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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.

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.

A fundamental tenet of neuroscience is that cortical functional differentiation is related to the cross-areal differences in cyto-, receptor-, and myeloarchitectonics that are observed in ex-vivo preparations. An ongoing challenge is to create noninvasive magnetic resonance (MR) imaging techniques that offer sufficient resolution, tissue contrast, accuracy and precision to allow for characterization of cortical architecture over an entire living human brain. One exciting development is the advent of fast, high-resolution quantitative mapping of basic MR parameters that reflect cortical myeloarchitecture. Here, we outline some of the theoretical and technical advances underlying this technique, particularly in terms of measuring and correcting for transmit and receive radio frequency field inhomogeneities. We also discuss new directions in analytic techniques, including higher resolution reconstructions of the cortical surface. We then discuss two recent applications of this technique. The first compares individual and group myelin maps to functional retinotopic maps in the same individuals, demonstrating a close relationship between functionally and myeloarchitectonically defined areal boundaries (as well as revealing an interesting disparity in a highly studied visual area). The second combines tonotopic and myeloarchitectonic mapping to localize primary auditory areas in individual healthy adults, using a similar strategy as combined electrophysiological and post-mortem myeloarchitectonic studies in non-human primates.

In macaque monkeys, V6A is a visuomotor area located in the anterior bank of the POs, dorsal and anterior to retinotopically-organized extrastriate area V6 (Galletti et al 1996). Unlike V6, V6A represents both contra- and ipsilateral visual fields and is broadly retinotopically organized (Galletti et al 1999b). The contralateral lower visual field is over-represented in V6A. The central 20°-30° of the visual field are mainly represented dorsally (V6Ad) and the periphery ventrally (V6Av), at the border with V6. Both sectors of area V6A contain arm movement-related cells, active during spatially-directed reaching movements (Gamberini et al., 2011).\ud \ud In humans, we previously mapped the retinotopic organization of area V6 (Pitzalis et al., 2006). Here, using phase-encoded fMRI, cortical surface-based analysis and wide-field retinotopic mapping, we define a new cortical region that borders V6 anteriorly and shows a clear over-representation of the contralateral lower visual field and of the periphery. As with macaque V6A, the eccentricity increases moving ventrally within the area. The new region contains a non-mirror-image representation of the visual field. Functional mapping reveals that, as in macaque V6A, the new region, but not the nearby area V6, responds during finger pointing and reaching movements. Based on similarity in position, retinotopic properties, functional organization and relationship with the neighbouring extrastriate visual areas, we propose that the new cortical region is the human homologue of macaque area V6A.

Temporal conjunctions ‘before’ and ‘after’ give us freedom to describe a series of events in different orders. Previous studies suggested ‘before’ sentences in which events were expresses in an order inconsistent with their actual order of occurrence need additional computations, i.e. reversing the order of event mention to reach the actual order of event occurrence. This study found the additional computations may be supported by a neural network connecting the caudate nucleus with the medial prefrontal cortex, middle frontal gyrus (MFG), precuneus and occipital cortex. The connectivity in this network was strongly enhanced for ‘before’ than ‘after’ sentences. Meanwhile, another network was observed to support the memory retrieval, connecting the hippocampus with the MFG via the orbital inferior frontal gyrus and temporal pole. The connectivity pattern of this network was not different between conditions. With the common node MFG, these two networks may communicate in working memory to work together.

We used functional magnetic resonance imaging (fMRI) to map the cortical representations of executed reaching, observed reaching, and imagined reaching in humans. Whereas previous studies have mostly examined hand actions related to grasping, hand–object interactions, or local finger movements, here we were interested in reaching only (i.e. the transport phase of the hand to a particular location in space), without grasping. We hypothesized that mirror neuron areas specific to reaching-related representations would be active in all three conditions. An overlap between executed, observed, and imagined reaching activations was found in dorsal premotor cortex as well as in the superior parietal lobe and the intraparietal sulcus, in accord with our hypothesis. Activations for observed reaching were more dorsal than activations typically reported in the literature for observation of hand–object interactions (grasping). Our results suggest that the mirror neuron system is specific to the type of hand action performed, and that these fronto-parietal activations are a putative human homologue of the neural circuits underlying reaching in macaques. The parietal activations reported here for executed, imagined, and observed reaching are also consistent with previous functional imaging studies on planned reaching and delayed pointing movements, and extend the proposed localization of human reach-related brain areas to observation as well as imagery of reaching.

