomaps."

Reconstructing neural representations of tactile space.

Psychophysical experiments have demonstrated large and highly systematic perceptual distortions of tactile space. Such a space can be referred to our experience of the spatial organisation of objects, at representational level, through touch, in analogy with the familiar concept of visual space. We investigated the neural basis of tactile space by analysing activity patterns induced by tactile stimulation of nine points on a 3 × 3 square grid on the hand dorsum using functional magnetic resonance imaging. We used a searchlight approach within pre-defined regions of interests to compute the pairwise Euclidean distances between the activity patterns elicited by tactile stimulation. Then, we used multidimensional scaling to reconstruct tactile space at the neural level and compare it with skin space at the perceptual level. Our reconstructions of the shape of skin space in contralateral primary somatosensory and motor cortices reveal that it is distorted in a way that matches the perceptual shape of skin space. This suggests that early sensorimotor areas critically contribute to the distorted internal representation of tactile space on the hand dorsum.

Multiple b-values improve discrimination of cortical gray matter regions using diffusion MRI: an experimental validation with a data-driven approach.

OBJECTIVE: To investigate whether varied or repeated b-values provide better diffusion MRI data for discriminating cortical areas with a data-driven approach. METHODS: Data were acquired from three volunteers at 1.5T with b-values of 800, 1400, 2000 s/mm2 along 64 diffusion-encoding directions. The diffusion signal was sampled from gray matter in seven regions of interest (ROIs). Rotational invariants of the local diffusion profile were extracted as features that characterize local tissue properties. Random forest classification experiments assessed whether classification accuracy improved when data with multiple b-values were used over repeated acquisition of the same (1400 s/mm2) b-value to compare all possible pairs of the seven ROIs. Three data sets from the Human Connectome Project were subjected to similar processing and analysis pipelines in eight ROIs. RESULTS: Three different b-values showed an average improvement in correct classification rates of 5.6% and 4.6%, respectively, in the local and HCP data over repeated measurements of the same b-value. The improvement in correct classification rate reached as high as 16% for individual binary classification experiments between two ROIs. Often using only two of the available three b-values were adequate to make such an improvement in classification rates. CONCLUSION: Acquisitions with varying b-values are more suitable for discriminating cortical areas.

Fine-Grained Mapping of Cortical Somatotopies in Chronic Complex Regional Pain Syndrome.

It has long been thought that severe chronic pain conditions, such as complex regional pain syndrome (CRPS), are not only associated with, but even maintained by a reorganization of the somatotopic representation of the affected limb in primary somatosensory cortex (S1). This notion has driven treatments that aim to restore S1 representations in CRPS patients, such as sensory discrimination training and mirror therapy. However, this notion is based on both indirect and incomplete evidence obtained with imaging methods with low spatial resolution. Here, we used fMRI to characterize the S1 representation of the affected and unaffected hand in humans (of either sex) with unilateral CRPS. The cortical area, location, and geometry of the S1 representation of the CRPS hand were largely comparable with those of both the unaffected hand and healthy controls. We found no differential relation between affected versus unaffected hand map measures and clinical measures (pain severity, upper limb disability, disease duration). Thus, if any map reorganization occurs, it does not appear to be directly related to pain and disease severity. These findings compel us to reconsider the cortical mechanisms underlying CRPS and the rationale for interventions that aim to "restore" somatotopic representations to treat pain.SIGNIFICANCE STATEMENT This study shows that the spatial map of the fingers in somatosensory cortex is largely preserved in chronic complex regional pain syndrome (CRPS). These findings challenge the treatment rationale for restoring somatotopic representations in complex regional pain syndrome patients.

Detectability of cerebellar activity with magnetoencephalography and electroencephalography.

