Two-Dimensional Population Receptive Field Mapping of Human Primary Somatosensory Cortex.

Functional magnetic resonance imaging can provide detailed maps of how sensory space is mapped in the human brain. Here, we use a novel 16 stimulator setup (a 4 × 4 grid) to measure two-dimensional sensory maps of between and within-digit (D2-D4) space using high spatial-resolution (1.25 mm isotropic) imaging at 7 Tesla together with population receptive field (pRF) mapping in 10 participants. Using a 2D Gaussian pRF model, we capture maps of the coverage of digits D2-D5 across Brodmann areas and estimate pRF size and shape. In addition, we compare results to previous studies that used fewer stimulators by constraining pRF models to a 1D Gaussian Between Digit or 1D Gaussian Within Digit model. We show that pRFs across somatosensory areas tend to have a strong preference to cover the within-digit axis. We show an increase in pRF size moving from D2-D5. We quantify pRF shapes in Brodmann area (BA) 3b, 3a, 1, 2 and show differences in pRF size in Brodmann areas 3a-2, with larger estimates for BA2. Generally, the 2D Gaussian pRF model better represents pRF coverage maps generated by our data, which itself is produced from a 2D stimulation grid.

A probabilistic atlas of finger dominance in the primary somatosensory cortex.

With the advent of ultra-high field (7T), high spatial resolution functional MRI (fMRI) has allowed the differentiation of the cortical representations of each of the digits at an individual-subject level in human primary somatosensory cortex (S1). Here we generate a probabilistic atlas of the contralateral SI representations of the digits of both the left and right hand in a group of 22 right-handed individuals. The atlas is generated in both volume and surface standardised spaces from somatotopic maps obtained by delivering vibrotactile stimulation to each distal phalangeal digit using a travelling wave paradigm. Metrics quantify the likelihood of a given position being assigned to a digit (full probability map) and the most probable digit for a given spatial location (maximum probability map). The atlas is validated using a leave-one-out cross validation procedure. Anatomical variance across the somatotopic map is also assessed to investigate whether the functional variability across subjects is coupled to structural differences. This probabilistic atlas quantifies the variability in digit representations in healthy subjects, finding some quantifiable separability between digits 2, 3 and 4, a complex overlapping relationship between digits 1 and 2, and little agreement of digit 5 across subjects. The atlas and constituent subject maps are available online for use as a reference in future neuroimaging studies.

Exploring structure and function of sensory cortex with 7T MRI.

In this paper, we present an overview of 7T magnetic resonance imaging (MRI) studies of the detailed function and anatomy of sensory areas of the human brain. We discuss the motivation for the studies, with particular emphasis on increasing the spatial resolution of functional MRI (fMRI) using reduced field-of-view (FOV) data acquisitions. MRI at ultra-high-field (UHF) - defined here as 7T and above - has several advantages over lower field strengths. The intrinsic signal-to-noise ratio (SNR) of images is higher at UHF, and coupled with the increased blood-oxygen-level-dependent (BOLD) signal change, this results in increased BOLD contrast-to-noise ratio (CNR), which can be exploited to improve spatial resolution or detect weaker signals. Additionally, the BOLD signal from the intra-vascular (IV) compartment is relatively diminished compared to lower field strengths. Together, these properties make 7T functional MRI an attractive proposition for high spatial specificity measures. But with the advantages come some challenges. For example, increased vulnerability to susceptibility-induced geometric distortions and signal loss in EPI acquisitions tend to be much larger. Some of these technical issues can be addressed with currently available tools and will be discussed. We highlight the key methodological considerations for high resolution functional and structural imaging at 7 T. We then present recent data using the high spatial resolution available at UHF in studies of the visual and somatosensory cortex to highlight promising developments in this area.

7 tesla FMRI reveals systematic functional organization for binocular disparity in dorsal visual cortex.

The binocular disparity between the views of the world registered by the left and right eyes provides a powerful signal about the depth structure of the environment. Despite increasing knowledge of the cortical areas that process disparity from animal models, comparatively little is known about the local architecture of stereoscopic processing in the human brain. Here, we take advantage of the high spatial specificity and image contrast offered by 7 tesla fMRI to test for systematic organization of disparity representations in the human brain. Participants viewed random dot stereogram stimuli depicting different depth positions while we recorded fMRI responses from dorsomedial visual cortex. We repeated measurements across three separate imaging sessions. Using a series of computational modeling approaches, we report three main advances in understanding disparity organization in the human brain. First, we show that disparity preferences are clustered and that this organization persists across imaging sessions, particularly in area V3A. Second, we observe differences between the local distribution of voxel responses in early and dorsomedial visual areas, suggesting different cortical organization. Third, using modeling of voxel responses, we show that higher dorsal areas (V3A, V3B/KO) have properties that are characteristic of human depth judgments: a simple model that uses tuning parameters estimated from fMRI data captures known variations in human psychophysical performance. Together, these findings indicate that human dorsal visual cortex contains selective cortical structures for disparity that may support the neural computations that underlie depth perception.

