Kalanit Grill-Spector is a Professor in Psychology and the Stanford Neurosciences Institute. Her research examines how the brain processes visual information and perceives it. She uses functional imaging techniques to visualize the living brain in action and understand how it functions to recognize people, objects and places. Additionally, she investigates how the anatomical and functional properties of the brain change from childhood through adolescences to adulthood, and how this development is related to improved recognition abilities. She received her PhD from the Weizmann Institute of Science in Israel and was a postdoctoral fellow in Brain and Cognitive Sciences at MIT before joining Stanford University. She has received several awards and honors including the Human Sciences Frontier Fellowship, the Sloan Fellowship, and the Klingenstein Fellowship in Neuroscience. She has served as an Editor for the Journal of Vision and Neuropsychologia. Presently, she has an active and diverse laboratory at the Psychology Department at Stanford University, she is a leader on the Wu Tsai Big idea project on Neurodevelopment, is a board member of the Center for Cognitive and Neurobiological Imaging at Stanford University, and is the director of the graduate studies in the Department of Psychology.

Academic Appointments

Administrative Appointments

  • Graduate Committee Chair, Department of Psychology, Stanford University (2017 - Present)

Boards, Advisory Committees, Professional Organizations

  • Editor, Neuropsychologia (2016 - Present)
  • Editor, Journal of Vision (2008 - 2012)
  • Editorial Board, NeuroImage (2005 - 2008)

Program Affiliations

  • Symbolic Systems Program

Current Research and Scholarly Interests

For humans, recognition is a natural, effortless skill that occurs within a few hundreds of milliseconds, yet it is one of the least understood aspects of visual perception. Our research utilizes functional imaging (fMRI),diffusion weighted imaging (DWI), computational techniques, and behavioral methods to investigate the neural mechanisms underlying visual recognition in humans. We also examine the development of these mechanisms from childhood to adulthood as well as between populations.

2018-19 Courses

Stanford Advisees

Graduate and Fellowship Programs

All Publications

  • Development differentially sculpts receptive fields across early and high-level human visual cortex Nature Communications Gomez, J., Natu, V., Jeska, B., Barnett, M., Grill-Spector, K. 2018; 9: 788


    Receptive fields (RFs) processing information in restricted parts of the visual field are a key property of visual system neurons. However, how RFs develop in humans is unknown. Using fMRI and population receptive field (pRF) modeling in children and adults, we determine where and how pRFs develop across the ventral visual stream. Here we report that pRF properties in visual field maps, from the first visual area, V1, through the first ventro-occipital area, VO1, are adult-like by age 5. However, pRF properties in face-selective and character-selective regions develop into adulthood, increasing the foveal coverage bias for faces in the right hemisphere and words in the left hemisphere. Eye-tracking indicates that pRF changes are related to changing fixation patterns on words and faces across development. These findings suggest a link between face and word viewing behavior and the differential development of pRFs across visual cortex, potentially due to competition on foveal coverage.

    View details for DOI 10.1038/s41467-018-03166-3

    View details for PubMedCentralID PMC5824941

  • Microstructural proliferation in human cortex is coupled with the development of face processing SCIENCE Gomez, J., Barnett, M. A., Natu, V., Mezer, A., Palomero-Gallagher, N., Weiner, K. S., Amunts, K., Zilles, K., Grill-Spector, K. 2017; 355 (6320): 68-?


    How does cortical tissue change as brain function and behavior improve from childhood to adulthood? By combining quantitative and functional magnetic resonance imaging in children and adults, we find differential development of high-level visual areas that are involved in face and place recognition. Development of face-selective regions, but not place-selective regions, is dominated by microstructural proliferation. This tissue development is correlated with specific increases in functional selectivity to faces, as well as improvements in face recognition, and ultimately leads to differentiated tissue properties between face- and place-selective regions in adulthood, which we validate with postmortem cytoarchitectonic measurements. These data suggest a new model by which emergent brain function and behavior result from cortical tissue proliferation rather than from pruning exclusively.

    View details for DOI 10.1126/science.aag0311

    View details for Web of Science ID 000391739900044

    View details for PubMedID 28059764

    View details for PubMedCentralID PMC5373008

  • The Cytoarchitecture of Domain-specific Regions in Human High-level Visual Cortex CEREBRAL CORTEX Weiner, K. S., Barnett, M. A., Lorenz, S., Caspers, J., Stigliani, A., Amunts, K., Zilles, K., Fischl, B., Grill-Spector, K. 2017; 27 (1): 146-161
  • Encoding model of temporal processing in human visual cortex. Proceedings of the National Academy of Sciences of the United States of America Stigliani, A., Jeska, B., Grill-Spector, K. 2017; 114 (51): E11047–E11056


    How is temporal information processed in human visual cortex? Visual input is relayed to V1 through segregated transient and sustained channels in the retina and lateral geniculate nucleus (LGN). However, there is intense debate as to how sustained and transient temporal channels contribute to visual processing beyond V1. The prevailing view associates transient processing predominately with motion-sensitive regions and sustained processing with ventral stream regions, while the opposing view suggests that both temporal channels contribute to neural processing beyond V1. Using fMRI, we measured cortical responses to time-varying stimuli and then implemented a two temporal channel-encoding model to evaluate the contributions of each channel. Different from the general linear model of fMRI that predicts responses directly from the stimulus, the encoding approach first models neural responses to the stimulus from which fMRI responses are derived. This encoding approach not only predicts cortical responses to time-varying stimuli from milliseconds to seconds but also, reveals differential contributions of temporal channels across visual cortex. Consistent with the prevailing view, motion-sensitive regions and adjacent lateral occipitotemporal regions are dominated by transient responses. However, ventral occipitotemporal regions are driven by both sustained and transient channels, with transient responses exceeding the sustained. These findings propose a rethinking of temporal processing in the ventral stream and suggest that transient processing may contribute to rapid extraction of the content of the visual input. Importantly, our encoding approach has vast implications, because it can be applied with fMRI to decipher neural computations in millisecond resolution in any part of the brain.