Somatotopic mapping of human body surface using fMRI is challenging. First, it is difficult to deliver tactile stimuli in the scanner. Second, multiple stimulators are often required to cover enough area of the complex-shaped body surface, such as the face. In this study, a computer-controlled pneumatic system was constructed to automatically deliver air puffs to 12 locations on the body surface through an MR-compatible manifold (Dodecapus) mounted on a head coil inside the scanner bore. The timing of each air-puff channel is completely programmable and this allows systematic and precise stimulation on multiple locations on the body surface during functional scans. Three two-condition block-design "Localizer" paradigms were employed to localize the cortical representations of the face, lips, and fingers, respectively. Three "Phase-encoded" paradigms were employed to map the detailed somatotopic organizations of the face, lips, and fingers following each "Localizer" paradigm. Multiple somatotopic representations of the face, lips, and fingers were localized and mapped in primary motor cortex (MI), ventral premotor cortex (PMv), polysensory zone (PZ), primary (SI) and secondary (SII) somatosensory cortex, parietal ventral area (PV) and 7b, as well as anterior and ventral intraparietal areas (AIP and VIP). The Dodecapus system is portable, easy to setup, generates no radio frequency interference, and can also be used for EEG and MEG experiments. This system could be useful for non-invasive somatotopic mapping in both basic and clinical studies.

There is evidence of both crossmodal and intermodal plasticity in the deaf brain. Here, we investigated whether sub-cortical plasticity, specifically of the thalamus, contributed to this reorganisation. We contrasted diffusion weighted magnetic resonance imaging data from 13 congenitally deaf and 13 hearing participants, all of whom had learnt British Sign Language after 10 years of age. Connectivity based segmentation of the thalamus revealed changes to mean and radial diffusivity in occipital and frontal regions, which may be linked to enhanced peripheral visual acuity, and differences in how visual attention is deployed in the deaf group. Using probabilistic tractography, tracts were traced between the thalamus and its cortical targets, and microstructural measurements were extracted from these tracts. Group differences were found in microstructural measurements of occipital, frontal, somatosensory, motor and parietal thalamo-cortical tracts. Our findings suggest that there is sub-cortical plasticity in the deaf brain, and that white matter alterations can be found throughout the deaf brain, rather than being restricted to, or focussed in the auditory cortex., Highlights • Diffusion weighted MRI data was acquired from congenitally deaf and hearing groups. • Mean and radial diffusivity is altered in the thalamus of congenitally deaf people. • Diffusivity in white matter thalamo-cortical radiations is altered in deafness.

Several properties of the cerebral cortex, including its columnar and laminar organization, as well as the topographic organization of cortical areas, can only be properly understood in the context of the intrinsic two-dimensional structure of the cortical surface. In order to study such cortical properties in humans, it is necessary to obtain an accurate and explicit representation of the cortical surface in individual subjects. Here we describe a set of automated procedures for obtaining accurate reconstructions of the cortical surface, which have been applied to data from more than 100 subjects, requiring little or no manual intervention. Automated routines for unfolding and flattening the cortical surface are described in a companion paper. These procedures allow for the routine use of cortical surface-based analysis and visualization methods in functional brain imaging. r 1999 Academic Press