Do perceptual biases emerge early or late in visual processing? Decision-biases in motion perception.

Visual perception is strongly influenced by contextual information. A good example is reference repulsion, where subjective reports about the direction of motion of a stimulus are significantly biased by the presence of an explicit reference. These perceptual biases could arise early, during sensory encoding, or alternatively, they may reflect decision-related processes occurring relatively late in the task sequence. To separate these two competing possibilities, we asked (human) subjects to perform a fine motion-discrimination task and then estimate the direction of motion in the presence or absence of an oriented reference line. When subjects performed the discrimination task with the reference, but subsequently estimated motion direction in its absence, direction estimates were unbiased. However, when subjects viewed the same stimuli but performed the estimation task only, with the orientation of the reference line jittered on every trial, the directions estimated by subjects were biased and yoked to the orientation of the shifted reference line. These results show that judgements made relative to a reference are subject to late, decision-related biases A model in which information about motion is integrated with that of an explicit reference cue, resulting in a late, decision-related re-weighting of the sensory representation, can account for these results.

Assessing the spatial precision of SE and GE-BOLD contrast at 7 Tesla.

Spin echo (SE) EPI offers an alternative to standard gradient echo (GE) EPI for functional MRI. SE-EPI offers improved spatial specificity, since signal changes originate from the microvasculature, but its lower functional sensitivity has limited the usage of this sequence in fMRI experiments. Differential fMRI paradigms, in which two closely matched stimulus conditions are used, can suppress the contribution from veins, thus also offering improved spatial specificity compared to conventional block or event-related designs with long "rest" periods. In this study, we employed a differential fMRI paradigm to stimulate bands of primary visual cortex with pre-defined widths by using visual stimuli comprised of complementary rings of contrast-reversing checkerboard patterns (8 Hz). This paradigm was used to investigate the spatial specificity of GE and SE-BOLD contrast at 7T. Results show that the contrast-to-noise ratio (CNR) is larger for GE-EPI data than for the SE-EPI data for band widths in the range 1.7-6.6 mm, however as the width of the band decreases the CNR for GE and SE sequences converges. These results suggest that when using a differential mapping paradigm, GE-BOLD contrast is better for studying functional features that are larger than ~1.5 mm in size.

Decoding working memory of stimulus contrast in early visual cortex.

Most studies of the early stages of visual analysis (V1-V3) have focused on the properties of neurons that support processing of elemental features of a visual stimulus or scene, such as local contrast, orientation, or direction of motion. Recent evidence from electrophysiology and neuroimaging studies, however, suggests that early visual cortex may also play a role in retaining stimulus representations in memory for short periods. For example, fMRI responses obtained during the delay period between two presentations of an oriented visual stimulus can be used to decode the remembered stimulus orientation with multivariate pattern analysis. Here, we investigated whether orientation is a special case or if this phenomenon generalizes to working memory traces of other visual features. We found that multivariate classification of fMRI signals from human visual cortex could be used to decode the contrast of a perceived stimulus even when the mean response changes were accounted for, suggesting some consistent spatial signal for contrast in these areas. Strikingly, we found that fMRI responses also supported decoding of contrast when the stimulus had to be remembered. Furthermore, classification generalized from perceived to remembered stimuli and vice versa, implying that the corresponding pattern of responses in early visual cortex were highly consistent. In additional analyses, we show that stimulus decoding here is driven by biases depending on stimulus eccentricity. This places important constraints on the interpretation for decoding stimulus properties for which cortical processing is known to vary with eccentricity, such as contrast, color, spatial frequency, and temporal frequency.

Regional structural differences across functionally parcellated Brodmann areas of human primary somatosensory cortex.