Electrophysiological signals from the cerebellum have traditionally been viewed as inaccessible to magnetoencephalography (MEG) and electroencephalography (EEG). Here, we challenge this position by investigating the ability of MEG and EEG to detect cerebellar activity using a model that employs a high-resolution tessellation of the cerebellar cortex. The tessellation was constructed from repetitive high-field (9.4T) structural magnetic resonance imaging (MRI) of an ex vivo human cerebellum. A boundary-element forward model was then used to simulate the M/EEG signals resulting from neural activity in the cerebellar cortex. Despite significant signal cancelation due to the highly convoluted cerebellar cortex, we found that the cerebellar signal was on average only 30-60% weaker than the cortical signal. We also made detailed M/EEG sensitivity maps and found that MEG and EEG have highly complementary sensitivity distributions over the cerebellar cortex. Based on previous fMRI studies combined with our M/EEG sensitivity maps, we discuss experimental paradigms that are likely to offer high M/EEG sensitivity to cerebellar activity. Taken together, these results show that cerebellar activity should be clearly detectable by current M/EEG systems with an appropriate experimental setup.

Visually-Driven Maps in Area 3b.

Sensory perception relies on the precise neuronal encoding of modality-specific environmental features in primary sensory cortices. Some studies have reported the penetration of signals from other modalities even into early sensory areas. So far, no comprehensive account of maps induced by "foreign sources" exists. We addressed this question using surface-based topographic mapping techniques applied to ultra-high resolution fMRI neuroimaging data, measured in female participants. We show that fine-grained finger maps in human primary somatosensory cortex, area 3b, are somatotopically activated not only during tactile mechanical stimulation, but also when viewing the same fingers being touched. Visually-induced maps were weak in amplitude, but overlapped with the stronger tactile maps tangential to the cortical sheet when finger touches were observed in both first- and third-person perspectives. However, visually-induced maps did not overlap tactile maps when the observed fingers were only approached by an object but not actually touched. Our data provide evidence that "foreign source maps" in early sensory cortices are present in the healthy human brain, that their arrangement is precise, and that their induction is feature-selective. The computations required to generate such specific responses suggest that counterflow (feedback) processing may be much more spatially specific than has been often assumed.SIGNIFICANCE STATEMENT Using ultra-high field fMRI, we provide empirical evidence that viewing touches activates topographically aligned single finger maps in human primary somatosensory cortical area 3b. This shows that "foreign source maps" in early sensory cortices are topographic, precise, and feature-selective in healthy human participants with intact sensory pathways.

Unraveling the spatiotemporal brain dynamics during a simulated reach-to-eat task.

The reach-to-eat task involves a sequence of action components including looking, reaching, grasping, and feeding. While cortical representations of individual action components have been mapped in human functional magnetic resonance imaging (fMRI) studies, little is known about the continuous spatiotemporal dynamics among these representations during the reach-to-eat task. In a periodic event-related fMRI experiment, subjects were scanned while they reached toward a food image, grasped the virtual food, and brought it to their mouth within each 16-s cycle. Fourier-based analysis of fMRI time series revealed periodic signals and noise distributed across the brain. Independent component analysis was used to remove periodic or aperiodic motion artifacts. Time-frequency analysis was used to analyze the temporal characteristics of periodic signals in each voxel. Circular statistics was then used to estimate mean phase angles of periodic signals and select voxels based on the distribution of phase angles. By sorting mean phase angles across regions, we were able to show the real-time spatiotemporal brain dynamics as continuous traveling waves over the cortical surface. The activation sequence consisted of approximately the following stages: (1) stimulus related activations in occipital and temporal cortices; (2) movement planning related activations in dorsal premotor and superior parietal cortices; (3) reaching related activations in primary sensorimotor cortex and supplementary motor area; (4) grasping related activations in postcentral gyrus and sulcus; (5) feeding related activations in orofacial areas. These results suggest that phase-encoded design and analysis can be used to unravel sequential activations among brain regions during a simulated reach-to-eat task.

Neural correlates of the LSD experience revealed by multimodal neuroimaging.