    View details for DOI 10.1073/pnas.1704877114

    View details for PubMedID 29208714

    View details for PubMedCentralID PMC5754759

  • Development of Neural Sensitivity to Face Identity Correlates with Perceptual Discriminability. journal of neuroscience Natu, V. S., Barnett, M. A., Hartley, J., Gomez, J., Stigliani, A., Grill-Spector, K. 2016; 36 (42): 10893-10907


    Face perception is subserved by a series of face-selective regions in the human ventral stream, which undergo prolonged development from childhood to adulthood. However, it is unknown how neural development of these regions relates to the development of face-perception abilities. Here, we used functional magnetic resonance imaging (fMRI) to measure brain responses of ventral occipitotemporal regions in children (ages, 5-12 years) and adults (ages, 19-34 years) when they viewed faces that parametrically varied in dissimilarity. Since similar faces generate lower responses than dissimilar faces due to fMRI adaptation, this design objectively evaluates neural sensitivity to face identity across development. Additionally, a subset of subjects participated in a behavioral experiment to assess perceptual discriminability of face identity. Our data reveal three main findings: (1) neural sensitivity to face identity increases with age in face-selective but not object-selective regions; (2) the amplitude of responses to faces increases with age in both face-selective and object-selective regions; and (3) perceptual discriminability of face identity is correlated with the neural sensitivity to face identity of face-selective regions. In contrast, perceptual discriminability is not correlated with the amplitude of response in face-selective regions or of responses of object-selective regions. These data suggest that developmental increases in neural sensitivity to face identity in face-selective regions improve perceptual discriminability of faces. Our findings significantly advance the understanding of the neural mechanisms of development of face perception and open new avenues for using fMRI adaptation to study the neural development of high-level visual and cognitive functions more broadly.Face perception, which is critical for daily social interactions, develops from childhood to adulthood. However, it is unknown what developmental changes in the brain lead to improved performance. Using fMRI in children and adults, we find that from childhood to adulthood, neural sensitivity to changes in face identity increases in face-selective regions. Critically, subjects' perceptual discriminability among faces is linked to neural sensitivity: participants with higher neural sensitivity in face-selective regions demonstrate higher perceptual discriminability. Thus, our results suggest that developmental increases in face-selective regions' sensitivity to face identity improve perceptual discrimination of faces. These findings significantly advance understanding of the neural mechanisms underlying the development of face perception and have important implications for assessing both typical and atypical development.

    View details for PubMedID 27798143

    View details for PubMedCentralID PMC5083016

  • Attention reduces spatial uncertainty in human ventral temporal cortex. Current biology Kay, K. N., Weiner, K. S., Grill-Spector, K. 2015; 25 (5): 595-600


    Ventral temporal cortex (VTC) is the latest stage of the ventral "what" visual pathway, which is thought to code the identity of a stimulus regardless of its position or size [1, 2]. Surprisingly, recent studies show that position information can be decoded from VTC [3-5]. However, the computational mechanisms by which spatial information is encoded in VTC are unknown. Furthermore, how attention influences spatial representations in human VTC is also unknown because the effect of attention on spatial representations has only been examined in the dorsal "where" visual pathway [6-10]. Here, we fill these significant gaps in knowledge using an approach that combines functional magnetic resonance imaging and sophisticated computational methods. We first develop a population receptive field (pRF) model [11, 12] of spatial responses in human VTC. Consisting of spatial summation followed by a compressive nonlinearity, this model accurately predicts responses of individual voxels to stimuli at any position and size, explains how spatial information is encoded, and reveals a functional hierarchy in VTC. We then manipulate attention and use our model to decipher the effects of attention. We find that attention to the stimulus systematically and selectively modulates responses in VTC, but not early visual areas. Locally, attention increases eccentricity, size, and gain of individual pRFs, thereby increasing position tolerance. However, globally, these effects reduce uncertainty regarding stimulus location and actually increase position sensitivity of distributed responses across VTC. These results demonstrate that attention actively shapes and enhances spatial representations in the ventral visual pathway.

    View details for DOI 10.1016/j.cub.2014.12.050

    View details for PubMedID 25702580

    View details for PubMedCentralID PMC4348205

  • Functionally defined white matter reveals segregated pathways in human ventral temporal cortex associated with category-specific processing. Neuron Gomez, J., Pestilli, F., Witthoft, N., Golarai, G., Liberman, A., Poltoratski, S., Yoon, J., Grill-Spector, K. 2015; 85 (1): 216-227


    It is unknown if the white-matter properties associated with specific visual networks selectively affect category-specific processing. In a novel protocol we combined measurements of white-matter structure, functional selectivity, and behavior in the same subjects. We find two parallel white-matter pathways along the ventral temporal lobe connecting to either face-selective or place-selective regions. Diffusion properties of portions of these tracts adjacent to face- and place-selective regions of ventral temporal cortex correlate with behavioral performance for face or place processing, respectively. Strikingly, adults with developmental prosopagnosia (face blindness) express an atypical structure-behavior relationship near face-selective cortex, suggesting that white-matter atypicalities in this region may have behavioral consequences. These data suggest that examining the interplay between cortical function, anatomical connectivity, and visual behavior is integral to understanding functional networks and their role in producing visual abilities and deficits.

    View details for DOI 10.1016/j.neuron.2014.12.027

    View details for PubMedID 25569351

    View details for PubMedCentralID PMC4287959

  • The functional architecture of the ventral temporal cortex and its role in categorization. Nature reviews. Neuroscience Grill-Spector, K., Weiner, K. S. 2014; 15 (8): 536-548


    Visual categorization is thought to occur in the human ventral temporal cortex (VTC), but how this categorization is achieved is still largely unknown. In this Review, we consider the computations and representations that are necessary for categorization and examine how the microanatomical and macroanatomical layout of the VTC might optimize them to achieve rapid and flexible visual categorization. We propose that efficient categorization is achieved by organizing representations in a nested spatial hierarchy in the VTC. This spatial hierarchy serves as a neural infrastructure for the representational hierarchy of visual information in the VTC and thereby enables flexible access to category information at several levels of abstraction.

    View details for DOI 10.1038/nrn3747

    View details for PubMedID 24962370

  • Data on a cytoarchitectonic brain atlas: effects of brain template and a comparison to a multimodal atlas. Data in brief Rosenke, M., Weiner, K. S., Barnett, M. A., Zilles, K., Amunts, K., Goebel, R., Grill-Spector, K. 2017; 12: 327-332


    The data presented here are related to the research article: "A cross-validated cytoarchitectonic atlas of the human ventral visual stream" in which we developed a cytoarchitectonic atlas of ventral visual cortex. Here, we provide two additional quantifications of this cytoarchitectonic atlas: First, we quantify the effect of brain template on cross-validation performance. The data show a comparison between cortex-based alignment to two templates: the postmortem average brain and the FreeSurfer average brain. Second, we quantify the relationship between this cytoarchitectonic atlas and a recently published multimodal atlas of the human brain (Glasser et al., 2016).