We explored how developing neural artifact and animal representations in the dorsal and ventral stream play a role in children's increasingly more proficient interactions with objects. In thirty-three 6- to 10-year-old children and 11 adults, we used fMRI to track the development of (1) the cortical category preference for tools compared to animals and (2) the response to complex objects (as compared to scrambled objects) during a passive viewing task. In addition, we related a cognitive skill that improved substantially from age 6 to 10, namely the ability to recognize tools from unusual viewpoints, to the development of cortical object processing. In multiple complementary analyses we showed that those children who were better at recognizing tools from unusual viewpoints outside the scanner showed a reduced cortical response to tools and animals when viewed inside the scanner, bilaterally in intraparietal and inferotemporal cortex. In contrast, the cortical preference for tools in the dorsal and ventral visual stream did not predict object recognition performance, and was organized in an adult-like manner at six. While cortical tool preference did not change with age, the findings suggest that animal-preferring regions in the ventral visual stream may develop later, concordant with previous reports of a protracted development in similar regions for faces. We thus conclude that intraparietal and inferotemporal cortical networks that support aspects of object processing irrespective of tool or animal category, continue to develop during the school-age years and contribute to the development of object recognition skills during this period.

Cortical surface-based analysis of fMRI data has proven to be a useful method with several advantages over 3-dimensional volumetric analyses. Many of the statistical methods used in 3D analyses can be adapted for use with surface-based analyses. Operating within the framework of the FreeSurfer software package, we have implemented a surface-based version of the cluster size exclusion method used for multiple comparisons correction. Furthermore, we have a developed a new method for generating regions of interest on the cortical surface using a sliding threshold of cluster exclusion followed by cluster growth. Cluster size limits for multiple probability thresholds were estimated using random field theory and validated with Monte Carlo simulation. A prerequisite of RFT or cluster size simulation is an estimate of the smoothness of the data. In order to estimate the intrinsic smoothness of group analysis statistics, independent of true activations, we conducted a group analysis of simulated noise data sets. Because smoothing on a cortical surface mesh is typically implemented using an iterative method, rather than directly applying a Gaussian blurring kernel, it is also necessary to determine the width of the equivalent Gaussian blurring kernel as a function of smoothing steps. Iterative smoothing has previously been modeled as continuous heat diffusion, providing a theoretical basis for predicting the equivalent kernel width, but the predictions of the model were not empirically tested. We generated an empirical heat diffusion kernel width function by performing surface-based smoothing simulations and found a large disparity between the expected and actual kernel widths.

A recent study from our laboratory demonstrated that parietal cortex contains a map of visual space related to saccades and spatial attention and identified this area as the likely human homologue of the lateral intraparietal (LIP). A human homologue for the parietal reach region (PRR), thought to preferentially encode planned hand movements, has also been recently proposed. Both of these areas, originally identified in the macaque monkey, have been shown to encode space with eye-centered coordinates. Functional magnetic resonance imaging (fMRI) of humans was used to test the hypothesis that the putative human PRR contains a retinotopic map recruited by finger pointing but not saccades and to test more generally for differences in the visuospatial maps recruited by pointing and saccades. We identified multiple maps in both posterior parietal cortex and superior frontal cortex recruited for eye and hand movements, including maps not observed in previous mapping studies. Pointing and saccade maps were generally consistent within single subjects. We have developed new group analysis methods for phase-encoded data, which revealed subtle differences between pointing and saccades, including hemispheric asymmetries, but we did not find evidence of pointing-specific maps of visual space.

Though the function of prefrontal cortex has been extensively investigated, little is known about the internal organization of individual prefrontal areas. Functional magnetic resonance imaging was used to show that some frontal and prefrontal cortical areas represent visual space in orderly, reproducible, topographic maps. The map-containing areas partly overlap dorsolateral prefrontal areas engaged by working memory tasks. These maps may be useful for attending to task-relevant objects at various spatial locations, an aspect of the executive control of attention.

Automatic Reconstruction of the Cortical Surface To apply surface-based techniques routinely, the cortical surface reconstruction process must be automated. Several years ago, a method was devised for recovering the complete cortical surface of each hemisphere from high resolution MRI images in the context of improved solutions to the inverse problem for EEG and MEG (2). Since then, these programs have been improved and extended to incorporate functional MRI imaging data (3) (see also http://cogsci.ucsd.edu/~sereno).