Lysergic acid diethylamide (LSD) is the prototypical psychedelic drug, but its effects on the human brain have never been studied before with modern neuroimaging. Here, three complementary neuroimaging techniques: arterial spin labeling (ASL), blood oxygen level-dependent (BOLD) measures, and magnetoencephalography (MEG), implemented during resting state conditions, revealed marked changes in brain activity after LSD that correlated strongly with its characteristic psychological effects. Increased visual cortex cerebral blood flow (CBF), decreased visual cortex alpha power, and a greatly expanded primary visual cortex (V1) functional connectivity profile correlated strongly with ratings of visual hallucinations, implying that intrinsic brain activity exerts greater influence on visual processing in the psychedelic state, thereby defining its hallucinatory quality. LSD's marked effects on the visual cortex did not significantly correlate with the drug's other characteristic effects on consciousness, however. Rather, decreased connectivity between the parahippocampus and retrosplenial cortex (RSC) correlated strongly with ratings of "ego-dissolution" and "altered meaning," implying the importance of this particular circuit for the maintenance of "self" or "ego" and its processing of "meaning." Strong relationships were also found between the different imaging metrics, enabling firmer inferences to be made about their functional significance. This uniquely comprehensive examination of the LSD state represents an important advance in scientific research with psychedelic drugs at a time of growing interest in their scientific and therapeutic value. The present results contribute important new insights into the characteristic hallucinatory and consciousness-altering properties of psychedelics that inform on how they can model certain pathological states and potentially treat others.

Extensive Tonotopic Mapping across Auditory Cortex Is Recapitulated by Spectrally Directed Attention and Systematically Related to Cortical Myeloarchitecture.

Auditory selective attention is vital in natural soundscapes. But it is unclear how attentional focus on the primary dimension of auditory representation-acoustic frequency-might modulate basic auditory functional topography during active listening. In contrast to visual selective attention, which is supported by motor-mediated optimization of input across saccades and pupil dilation, the primate auditory system has fewer means of differentially sampling the world. This makes spectrally-directed endogenous attention a particularly crucial aspect of auditory attention. Using a novel functional paradigm combined with quantitative MRI, we establish in male and female listeners that human frequency-band-selective attention drives activation in both myeloarchitectonically estimated auditory core, and across the majority of tonotopically mapped nonprimary auditory cortex. The attentionally driven best-frequency maps show strong concordance with sensory-driven maps in the same subjects across much of the temporal plane, with poor concordance in areas outside traditional auditory cortex. There is significantly greater activation across most of auditory cortex when best frequency is attended, versus ignored; the same regions do not show this enhancement when attending to the least-preferred frequency band. Finally, the results demonstrate that there is spatial correspondence between the degree of myelination and the strength of the tonotopic signal across a number of regions in auditory cortex. Strong frequency preferences across tonotopically mapped auditory cortex spatially correlate with R1-estimated myeloarchitecture, indicating shared functional and anatomical organization that may underlie intrinsic auditory regionalization.SIGNIFICANCE STATEMENT Perception is an active process, especially sensitive to attentional state. Listeners direct auditory attention to track a violin's melody within an ensemble performance, or to follow a voice in a crowded cafe. Although diverse pathologies reduce quality of life by impacting such spectrally directed auditory attention, its neurobiological bases are unclear. We demonstrate that human primary and nonprimary auditory cortical activation is modulated by spectrally directed attention in a manner that recapitulates its tonotopic sensory organization. Further, the graded activation profiles evoked by single-frequency bands are correlated with attentionally driven activation when these bands are presented in complex soundscapes. Finally, we observe a strong concordance in the degree of cortical myelination and the strength of tonotopic activation across several auditory cortical regions.

Microstructural parcellation of the human brain.

The human cerebral cortex is composed of a mosaic of areas thought to subserve different functions. The parcellation of the cortex into areas has a long history and has been carried out using different combinations of structural, connectional, receptotopic, and functional properties. Here we give a brief overview of the history of cortical parcellation, and explore different microstructural properties and analysis techniques that can be used to define the borders between different regions. We show that accounting for the 3D geometry of the highly folded human cortex is especially critical for accurate parcellation. We close with some thoughts on future directions and best practices for combining modalities.

Using diffusion MRI to discriminate areas of cortical grey matter.