    View details for DOI 10.1016/j.dib.2017.04.007

    View details for PubMedID 28487876

  • Defining the most probable location of the parahippocampal place area using cortex-based alignment and cross-validation. NeuroImage Weiner, K. S., Barnett, M. A., Witthoft, N., Golarai, G., Stigliani, A., Kay, K. N., Gomez, J., Natu, V. S., Amunts, K., Zilles, K., Grill-Spector, K. 2017


    The parahippocampal place area (PPA) is a widely studied high-level visual region in the human brain involved in place and scene processing. The goal of the present study was to identify the most probable location of place-selective voxels in medial ventral temporal cortex. To achieve this goal, we first used cortex-based alignment (CBA) to create a probabilistic place-selective region of interest (ROI) from one group of 12 participants. We then tested how well this ROI could predict place selectivity in each hemisphere within a new group of 12 participants. Our results reveal that a probabilistic ROI (pROI) generated from one group of 12 participants accurately predicts the location and functional selectivity in individual brains from a new group of 12 participants, despite between subject variability in the exact location of place-selective voxels relative to the folding of parahippocampal cortex. Additionally, the prediction accuracy of our pROI is significantly higher than that achieved by volume-based Talairach alignment. Comparing the location of the pROI of the PPA relative to published data from over 500 participants, including data from the Human Connectome Project, shows a striking convergence of the predicted location of the PPA and the cortical location of voxels exhibiting the highest place selectivity across studies using various methods and stimuli. Specifically, the most predictive anatomical location of voxels exhibiting the highest place selectivity in medial ventral temporal cortex is the junction of the collateral and anterior lingual sulci. Methodologically, we make this pROI freely available (, which provides a means to accurately identify a functional region from anatomical MRI data when fMRI data are not available (for example, in patient populations). Theoretically, we consider different anatomical and functional factors that may contribute to the consistent anatomical location of place selectivity relative to the folding of high-level visual cortex.

    View details for DOI 10.1016/j.neuroimage.2017.04.040

    View details for PubMedID 28435097

  • Task alters category representations in prefrontal but not high-level visual cortex. NeuroImage Bugatus, L., Weiner, K. S., Grill-Spector, K. 2017


    A central question in neuroscience is how cognitive tasks affect category representations across the human brain. Regions in lateral occipito-temporal cortex (LOTC), ventral temporal cortex (VTC), and ventro-lateral prefrontal cortex (VLFPC) constitute the extended "what" pathway, which is considered instrumental for visual category processing. However, it is unknown (1) whether distributed responses across LOTC, VTC, and VLPFC explicitly represent category, task, or some combination of both, and (2) in what way representations across these subdivisions of the extended 'what' pathway may differ. To fill these gaps in knowledge, we scanned 12 participants using fMRI to test the effect of category and task on distributed responses across LOTC, VTC, and VLPFC. Results reveal that task and category modulate responses in both high-level visual regions, as well as prefrontal cortex. However, we found fundamentally different types of representations across the brain. Distributed responses in high-level visual regions are more strongly driven by category than task, and exhibit task-independent category representations. In contrast, distributed responses in prefrontal cortex are more strongly driven by task than category, and contain task-dependent category representations. Together, these findings of differential representations across the brain support a new idea that LOTC and VTC maintain stable category representations allowing efficient processing of visual information, while prefrontal cortex contains flexible representations in which category information may emerge only when relevant to the task.

    View details for DOI 10.1016/j.neuroimage.2017.03.062

    View details for PubMedID 28389381

  • A cross-validated cytoarchitectonic atlas of the human ventral visual stream. NeuroImage Rosenke, M., Weiner, K. S., Barnett, M. A., Zilles, K., Amunts, K., Goebel, R., Grill-Spector, K. 2017


    The human ventral visual stream consists of several areas that are considered processing stages essential for perception and recognition. A fundamental microanatomical feature differentiating areas is cytoarchitecture, which refers to the distribution, size, and density of cells across cortical layers. Because cytoarchitectonic structure is measured in 20-micron-thick histological slices of postmortem tissue, it is difficult to assess (a) how anatomically consistent these areas are across brains and (b) how they relate to brain parcellations obtained with prevalent neuroimaging methods, acquired at the millimeter and centimeter scale. Therefore, the goal of this study was to (a) generate a cross-validated cytoarchitectonic atlas of the human ventral visual stream on a whole brain template that is commonly used in neuroimaging studies and (b) to compare this atlas to a recently published retinotopic parcellation of visual cortex (Wang et al., 2014). To achieve this goal, we generated an atlas of eight cytoarchitectonic areas: four areas in the occipital lobe (hOc1-hOc4v) and four in the fusiform gyrus (FG1-FG4), then we tested how the different alignment techniques affect the accuracy of the resulting atlas. Results show that both cortex-based alignment (CBA) and nonlinear volumetric alignment (NVA) generate an atlas with better cross-validation performance than affine volumetric alignment (AVA). Additionally, CBA outperformed NVA in 6/8 of the cytoarchitectonic areas. Finally, the comparison of the cytoarchitectonic atlas to a retinotopic atlas shows a clear correspondence between cytoarchitectonic and retinotopic areas in the ventral visual stream. The successful performance of CBA suggests a coupling between cytoarchitectonic areas and macroanatomical landmarks in the human ventral visual stream, and furthermore, that this coupling can be utilized for generating an accurate group atlas. In addition, the coupling between cytoarchitecture and retinotopy highlights the potential use of this atlas in understanding how anatomical features contribute to brain function. We make this cytoarchitectonic atlas freely available in both BrainVoyager and FreeSurfer formats ( The availability of this atlas will enable future studies to link cytoarchitectonic organization to other parcellations of the human ventral visual stream with potential to advance the understanding of this pathway in typical and atypical populations.

    View details for DOI 10.1016/j.neuroimage.2017.02.040

    View details for PubMedID 28213120

  • Experience Shapes the Development of Neural Substrates of Face Processing in Human Ventral Temporal Cortex CEREBRAL CORTEX Golarai, G., Liberman, A., Grill-Spector, K. 2017; 27 (2): 1229-1244
  • Two New Cytoarchitectonic Areas on the Human Mid-Fusiform Gyrus CEREBRAL CORTEX Lorenz, S., Weiner, K. S., Caspers, J., Mohlberg, H., Schleicher, A., Bludau, S., Eickhoff, S., Grill-Spector, K., Zilles, K., Amunts, K. 2017; 27 (1): 373-385
  • The Cytoarchitecture of Domain-specific Regions in Human High-level Visual Cortex. Cerebral cortex Weiner, K. S., Barnett, M. A., Lorenz, S., Caspers, J., Stigliani, A., Amunts, K., Zilles, K., Fischl, B., Grill-Spector, K. 2016: -?


    A fundamental hypothesis in neuroscience proposes that underlying cellular architecture (cytoarchitecture) contributes to the functionality of a brain area. However, this hypothesis has not been tested in human ventral temporal cortex (VTC) that contains domain-specific regions causally involved in perception. To fill this gap in knowledge, we used cortex-based alignment to register functional regions from living participants to cytoarchitectonic areas in ex vivo brains. This novel approach reveals 3 findings. First, there is a consistent relationship between domain-specific regions and cytoarchitectonic areas: each functional region is largely restricted to 1 cytoarchitectonic area. Second, extracting cytoarchitectonic profiles from face- and place-selective regions after back-projecting each region to 20-μm thick histological sections indicates that cytoarchitectonic properties distinguish these regions from each other. Third, some cytoarchitectonic areas contain more than 1 domain-specific region. For example, face-, body-, and character-selective regions are located within the same cytoarchitectonic area. We summarize these findings with a parsimonious hypothesis incorporating how cellular properties may contribute to functional specialization in human VTC. Specifically, we link computational principles to correlated axes of functional and cytoarchitectonic segregation in human VTC, in which parallel processing across domains occurs along a lateral-medial axis while transformations of information within domain occur along an anterior-posterior axis.