Ultra-high-field (UHF) MRI is ideally suited for structural and functional imaging of the brain. High-resolution structural MRI can be used to map the anatomical boundaries between functional domains of the brain by identifying changes related to the pattern of myelination within cortical gray matter, opening up the possibility to study the relationship between functional domains and underlying structure in vivo. In a recent study, we demonstrated the correspondence between functional (based on retinotopic mapping) and structural (based on changes in T2(⁎)-weighted images linked to myelination) parcellations of the primary visual cortex (V1) in vivo at 7T (Sanchez-Panchuelo et al., 2012b). Here, we take advantage of the improved BOLD CNR and high spatial resolution achievable at 7T to study regional structural variations across the functionally defined areas within the primary somatosensory cortex (S1) in individual subjects. Using a traveling wave fMRI paradigm to map the internal somatotopic representation of the index, middle, and ring fingers in S1, we were able to identify multiple map reversals at the tip and base, corresponding to the boundaries between Brodmann areas 3a, 3b, 1 and 2. Based on high resolution structural MRI data acquired in the same subjects, we inspected these functionally-parcellated Brodmann areas for differences in cortical thickness and MR contrast measures (magnetization transfer ratio (MTR) and signal intensity in phase sensitive inversion recovery (PSIR) images) that are sensitive to myelination. Consistent area-related differences in cortical thickness and MTR/PSIR measurements were found across subjects. However these measures did not have sufficient sensitivity to allow definition of areal boundaries.

Event-related fMRI at 7T reveals overlapping cortical representations for adjacent fingertips in S1 of individual subjects.

Recent fMRI studies of the human primary somatosensory cortex have been able to differentiate the cortical representations of different fingertips at a single-subject level. These studies did not, however, investigate the expected overlap in cortical activation due to the stimulation of different fingers. Here, we used an event-related design in six subjects at 7 Tesla to explore the overlap in cortical responses elicited in S1 by vibrotactile stimulation of the five fingertips. We found that all parts of S1 show some degree of spatial overlap between the cortical representations of adjacent or even nonadjacent fingertips. In S1, the posterior bank of the central sulcus showed less overlap than regions in the post-central gyrus, which responded to up to five fingertips. The functional properties of these two areas are consistent with the known layout of cytoarchitectonically defined subareas, and we speculate that they correspond to subarea 3b (S1 proper) and subarea 1, respectively. In contrast with previous fMRI studies, however, we did not observe discrete activation clusters that could unequivocally be attributed to different subareas of S1. Venous maps based on T2*-weighted structural images suggest that the observed overlap is not driven by extra-vascular contributions from large veins.

Within-digit functional parcellation of Brodmann areas of the human primary somatosensory cortex using functional magnetic resonance imaging at 7 tesla.

The primary somatosensory cortex (S1) can be subdivided cytoarchitectonically into four distinct Brodmann areas (3a, 3b, 1, and 2), but these areas have never been successfully delineated in vivo in single human subjects. Here, we demonstrate the functional parcellation of four areas of S1 in individual human subjects based on high-resolution functional MRI measurements made at 7 T using vibrotactile stimulation. By stimulating four sites along the length of the index finger, we were able to identify and locate map reversals of the base to tip representation of the index finger in S1. We suggest that these reversals correspond to the areal borders between the mirrored representations in the four Brodmann areas, as predicted from electrophysiology measurements in nonhuman primates. In all subjects, maps were highly reproducible across scanning sessions and stable over weeks. In four of the six subjects scanned, four, mirrored, within-finger somatotopic maps defining the extent of the Brodmann areas could be directly observed on the cortical surface. In addition, by using multivariate classification analysis, the location of stimulation on the index finger (four distinct sites) could be decoded with a mean accuracy of 65% across subjects. Our measurements thus show that within-finger topography is present at the millimeter scale in the cortex and is highly reproducible. The ability to identify functional areas of S1 in vivo in individual subjects will provide a framework for investigating more complex aspects of tactile representation in S1.

Single-subject fMRI mapping at 7 T of the representation of fingertips in S1: a comparison of event-related and phase-encoding designs.