Cortical area parcellation is a challenging problem that is often approached by combining structural imaging (e.g., quantitative T1, diffusion-based connectivity) with functional imaging (e.g., task activations, topological mapping, resting state correlations). Diffusion MRI (dMRI) has been widely adopted to analyse white matter microstructure, but scarcely used to distinguish grey matter regions because of the reduced anisotropy there. Nevertheless, differences in the texture of the cortical 'fabric' have long been mapped by histologists to distinguish cortical areas. Reliable area-specific contrast in the dMRI signal has previously been demonstrated in selected occipital and sensorimotor areas. We expand upon these findings by testing several diffusion-based feature sets in a series of classification tasks. Using Human Connectome Project (HCP) 3T datasets and a supervised learning approach, we demonstrate that diffusion MRI is sensitive to architectonic differences between a large number of different cortical areas defined in the HCP parcellation. By employing a surface-based cortical imaging pipeline, which defines diffusion features relative to local cortical surface orientation, we show that we can differentiate areas from their neighbours with higher accuracy than when using only fractional anisotropy or mean diffusivity. The results suggest that grey matter diffusion may provide a new, independent source of information for dividing up the cortex.

Learning of goal-relevant and -irrelevant complex visual sequences in human V1.

Learning and memory are supported by a network involving the medial temporal lobe and linked neocortical regions. Emerging evidence indicates that primary visual cortex (i.e., V1) may contribute to recognition memory, but this has been tested only with a single visuospatial sequence as the target memorandum. The present study used functional magnetic resonance imaging to investigate whether human V1 can support the learning of multiple, concurrent complex visual sequences involving discontinous (second-order) associations. Two peripheral, goal-irrelevant but structured sequences of orientated gratings appeared simultaneously in fixed locations of the right and left visual fields alongside a central, goal-relevant sequence that was in the focus of spatial attention. Pseudorandom sequences were introduced at multiple intervals during the presentation of the three structured visual sequences to provide an online measure of sequence-specific knowledge at each retinotopic location. We found that a network involving the precuneus and V1 was involved in learning the structured sequence presented at central fixation, whereas right V1 was modulated by repeated exposure to the concurrent structured sequence presented in the left visual field. The same result was not found in left V1. These results indicate for the first time that human V1 can support the learning of multiple concurrent sequences involving complex discontinuous inter-item associations, even peripheral sequences that are goal-irrelevant.

Spatiotemporal integration of looming visual and tactile stimuli near the face.

Real-world objects approaching or passing by an observer often generate visual, auditory, and tactile signals with different onsets and durations. Prompt detection and avoidance of an impending threat depend on precise binding of looming signals across modalities. Here we constructed a multisensory apparatus to study the spatiotemporal integration of looming visual and tactile stimuli near the face. In a psychophysical experiment, subjects assessed the subjective synchrony between a looming ball and an air puff delivered to the same side of the face with a varying temporal offset. Multisensory stimuli with similar onset times were perceived as completely out of sync and assessed with the lowest subjective synchrony index (SSI). Across subjects, the SSI peaked at an offset between 800 and 1,000 ms, where the multisensory stimuli were perceived as optimally in sync. In an fMRI experiment, tactile, visual, tactile-visual out-of-sync (TVoS), and tactile-visual in-sync (TViS) stimuli were delivered to either side of the face in randomized events. Group-average statistical responses to different stimuli were compared within each surface-based region of interest (sROI) outlined on the cortical surface. Most sROIs showed a preference for contralateral stimuli and higher responses to multisensory than unisensory stimuli. In several bilateral sROIs, particularly the human MT+ complex and V6A, responses to spatially aligned multisensory stimuli (TVoS) were further enhanced when the stimuli were in-sync (TViS), as expressed by TVoS < TViS. This study demonstrates the perceptual and neural mechanisms of multisensory integration near the face, which has potential applications in the development of multisensory entertainment systems and media.

Functional and Quantitative MRI Mapping of Somatomotor Representations of Human Supralaryngeal Vocal Tract.

Speech articulation requires precise control of and coordination between the effectors of the vocal tract (e.g., lips, tongue, soft palate, and larynx). However, it is unclear how the cortex represents movements of and contact between these effectors during speech, or how these cortical responses relate to inter-regional anatomical borders. Here, we used phase-encoded fMRI to map somatomotor representations of speech articulations. Phonetically trained participants produced speech phones, progressing from front (bilabial) to back (glottal) place of articulation. Maps of cortical myelin proxies (R1 = 1/T1) further allowed us to situate functional maps with respect to anatomical borders of motor and somatosensory regions. Across participants, we found a consistent topological map of place of articulation, spanning the central sulcus and primary motor and somatosensory areas, that moved from lateral to inferior as place of articulation progressed from front to back. Phones produced at velar and glottal places of articulation activated the inferior aspect of the central sulcus, but with considerable across-subject variability. R1 maps for a subset of participants revealed that articulator maps extended posteriorly into secondary somatosensory regions. These results show consistent topological organization of cortical representations of the vocal apparatus in the context of speech behavior.