    View details for PubMedID 27909003

  • The Face-Processing Network Is Resilient to Focal Resection of Human Visual Cortex. journal of neuroscience Weiner, K. S., Jonas, J., Gomez, J., Maillard, L., Brissart, H., Hossu, G., Jacques, C., Loftus, D., Colnat-Coulbois, S., Stigliani, A., Barnett, M. A., Grill-Spector, K., Rossion, B. 2016; 36 (32): 8425-8440


    Human face perception requires a network of brain regions distributed throughout the occipital and temporal lobes with a right hemisphere advantage. Present theories consider this network as either a processing hierarchy beginning with the inferior occipital gyrus (occipital face area; IOG-faces/OFA) or a multiple-route network with nonhierarchical components. The former predicts that removing IOG-faces/OFA will detrimentally affect downstream stages, whereas the latter does not. We tested this prediction in a human patient (Patient S.P.) requiring removal of the right inferior occipital cortex, including IOG-faces/OFA. We acquired multiple fMRI measurements in Patient S.P. before and after a preplanned surgery and multiple measurements in typical controls, enabling both within-subject/across-session comparisons (Patient S.P. before resection vs Patient S.P. after resection) and between-subject/across-session comparisons (Patient S.P. vs controls). We found that the spatial topology and selectivity of downstream ipsilateral face-selective regions were stable 1 and 8 month(s) after surgery. Additionally, the reliability of distributed patterns of face selectivity in Patient S.P. before versus after resection was not different from across-session reliability in controls. Nevertheless, postoperatively, representations of visual space were typical in dorsal face-selective regions but atypical in ventral face-selective regions and V1 of the resected hemisphere. Diffusion weighted imaging in Patient S.P. and controls identifies white matter tracts connecting retinotopic areas to downstream face-selective regions, which may contribute to the stable and plastic features of the face network in Patient S.P. after surgery. Together, our results support a multiple-route network of face processing with nonhierarchical components and shed light on stable and plastic features of high-level visual cortex following focal brain damage.Brain networks consist of interconnected functional regions commonly organized in processing hierarchies. Prevailing theories predict that damage to the input of the hierarchy will detrimentally affect later stages. We tested this prediction with multiple brain measurements in a rare human patient requiring surgical removal of the putative input to a network processing faces. Surprisingly, the spatial topology and selectivity of downstream face-selective regions are stable after surgery. Nevertheless, representations of visual space were typical in dorsal face-selective regions but atypical in ventral face-selective regions and V1. White matter connections from outside the face network may support these stable and plastic features. As processing hierarchies are ubiquitous in biological and nonbiological systems, our results have pervasive implications for understanding the construction of resilient networks.

    View details for DOI 10.1523/JNEUROSCI.4509-15.2016

    View details for PubMedID 27511014

    View details for PubMedCentralID PMC4978802

  • Learning the 3-D structure of objects from 2-D views depends on shape, not format JOURNAL OF VISION Tian, M., Yamins, D., Grill-Spector, K. 2016; 16 (7)


    Humans can learn to recognize new objects just from observing example views. However, it is unknown what structural information enables this learning. To address this question, we manipulated the amount of structural information given to subjects during unsupervised learning by varying the format of the trained views. We then tested how format affected participants' ability to discriminate similar objects across views that were rotated 90° apart. We found that, after training, participants' performance increased and generalized to new views in the same format. Surprisingly, the improvement was similar across line drawings, shape from shading, and shape from shading + stereo even though the latter two formats provide richer depth information compared to line drawings. In contrast, participants' improvement was significantly lower when training used silhouettes, suggesting that silhouettes do not have enough information to generate a robust 3-D structure. To test whether the learned object representations were format-specific or format-invariant, we examined if learning novel objects from example views transfers across formats. We found that learning objects from example line drawings transferred to shape from shading and vice versa. These results have important implications for theories of object recognition because they suggest that (a) learning the 3-D structure of objects does not require rich structural cues during training as long as shape information of internal and external features is provided and (b) learning generates shape-based object representations independent of the training format.

    View details for DOI 10.1167/16.7.7

    View details for Web of Science ID 000377938400007

    View details for PubMedID 27153196

    View details for PubMedCentralID PMC4898268

  • Corresponding ECoG and fMRI category-selective signals in human ventral temporal cortex. Neuropsychologia Jacques, C., Witthoft, N., Weiner, K. S., Foster, B. L., Rangarajan, V., Hermes, D., Miller, K. J., Parvizi, J., Grill-Spector, K. 2016; 83: 14-28


    Functional magnetic resonance imaging (fMRI) and electrocorticography (ECoG) research have been influential in revealing the functional characteristics of category-selective responses in human ventral temporal cortex (VTC). One important, but unanswered, question is how these two types of measurements might be related with respect to the VTC. Here we examined which components of the ECoG signal correspond to the fMRI response, by using a rare opportunity to measure both fMRI and ECoG responses from the same individuals to images of exemplars of various categories including faces, limbs, cars and houses. Our data reveal three key findings. First, we discovered that the coupling between fMRI and ECoG responses is frequency and time dependent. The strongest and most sustained correlation is observed between fMRI and high frequency broadband (HFB) ECoG responses (30-160hz). In contrast, the correlation between fMRI and ECoG signals in lower frequency bands is temporally transient, where the correlation is initially positive, but then tapers off or becomes negative. Second, we find that the strong and positive correlation between fMRI and ECoG signals in all frequency bands emerges rapidly around 100ms after stimulus onset, together with the onset of the first stimulus-driven neural signals in VTC. Third, we find that the spatial topology and representational structure of category-selectivity in VTC reflected in ECoG HFB responses mirrors the topology and structure observed with fMRI. These findings of a strong and rapid coupling between fMRI and HFB responses validate fMRI measurements of functional selectivity with recordings of direct neural activity and suggest that fMRI category-selective signals in VTC are associated with feed-forward neural processing.

    View details for DOI 10.1016/j.neuropsychologia.2015.07.024

    View details for PubMedID 26212070

  • Experience Shapes the Development of Neural Substrates of Face Processing in Human Ventral Temporal Cortex. Cerebral cortex Golarai, G., Liberman, A., Grill-Spector, K. 2015


    In adult humans, the ventral temporal cortex (VTC) represents faces in a reproducible topology. However, it is unknown what role visual experience plays in the development of this topology. Using functional magnetic resonance imaging in children and adults, we found a sequential development, in which the topology of face-selective activations across the VTC was matured by age 7, but the spatial extent and degree of face selectivity continued to develop past age 7 into adulthood. Importantly, own- and other-age faces were differentially represented, both in the distributed multivoxel patterns across the VTC, and also in the magnitude of responses of face-selective regions. These results provide strong evidence that experience shapes cortical representations of faces during development from childhood to adulthood. Our findings have important implications for the role of experience and age in shaping the neural substrates of face processing in the human VTC.