A desirable goal of functional MRI (fMRI), both clinically and for basic research, is to produce detailed maps of cortical function in individual subjects. Single-subject mapping of the somatotopic hand representation in the human primary somatosensory cortex (S1) has been performed using both phase-encoding and block/event-related designs. Here, we review the theoretical strengths and limits of each method and empirically compare high-resolution (1.5 mm isotropic) somatotopic maps obtained using fMRI at ultrahigh magnetic field (7 T) with phase-encoding and event-related designs in six subjects in response to vibrotactile stimulation of the five fingertips. Results show that the phase-encoding design is more efficient than the event-related design for mapping fingertip-specific responses and in particular allows us to describe a new additional somatotopic representation of fingertips on the precentral gyrus. However, with sufficient data, both designs yield very similar fingertip-specific maps in S1, which confirms that the assumption of local representational continuity underlying phase-encoding designs is largely valid at the level of the fingertips in S1. In addition, it is shown that the event-related design allows the mapping of overlapping cortical representations that are difficult to estimate using the phase-encoding design. The event-related data show a complex pattern of overlapping cortical representations for different fingertips within S1 and demonstrate that regions of S1 responding to several adjacent fingertips can incorrectly be identified as responding preferentially to one fingertip in the phase-encoding data.

The effects of simulated hemianopia on eye movements during text reading.

Vision loss is a common, devastating complication of cerebral strokes. In some cases the complete contra-lesional visual field is affected, leading to problems with routine tasks and, notably, the ability to read. Although visual information crucial for reading is imaged on the foveal region, readers often extract useful parafoveal information from the next word or two in the text. In hemianopic field loss, parafoveal processing is compromised, shrinking the visual span and resulting in slower reading speeds. Recent approaches to rehabilitation using perceptual training have been able to demonstrate some recovery of useful visual capacity. As gains in visual sensitivity were most pronounced at the border of the scotoma, it may be possible to use training to restore some of the lost visual span for reading. As restitutive approaches often involve prolonged training sessions, it would be beneficial to know how much recovery is required to restore reading ability. To address this issue, we employed a gaze-contingent paradigm using a low-pass filter to blur one side of the text, functionally simulating a visual field defect. The degree of blurring acts as a proxy for visual function recovery that could arise from restitutive strategies, and allows us to evaluate and quantify the degree of visual recovery required to support normal reading fluency in patients. Because reading ability changes with age, we recruited a group of younger participants, and another with older participants who are closer in age to risk groups for ischaemic strokes. Our results show that changes in patterns of eye movement observed in hemianopic loss can be captured using this simulated reading environment. This opens up the possibility of using participants with normal visual function to help identify the most promising strategies for ameliorating hemianopic loss, before translation to patient groups.

Correspondence of human visual areas identified using functional and anatomical MRI in vivo at 7 T.

PURPOSE: To study the correspondence of anatomically and functionally defined visual areas (primary visual cortex, V1, and motion selective area V5/human MT+) by using structural magnetic resonance imaging (MRI) and functional MRI (fMRI) in vivo at 7 T. MATERIALS AND METHODS: Four subjects participated in this study. High-resolution (≈0.4 mm isotropic) anatomical MRI was used to identify cortical regions based on their distinct cortical lamination. The optimal contrast for identifying heavily myelinated layers within gray matter was quantitatively assessed by comparing T(1)-weighted magnetization-prepared rapid gradient echo (MPRAGE) and T(2)*-weighted, 3D fast-low angle shot (FLASH) imaging. Retinotopic mapping was performed using GE-based fMRI at 1.5 mm isotropic resolution to identify functional areas. RESULTS: T(2)*-weighted FLASH imaging was found to provide a significantly higher contrast-to-noise ratio, allowing visualization of the stria of Gennari in every slice of a volume covering the occipital cortex in each of the four subjects in this study. The independently derived boundary of V1, identified in the same subjects using retinotopic mapping by fMRI, closely matched the border of anatomically defined striate cortex in the human brain. Evidence of banding was also found within the functionally defined V5 area; however, we did not find a good correlation of this area, or the functionally identified subregion (MT), with the banded area. CONCLUSION: High-resolution T(2)*-weighted images acquired at 7 T can be used to identify myelinated bands within cortical gray matter in reasonable measurement times. Regions where a myelinated band was identified show a high degree of overlap with the functionally defined V1 area.

fMRI evidence that hyper-caricatured faces activate object-selective cortex.

Many brain imaging studies have looked at the cortical responses to object categories and faces. A popular way to manipulate face stimuli is by using a "face space," a high dimensional representation of individual face images, with the average face located at the origin. However, how the brain responds to faces that deviate substantially from average has not been much explored. Increasing the distance from the average (leading to increased caricaturing) could increase neural responses in face-selective regions, an idea supported by results from non-human primates. Here, we used a face space based on principal component analysis (PCA) to generate faces ranging from average to heavily caricatured. Using functional magnetic resonance imaging (fMRI), we first independently defined face-, object- and scene-selective areas with a localiser scan and then measured responses to parametrically caricatured faces. We also included conditions in which the images of faces were inverted. Interestingly in the right fusiform face area (FFA), we found that the patterns of fMRI response were more consistent as caricaturing increased. However, we found no consistent effect of either caricature level or facial inversion on the average fMRI response in the FFA or face-selective regions more broadly. In contrast, object-selective regions showed an increase in both the consistency of response pattern and the average fMRI response with increasing caricature level. This shows that caricatured faces recruit processing from regions typically defined as object-selective, possibly through enhancing low-level properties that are characteristic of objects.