Body Topography Parcellates Human Sensory and Motor Cortex.

The cytoarchitectonic map as proposed by Brodmann currently dominates models of human sensorimotor cortical structure, function, and plasticity. According to this model, primary motor cortex, area 4, and primary somatosensory cortex, area 3b, are homogenous areas, with the major division lying between the two. Accumulating empirical and theoretical evidence, however, has begun to question the validity of the Brodmann map for various cortical areas. Here, we combined in vivo cortical myelin mapping with functional connectivity analyses and topographic mapping techniques to reassess the validity of the Brodmann map in human primary sensorimotor cortex. We provide empirical evidence that area 4 and area 3b are not homogenous, but are subdivided into distinct cortical fields, each representing a major body part (the hand and the face). Myelin reductions at the hand-face borders are cortical layer-specific, and coincide with intrinsic functional connectivity borders as defined using large-scale resting state analyses. Our data extend the Brodmann model in human sensorimotor cortex and suggest that body parts are an important organizing principle, similar to the distinction between sensory and motor processing.

Mapping the complex topological organization of the human parietal face area.

The macaque monkey ventral intraparietal area (VIP) contains neurons with aligned visual-tactile receptive fields anchored to the face and upper body. Our previous fMRI studies using standard head coils found a human parietal face area (VIP+ complex; putative macaque VIP homologue) containing superimposed topological maps of the face and near-face visual space. Here, we construct high signal-to-noise surface coils and used phase-encoded air puffs and looming stimuli to map topological organization of the parietal face area at higher resolution. This area is consistently identified as a region extending between the superior postcentral sulcus and the upper bank of the anterior intraparietal sulcus (IPS), avoiding the fundus of IPS. Using smaller voxel sizes, our surface coils picked up strong fMRI signals in response to tactile and visual stimuli. By analyzing tactile and visual maps in our current and previous studies, we constructed a set of topological models illustrating commonalities and differences in map organization across subjects. The most consistent topological feature of the VIP+ complex is a central-anterior upper face (and upper visual field) representation adjoined by lower face (and lower visual field) representations ventrally (laterally) and/or dorsally (medially), potentially forming two subdivisions VIPv (ventral) and VIPd (dorsal). The lower visual field representations typically extend laterally into the anterior IPS to adjoin human area AIP, and medially to overlap with the parietal body areas at the superior parietal ridge. Significant individual variations are then illustrated to provide an accurate and comprehensive view of the topological organization of the parietal face area.

Validation of periodic fMRI signals in response to wearable tactile stimulation.

To map cortical representations of the body, we recently developed a wearable technology for automatic tactile stimulation in human functional magnetic resonance imaging (fMRI) experiments. In a two-condition block design experiment, air puffs were delivered to the face and hands periodically. Surface-based regions of interest (S-ROIs) were initially identified by thresholding a linear statistical measure of signal-to-noise ratio of periodic response. Across subjects, S-ROIs were found in the frontal, primary sensorimotor, posterior parietal, insular, temporal, cingulate, and occipital cortices. To validate and differentiate these S-ROIs, we develop a measure of temporal stability of response based on the assumption that a periodic stimulation evokes stable (low-variance) periodic fMRI signals throughout the entire scan. Toward this end, we apply time-frequency analysis to fMRI time series and use circular statistics to characterize the distribution of phase angles for data selection. We then assess the temporal variability of a periodic signal by measuring the path length of its trajectory in the complex plane. Both within and outside the primary sensorimotor cortex, S-ROIs with high temporal variability and deviant phase angles are rejected. A surface-based probabilistic group-average map is constructed for spatial screening of S-ROIs with low to moderate temporal variability in non-sensorimotor regions. Areas commonly activated across subjects are also summarized in the group-average map. In summary, this study demonstrates that analyzing temporal characteristics of the entire fMRI time series is essential for second-level selection and interpretation of S-ROIs initially defined by an overall linear statistical measure.