    View details for PubMedID 26683171

  • Two New Cytoarchitectonic Areas on the Human Mid-Fusiform Gyrus. Cerebral cortex Lorenz, S., Weiner, K. S., Caspers, J., Mohlberg, H., Schleicher, A., Bludau, S., Eickhoff, S. B., Grill-Spector, K., Zilles, K., Amunts, K. 2015


    Areas of the fusiform gyrus (FG) within human ventral temporal cortex (VTC) process high-level visual information associated with faces, limbs, words, and places. Since classical cytoarchitectonic maps do not adequately reflect the functional and structural heterogeneity of the VTC, we studied the cytoarchitectonic segregation in a region, which is rostral to the recently identified cytoarchitectonic areas FG1 and FG2. Using an observer-independent and statistically testable parcellation method, we identify 2 new areas, FG3 and FG4, in 10 human postmortem brains on the mid-FG. The mid-fusiform sulcus reliably identifies the cytoarchitectonic transition between FG3 and FG4. We registered these cytoarchitectonic areas to the common reference space of the single-subject Montreal Neurological Institute (MNI) template and generated probability maps, which reflect the intersubject variability of both areas. Future studies can relate in vivo neuroimaging data with these microscopically defined cortical areas to functional parcellations. We discuss these results in the context of both large-scale functional maps and fine-scale functional clusters that have been identified within the human VTC. We propose that our observer-independent cytoarchitectonic parcellation of the FG better explains the functional heterogeneity of the FG compared with the homogeneity of classic cytoarchitectonic maps.

    View details for PubMedID 26464475

  • Temporal Processing Capacity in High-Level Visual Cortex Is Domain Specific. journal of neuroscience Stigliani, A., Weiner, K. S., Grill-Spector, K. 2015; 35 (36): 12412-12424


    Prevailing hierarchical models propose that temporal processing capacity-the amount of information that a brain region processes in a unit time-decreases at higher stages in the ventral stream regardless of domain. However, it is unknown if temporal processing capacities are domain general or domain specific in human high-level visual cortex. Using a novel fMRI paradigm, we measured temporal capacities of functional regions in high-level visual cortex. Contrary to hierarchical models, our data reveal domain-specific processing capacities as follows: (1) regions processing information from different domains have differential temporal capacities within each stage of the visual hierarchy and (2) domain-specific regions display the same temporal capacity regardless of their position in the processing hierarchy. In general, character-selective regions have the lowest capacity, face- and place-selective regions have an intermediate capacity, and body-selective regions have the highest capacity. Notably, domain-specific temporal processing capacities are not apparent in V1 and have perceptual implications. Behavioral testing revealed that the encoding capacity of body images is higher than that of characters, faces, and places, and there is a correspondence between peak encoding rates and cortical capacities for characters and bodies. The present evidence supports a model in which the natural statistics of temporal information in the visual world may affect domain-specific temporal processing and encoding capacities. These findings suggest that the functional organization of high-level visual cortex may be constrained by temporal characteristics of stimuli in the natural world, and this temporal capacity is a characteristic of domain-specific networks in high-level visual cortex.Visual stimuli bombard us at different rates every day. For example, words and scenes are typically stationary and vary at slow rates. In contrast, bodies are dynamic and typically change at faster rates. Using a novel fMRI paradigm, we measured temporal processing capacities of functional regions in human high-level visual cortex. Contrary to prevailing theories, we find that different regions have different processing capacities, which have behavioral implications. In general, character-selective regions have the lowest capacity, face- and place-selective regions have an intermediate capacity, and body-selective regions have the highest capacity. These results suggest that temporal processing capacity is a characteristic of domain-specific networks in high-level visual cortex and contributes to the segregation of cortical regions.

    View details for DOI 10.1523/JNEUROSCI.4822-14.2015

    View details for PubMedID 26354910

  • The evolution of face processing networks. Trends in cognitive sciences Weiner, K. S., Grill-Spector, K. 2015; 19 (5): 240-241


    Recent studies in marmosets, macaques, and humans have begun to show commonalities and differences in the evolution of face processing networks. Despite differences in brain size and gyrification across species, myelination and motion may be key anatomical and functional features contributing to the surprising similarity of face networks across species.

    View details for DOI 10.1016/j.tics.2015.03.010

    View details for PubMedID 25840651

  • Feature saliency and feedback information interactively impact visual category learning FRONTIERS IN PSYCHOLOGY Hammer, R., Sloutsky, V., Grill-Spector, K. 2015; 6


    Visual category learning (VCL) involves detecting which features are most relevant for categorization. VCL relies on attentional learning, which enables effectively redirecting attention to object's features most relevant for categorization, while 'filtering out' irrelevant features. When features relevant for categorization are not salient, VCL relies also on perceptual learning, which enables becoming more sensitive to subtle yet important differences between objects. Little is known about how attentional learning and perceptual learning interact when VCL relies on both processes at the same time. Here we tested this interaction. Participants performed VCL tasks in which they learned to categorize novel stimuli by detecting the feature dimension relevant for categorization. Tasks varied both in feature saliency (low-saliency tasks that required perceptual learning vs. high-saliency tasks), and in feedback information (tasks with mid-information, moderately ambiguous feedback that increased attentional load, vs. tasks with high-information non-ambiguous feedback). We found that mid-information and high-information feedback were similarly effective for VCL in high-saliency tasks. This suggests that an increased attentional load, associated with the processing of moderately ambiguous feedback, has little effect on VCL when features are salient. In low-saliency tasks, VCL relied on slower perceptual learning; but when the feedback was highly informative participants were able to ultimately attain the same performance as during the high-saliency VCL tasks. However, VCL was significantly compromised in the low-saliency mid-information feedback task. We suggest that such low-saliency mid-information learning scenarios are characterized by a 'cognitive loop paradox' where two interdependent learning processes have to take place simultaneously.