Fast Event-Related Mapping of Population Fingertip Tuning Properties in Human Sensorimotor Cortex at 7T.

fMRI studies that investigate somatotopic tactile representations in the human cortex typically use either block or phase-encoded stimulation designs. Event-related (ER) designs allow for more flexible and unpredictable stimulation sequences than the other methods, but they are less efficient. Here, we compared an efficiency-optimized fast ER design (2.8-s average intertrial interval; ITI) to a conventional slow ER design (8-s average ITI) for mapping voxelwise fingertip tactile tuning properties in the sensorimotor cortex of six participants at 7 Tesla. The fast ER design yielded more reliable responses compared with the slow ER design, but with otherwise similar tuning properties. Concatenating the fast and slow ER data, we demonstrate in each individual brain the existence of two separate somatotopically-organized tactile representations of the fingertips, one in the primary somatosensory cortex (S1) on the postcentral gyrus, and the other shared across the motor and premotor cortices on the precentral gyrus. In both S1 and motor representations, fingertip selectivity decreased progressively, from narrowly-tuned Brodmann area (BA) 3b and BA4a, respectively, toward associative parietal and frontal regions that responded equally to all fingertips, suggesting increasing information integration along these two pathways. In addition, fingertip selectivity in S1 decreased from the cortical representation of the thumb to that of the pinky.

Linking Multi-Modal MRI to Clinical Measures of Visual Field Loss After Stroke.

Loss of vision across large parts of the visual field is a common and devastating complication of cerebral strokes. In the clinic, this loss is quantified by measuring the sensitivity threshold across the field of vision using static perimetry. These methods rely on the ability of the patient to report the presence of lights in particular locations. While perimetry provides important information about the intactness of the visual field, the approach has some shortcomings. For example, it cannot distinguish where in the visual pathway the key processing deficit is located. In contrast, brain imaging can provide important information about anatomy, connectivity, and function of the visual pathway following stroke. In particular, functional magnetic resonance imaging (fMRI) and analysis of population receptive fields (pRF) can reveal mismatches between clinical perimetry and maps of cortical areas that still respond to visual stimuli after stroke. Here, we demonstrate how information from different brain imaging modalities-visual field maps derived from fMRI, lesion definitions from anatomical scans, and white matter tracts from diffusion weighted MRI data-provides a more complete picture of vision loss. For any given location in the visual field, the combination of anatomical and functional information can help identify whether vision loss is due to absence of gray matter tissue or likely due to white matter disconnection from other cortical areas. We present a combined imaging acquisition and visual stimulus protocol, together with a description of the analysis methodology, and apply it to datasets from four stroke survivors with homonymous field loss (two with hemianopia, two with quadrantanopia). For researchers trying to understand recovery of vision after stroke and clinicians seeking to stratify patients into different treatment pathways, this approach combines multiple, convergent sources of data to characterize the extent of the stroke damage. We show that such an approach gives a more comprehensive measure of residual visual capacity-in two particular respects: which locations in the visual field should be targeted and what kind of visual attributes are most suited for rehabilitation.

A comparison of fMRI adaptation and multivariate pattern classification analysis in visual cortex.

Functional magnetic resonance imaging (fMRI) has become a ubiquitous tool in cognitive neuroscience. The technique allows noninvasive measurements of cortical responses in the human brain, but only on the millimeter scale. Because a typical voxel contains many thousands of neurons with varied properties, establishing the selectivity of their responses directly is impossible. In recent years, two methods using fMRI aimed at studying the selectivity of neuronal populations on a 'subvoxel' scale have been heavily used. The first technique, fMRI adaptation, relies on the observation that the blood oxygen level-dependent (BOLD) response in a given voxel is reduced after prolonged presentation of a stimulus, and that this reduction is selective to the characteristics of the repeated stimuli (adapters). The second technique, multivariate pattern analysis (MVPA), makes use of multivariate statistics to recover small biases in individual voxels in their responses to different stimuli. It is thought that these biases arise due to the uneven distribution of neurons (with different properties) sampled by the many voxels in the imaged volume. These two techniques have not been compared explicitly, however, and little is known about their relative sensitivities. Here, we compared fMRI results from orientation-specific visual adaptation and orientation-classification by MVPA, using optimized experimental designs for each, and found that the multivariate pattern classification approach was more sensitive to small differences in stimulus orientation than the adaptation paradigm. Estimates of orientation selectivity obtained with the two methods were, however, very highly correlated across visual areas.