    View details for DOI 10.3389/fpsyg.2015.00074

    View details for Web of Science ID 000349595900001

    View details for PubMedID 25745404

  • Spatiotemporal information during unsupervised learning enhances viewpoint invariant object recognition JOURNAL OF VISION Tian, M., Grill-Spector, K. 2015; 15 (6)


    Recognizing objects is difficult because it requires both linking views of an object that can be different and distinguishing objects with similar appearance. Interestingly, people can learn to recognize objects across views in an unsupervised way, without feedback, just from the natural viewing statistics. However, there is intense debate regarding what information during unsupervised learning is used to link among object views. Specifically, researchers argue whether temporal proximity, motion, or spatiotemporal continuity among object views during unsupervised learning is beneficial. Here, we untangled the role of each of these factors in unsupervised learning of novel three-dimensional (3-D) objects. We found that after unsupervised training with 24 object views spanning a 180° view space, participants showed significant improvement in their ability to recognize 3-D objects across rotation. Surprisingly, there was no advantage to unsupervised learning with spatiotemporal continuity or motion information than training with temporal proximity. However, we discovered that when participants were trained with just a third of the views spanning the same view space, unsupervised learning via spatiotemporal continuity yielded significantly better recognition performance on novel views than learning via temporal proximity. These results suggest that while it is possible to obtain view-invariant recognition just from observing many views of an object presented in temporal proximity, spatiotemporal information enhances performance by producing representations with broader view tuning than learning via temporal association. Our findings have important implications for theories of object recognition and for the development of computational algorithms that learn from examples.

    View details for DOI 10.1167/15.6.7

    View details for Web of Science ID 000357858600007

    View details for PubMedID 26024454

  • Electrical Stimulation of the Left and Right Human Fusiform Gyrus Causes Different Effects in Conscious Face Perception JOURNAL OF NEUROSCIENCE Rangarajan, V., Hermes, D., Foster, B. L., Weiner, K. S., Jacques, C., Grill-Spector, K., Parvizi, J. 2014; 34 (38): 12828-12836


    Neuroimaging and electrophysiological studies across species have confirmed bilateral face-selective responses in the ventral temporal cortex (VTC) and prosopagnosia is reported in patients with lesions in the VTC including the fusiform gyrus (FG). As imaging and electrophysiological studies provide correlative evidence, and brain lesions often comprise both white and gray matter structures beyond the FG, we designed the current study to explore the link between face-related electrophysiological responses in the FG and the causal effects of electrical stimulation of the left or right FG in face perception. We used a combination of electrocorticography (ECoG) and electrical brain stimulation (EBS) in 10 human subjects implanted with intracranial electrodes in either the left (5 participants, 30 FG sites) or right (5 participants, 26 FG sites) hemispheres. We identified FG sites with face-selective ECoG responses, and recorded perceptual reports during EBS of these sites. In line with existing literature, face-selective ECoG responses were present in both left and right FG sites. However, when the same sites were stimulated, we observed a striking difference between hemispheres. Only EBS of the right FG caused changes in the conscious perception of faces, whereas EBS of strongly face-selective regions in the left FG produced non-face-related visual changes, such as phosphenes. This study examines the relationship between correlative versus causal nature of ECoG and EBS, respectively, and provides important insight into the differential roles of the right versus left FG in conscious face perception.

    View details for DOI 10.1523/JNEUROSCI.0527-14.2014

    View details for Web of Science ID 000341767800019

    View details for PubMedCentralID PMC4166163

  • Where Is Human V4? Predicting the Location of hV4 and VO1 from Cortical Folding. Cerebral cortex Witthoft, N., Nguyen, M. L., Golarai, G., LaRocque, K. F., Liberman, A., Smith, M. E., Grill-Spector, K. 2014; 24 (9): 2401-2408


    A strong relationship between cortical folding and the location of primary sensory areas in the human brain is well established. However, it is unknown if coupling between functional responses and gross anatomy is found at higher stages of sensory processing. We examined the relationship between cortical folding and the location of the retinotopic maps hV4 and VO1, which are intermediate stages in the human ventral visual processing stream. Our data show a consistent arrangement of the eccentricity maps within hV4 and VO1 with respect to anatomy, with the consequence that the hV4/VO1 boundary is found consistently in the posterior transverse collateral sulcus (ptCoS) despite individual variability in map size and cortical folding. Understanding this relationship allowed us to predict the location of visual areas hV4 and VO1 in a separate set of individuals, using only their anatomies, with >85% accuracy. These findings have important implications for understanding the relation between cortical folding and functional maps as well as for defining visual areas from anatomical landmarks alone.

    View details for DOI 10.1093/cercor/bht092

    View details for PubMedID 23592823

  • The mid-fusiform sulcus: A landmark identifying both cytoarchitectonic and functional divisions of human ventral temporal cortex NEUROIMAGE Weiner, K. S., Golarai, G., Caspers, J., Chuapoco, M. R., Mohlberg, H., Zilles, K., Amunts, K., Grill-Spector, K. 2014; 84: 453-465


    Human ventral temporal cortex (VTC) plays a pivotal role in high-level vision. An under-studied macroanatomical feature of VTC is the mid-fusiform sulcus (MFS), a shallow longitudinal sulcus separating the lateral and medial fusiform gyrus (FG). Here, we quantified the morphological features of the MFS in 69 subjects (ages 7-40), and investigated its relationship to both cytoarchitectonic and functional divisions of VTC with four main findings. First, despite being a minor sulcus, we found that the MFS is a stable macroanatomical structure present in all 138 hemispheres with morphological characteristics developed by age 7. Second, the MFS is the locus of a lateral-medial cytoarchitectonic transition within the posterior FG serving as the boundary between cytoarchitectonic regions FG1 and FG2. Third, the MFS predicts a lateral-medial functional transition in eccentricity bias representations in children, adolescents, and adults. Fourth, the anterior tip of the MFS predicts the location of a face-selective region, mFus-faces/FFA-2. These findings are the first to illustrate that a macroanatomical landmark identifies both cytoarchitectonic and functional divisions of high-level sensory cortex in humans and have important implications for understanding functional and structural organization in the human brain.

    View details for DOI 10.1016/j.neuroimage.2013.08.068

    View details for Web of Science ID 000328868600042

    View details for PubMedID 24021838

  • Global Similarity and Pattern Separation in the Human Medial Temporal Lobe Predict Subsequent Memory JOURNAL OF NEUROSCIENCE LaRocque, K. F., Smith, M. E., Carr, V. A., Witthoft, N., Grill-Spector, K., Wagner, A. D. 2013; 33 (13): 5466-5474


    Intense debate surrounds the role of medial temporal lobe (MTL) structures in recognition memory. Using high-resolution fMRI and analyses of pattern similarity in humans, we examined the encoding computations subserved by MTL subregions. Specifically, we tested the theory that MTL cortex supports memory by encoding overlapping representations, whereas hippocampus supports memory by encoding pattern-separated representations. Consistent with this view, the relationship between encoding pattern similarity and subsequent memory dissociated MTL cortex and hippocampus: later memory was predicted by greater across-item pattern similarity in perirhinal cortex and in parahippocampal cortex, but greater pattern distinctiveness in hippocampus. Additionally, by comparing neural patterns elicited by individual stimuli regardless of subsequent memory, we found that perirhinal cortex and parahippocampal cortex exhibited differential content sensitivity for multiple stimulus categories, whereas hippocampus failed to demonstrate content sensitivity. These data provide novel evidence that complementary MTL encoding computations subserve declarative memory.

    View details for DOI 10.1523/JNEUROSCI.4293-12.2013

    View details for Web of Science ID 000316948600005

    View details for PubMedID 23536062

    View details for PubMedCentralID PMC3643502

  • Neural representations of faces and limbs neighbor in human high-level visual cortex: evidence for a new organization principle PSYCHOLOGICAL RESEARCH-PSYCHOLOGISCHE FORSCHUNG Weiner, K. S., Grill-Spector, K. 2013; 77 (1): 74-97


    Neurophysiology and optical imaging studies in monkeys and functional magnetic resonance imaging (fMRI) studies in both monkeys and humans have localized clustered neural responses in inferotemporal cortex selective for images of biologically relevant categories, such as faces and limbs. Using higher resolution (1.5 mm voxels) fMRI scanning methods than past studies (3-5 mm voxels), we recently reported a network of multiple face- and limb-selective regions that neighbor one another in human ventral temporal cortex (Weiner and Grill-Spector, Neuroimage, 52(4):1559-1573, 2010) and lateral occipitotemporal cortex (Weiner and Grill-Spector, Neuroimage, 56(4):2183-2199, 2011). Here, we expand on three basic organization principles of high-level visual cortex revealed by these findings: (1) consistency in the anatomical location of functional regions, (2) preserved spatial relationship among functional regions, and (3) a topographic organization of face- and limb-selective regions in adjacent and alternating clusters. We highlight the implications of this structure in comparing functional brain organization between typical and atypical populations. We conclude with a new model of high-level visual cortex consisting of ventral, lateral, and dorsal components, where multimodal processing related to vision, action, haptics, and language converges in the lateral pathway.

    View details for DOI 10.1007/s00426-011-0392-x

    View details for Web of Science ID 000313053700008

    View details for PubMedID 22139022

  • Electrical Stimulation of Human Fusiform Face-Selective Regions Distorts Face Perception JOURNAL OF NEUROSCIENCE Parvizi, J., Jacques, C., Foster, B. L., Withoft, N., Rangarajan, V., Weiner, K. S., Grill-Spector, K. 2012; 32 (43): 14915-14920


    Face-selective neural responses in the human fusiform gyrus have been widely examined. However, their causal role in human face perception is largely unknown. Here, we used a multimodal approach of electrocorticography (ECoG), high-resolution functional magnetic resonance imaging (fMRI), and electrical brain stimulation (EBS) to directly investigate the causal role of face-selective neural responses of the fusiform gyrus (FG) in face perception in a patient implanted with subdural electrodes in the right inferior temporal lobe. High-resolution fMRI identified two distinct FG face-selective regions [mFus-faces and pFus-faces (mid and posterior fusiform, respectively)]. ECoG revealed a striking anatomical and functional correspondence with fMRI data where a pair of face-selective electrodes, positioned 1 cm apart, overlapped mFus-faces and pFus-faces, respectively. Moreover, electrical charge delivered to this pair of electrodes induced a profound face-specific perceptual distortion during viewing of real faces. Specifically, the subject reported a "metamorphosed" appearance of faces of people in the room. Several controls illustrate the specificity of the effect to the perception of faces. EBS of mFus-faces and pFus-faces neither produced a significant deficit in naming pictures of famous faces on the computer, nor did it affect the appearance of nonface objects. Further, the appearance of faces remained unaffected during both sham stimulation and stimulation of a pair of nearby electrodes that were not face-selective. Overall, our findings reveal a striking convergence of fMRI, ECoG, and EBS, which together offer a rare causal link between functional subsets of the human FG network and face perception.

    View details for DOI 10.1523/JNEUROSCI.2609-12.2012

    View details for Web of Science ID 000310523900008

    View details for PubMedID 23100414

    View details for PubMedCentralID PMC3517886

  • Face-likeness and image variability drive responses in human face-selective ventral regions HUMAN BRAIN MAPPING Davidenko, N., Remus, D. A., Grill-Spector, K. 2012; 33 (10): 2334-2349


    The human ventral visual stream contains regions that respond selectively to faces over objects. However, it is unknown whether responses in these regions correlate with how face-like stimuli appear. Here, we use parameterized face silhouettes to manipulate the perceived face-likeness of stimuli and measure responses in face- and object-selective ventral regions with high-resolution fMRI. We first use "concentric hyper-sphere" (CH) sampling to define face silhouettes at different distances from the prototype face. Observers rate the stimuli as progressively more face-like the closer they are to the prototype face. Paradoxically, responses in both face- and object-selective regions decrease as face-likeness ratings increase. Because CH sampling produces blocks of stimuli whose variability is negatively correlated with face-likeness, this effect may be driven by more adaptation during high face-likeness (low-variability) blocks than during low face-likeness (high-variability) blocks. We tested this hypothesis by measuring responses to matched-variability (MV) blocks of stimuli with similar face-likeness ratings as with CH sampling. Critically, under MV sampling, we find a face-specific effect: responses in face-selective regions gradually increase with perceived face-likeness, but responses in object-selective regions are unchanged. Our studies provide novel evidence that face-selective responses correlate with the perceived face-likeness of stimuli, but this effect is revealed only when image variability is controlled across conditions. Finally, our data show that variability is a powerful factor that drives responses across the ventral stream. This indicates that controlling variability across conditions should be a critical tool in future neuroimaging studies of face and object representation.

    View details for DOI 10.1002/hbm.21367

    View details for Web of Science ID 000308589400007

    View details for PubMedID 21823208

  • White matter microstructure on diffusion tensor imaging is associated with conventional magnetic resonance imaging findings and cognitive function in adolescents born preterm DEVELOPMENTAL MEDICINE AND CHILD NEUROLOGY Feldman, H. M., Lee, E. S., Loe, I. M., Yeom, K. W., Grill-Spector, K., Luna, B. 2012; 54 (9): 809-814


    Diffusion tensor imaging (DTI) was used to evaluate white matter architecture after preterm birth. The goals were (1) to compare white matter microstructure in two cohorts of preterm- and term-born children; and (2) within preterm groups, to determine if sex, gestational age, birthweight, white matter injury score from conventional magnetic resonance imaging (MRI), or IQ was associated with DTI measures.Participants (n=121; 66 females, 55 males) were aged 9 to 16 years. They comprised 58 preterm children (site 1, n=25; and site 2, n=33) born at less than 36 weeks' gestation (mean 29.4 wks; birthweight 1289g) and 63 term children (site 1, n=40; site 2, n=23) born at more than 37 weeks' gestation. DTI was analyzed using tract-based spatial statistics. Diffusion measures were fractional anisotropy, axial, radial, and mean diffusivity.In no region of the white matter skeleton was fractional anisotropy lower in the preterm group at either site. Within the preterm groups, fractional anisotropy was significantly associated with white matter injury score, but not sex, gestational age, or birthweight. At site 1, fractional anisotropy was associated with IQ.DTI contributes to understanding individual differences after preterm birth but may not differentiate a relatively high-functioning group of preterm children from a matched group of term-born children.