Differential roles for frontal eye fields (FEFs) and intraparietal sulcus (IPS) in visual working memory and visual attention.

Cortical activity was measured with functional magnetic resonance imaging to probe the involvement of the superior precentral sulcus (including putative human frontal eye fields, FEFs) and the intraparietal sulcus (IPS) in visual short-term memory and visual attention. In two experimental tasks, human subjects viewed two visual stimuli separated by a variable delay period. The tasks placed differential demands on short-term memory and attention, but the stimuli were visually identical until after the delay period. An earlier study (S. Offen, D. Schluppeck, & D. J. Heeger, 2009) had found a dissociation in early visual cortex that suggested different computational mechanisms underlying the two processes. In contrast, the results reported here show that the patterns of activation in prefrontal and parietal cortex were different from one another but were similar for the two tasks. In particular, the FEF showed evidence for sustained delay period activity for both the working memory and the attention task, while the IPS did not show evidence for sustained delay period activity for either task. The results imply differential roles for the FEF and IPS in these tasks; the results also suggest that feedback of sustained activity from frontal cortex to visual cortex might be gated by task demands.

Temporal characteristics of global motion processing revealed by transcranial magnetic stimulation.

The ability to detect the motion of objects is critical to survival, and understanding the cortical mechanisms involved in this process remains a key challenge in sensory neuroscience. A relatively new approach to this problem is to temporarily disrupt processing at specific cortical sites and measure the behavioural consequences. Several previous studies have shown that transcranial magnetic stimulation (TMS) of human visual area V5/MT disrupts global motion perception, but reports vary widely in the timescale of this effect. To resolve this issue we employed psychophysical techniques to investigate how discrimination of translational, rotational and radial global motion is affected by TMS. Prior to applying TMS we established baseline coherence thresholds for global motion perception. Adopting each observer's coherence level at threshold we examined how TMS delivered to V5/MT modulated performance. Importantly, we measured the influence of single-pulse TMS over a broad temporal range to reveal the fine temporal structure of the disruption profile for global motion perception. Results show that the disruption profile consisted of two distinct epochs during which global direction judgments were reliably impaired, separated by an interval in which performance was unaffected. The bimodal nature of the distribution profiles is consistent with feedforward and feedback processing between visual areas mediating global motion processing. We present a novel quantitative model that characterizes the contribution of each process to visual motion perception.

Specificity of human cortical areas for reaches and saccades.

Electrophysiological studies in monkeys have identified effector-related regions in the posterior parietal cortex (PPC). The lateral intraparietal area, for example, responds preferentially for saccades, whereas the parietal reach region responds preferentially for arm movements. However, the degree of effector selectivity actually observed is limited; each area contains neurons selective for the nonpreferred effector, and many neurons in both areas respond for both effectors. We used functional magnetic resonance imaging to assess the degree of effector preference at the population level, focusing on topographically organized regions in the human PPC [visual area V7, intraparietal sulcus 1 (IPS1), and IPS2]. An event-related design adapted from monkey experiments was used. In each trial, an effector cue preceded the appearance of a spatial target, after which a Go signal instructed subjects to produce the specified movement with the specified effector. Our results show that the degree of effector specificity is limited in many cortical areas and transitions gradually from saccade to reach preference as one moves through the hierarchy of areas in the occipital, parietal, and frontal cortices. Saccade preference was observed in visual cortex, including early areas and V7. IPS1 exhibited balanced activation to saccades and reaches, whereas IPS2 showed a weak but significant preference for reaches. In frontal cortex, areas near the central sulcus showed a clear and absolute preference for reaches, whereas the frontal eye field showed little or no effector selectivity. Although these results contradict many theoretical conclusions about effector specificity, they are compatible with the complex picture arising from electrophysiological studies and also with previous imaging studies that reported mostly overlapping saccade- and arm-related activation. The results are also compatible with theories of efficient coding in cortex.