    View details for DOI 10.1111/j.1469-8749.2012.04378.x

    View details for Web of Science ID 000307468700011

    View details for PubMedID 22803787

    View details for PubMedCentralID PMC3683593

  • The improbable simplicity of the fusiform face area TRENDS IN COGNITIVE SCIENCES Weiner, K. S., Grill-Spector, K. 2012; 16 (5): 251-254


    The fusiform face area (FFA) is described as an easily identifiable module on the fusiform gyrus. However, the organization of face-selective regions in ventral temporal cortex (VTC) is more complex than this prevailing view. We highlight methodological factors contributing to these complexities and the extensive variability in how the FFA is identified. We suggest a series of constraints to aid researchers when defining any functionally specialized region with a pleasing realization: anatomy matters.

    View details for DOI 10.1016/j.tics.2012.03.003

    View details for Web of Science ID 000304026200001

    View details for PubMedID 22481071

  • The Interplay between Feature-Saliency and Feedback Information in Visual Category Learning Tasks. CogSci ... Annual Conference of the Cognitive Science Society. Cognitive Science Society (U.S.). Conference Hammer, R., Sloutsky, V., Grill-Spector, K. 2012; 2012: 420-425


    What is the role of feedback information in different visual category learning (VCL) scenarios? To address this question we tested participants' performance in VCL tasks in which stimuli varied in three feature dimensions, one of which was relevant for the task and the other two were irrelevant. The relevant feature could be identified based on trial-by-trial feedback. In one condition the task relevant and irrelevant features were highly-salient. In the second condition all features had low-visual-saliency. Feedback information was also manipulated: In the high-information condition the task relevant feature could be identified by the information provided in each trial whereas in the mid-information condition the feedback was ambiguous and information from several learning trials was required in order to confidently identify the relevant feature. Surprisingly, our data shows that mid- and high-information feedback are similarly effective in high-saliency VCL tasks. In contrast, in low-saliency VCL tasks, mid-information feedback impairs learning. We suggest that VCL can be done effectively either when feedback is ambiguous or in low-saliency conditions, but not in scenarios when both challenges occur concurrently.

    View details for PubMedID 25346948

    View details for PubMedCentralID PMC4208067

  • Not one extrastriate body area: Using anatomical landmarks, hMT+, and visual field maps to parcellate limb-selective activations in human lateral occipitotemporal cortex NEUROIMAGE Weiner, K. S., Grill-Spector, K. 2011; 56 (4): 2183-2199


    The prevailing view of human lateral occipitotemporal cortex (LOTC) organization suggests a single area selective for images of the human body (extrastriate body area, EBA) that highly overlaps with the human motion-selective complex (hMT+). Using functional magnetic resonance imaging with higher resolution (1.5mm voxels) than past studies (3-4mm voxels), we examined the fine-scale spatial organization of these activations relative to each other, as well as to visual field maps in LOTC. Rather than one contiguous EBA highly overlapping hMT+, results indicate three limb-selective activations organized in a crescent surrounding hMT+: (1) an activation posterior to hMT+ on the lateral occipital sulcus/middle occipital gyrus (LOS/MOG) overlapping the lower vertical meridian shared between visual field maps LO-2 and TO-1, (2) an activation anterior to hMT+ on the middle temporal gyrus (MTG) consistently overlapping the lower vertical meridian of TO-2 and extending outside presently defined visual field maps, and (3) an activation inferior to hMT+ on the inferotemporal gyrus (ITG) overlapping the parafoveal representation of the TO cluster. This crescent organization of limb-selective activations surrounding hMT+ is reproducible over a span of three years and is consistent across different image types used for localization. Further, these regions exhibit differential position properties: preference for contralateral image presentation decreases and preference for foveal presentation increases from the limb-selective LOS to the MTG. Finally, the relationship between limb-selective activations and visual field maps extends to the dorsal stream where a posterior IPS activation overlaps V7. Overall, our measurements demonstrate a series of LOTC limb-selective activations that 1) have separate anatomical and functional boundaries, 2) overlap distinct visual field maps, and 3) illustrate differential position properties. These findings indicate that category selectivity alone is an insufficient organization principle for defining brain areas. Instead, multiple properties are necessary in order to parcellate and understand the functional organization of high-level visual cortex.

    View details for DOI 10.1016/j.neuroimage.2011.03.041

    View details for Web of Science ID 000291457500029

    View details for PubMedID 21439386

  • Sparsely-distributed organization of face and limb activations in human ventral temporal cortex NEUROIMAGE Weiner, K. S., Grill-Spector, K. 2010; 52 (4): 1559-1573


    Functional magnetic resonance imaging (fMRI) has identified face- and body part-selective regions, as well as distributed activation patterns for object categories across human ventral temporal cortex (VTC), eliciting a debate regarding functional organization in VTC and neural coding of object categories. Using high-resolution fMRI, we illustrate that face- and limb-selective activations alternate in a series of largely nonoverlapping clusters in lateral VTC along the inferior occipital gyrus (IOG), fusiform gyrus (FG), and occipito-temporal sulcus (OTS). Both general linear model (GLM) and multivoxel pattern (MVP) analyses show that face- and limb-selective activations minimally overlap and that this organization is consistent across experiments and days. We provide a reliable method to separate two face-selective clusters on the middle and posterior FG (mFus and pFus), and another on the IOG using their spatial relation to limb-selective activations and retinotopic areas hV4, VO-1/2, and hMT+. Furthermore, these activations show a gradient of increasing face selectivity and decreasing limb selectivity from the IOG to the mFus. Finally, MVP analyses indicate that there is differential information for faces in lateral VTC (containing weakly- and highly-selective voxels) relative to non-selective voxels in medial VTC. These findings suggest a sparsely-distributed organization where sparseness refers to the presence of several face- and limb-selective clusters in VTC, and distributed refers to the presence of different amounts of information in highly-, weakly-, and non-selective voxels. Consequently, theories of object recognition should consider the functional and spatial constraints of neural coding across a series of minimally overlapping category-selective clusters that are themselves distributed.

    View details for DOI 10.1016/j.neuroimage.2010.04.262

    View details for Web of Science ID 000280695200044

    View details for PubMedID 20457261

  • fMRI-Adaptation and Category Selectivity in Human Ventral Temporal Cortex: Regional Differences Across Time Scales JOURNAL OF NEUROPHYSIOLOGY Weiner, K. S., Sayres, R., Vinberg, J., Grill-Spector, K. 2010; 103 (6): 3349-3365


    Repeating object images produces stimulus-specific repetition suppression referred to as functional magnetic resonance imaging-adaptation (fMRI-A) in ventral temporal cortex (VTC). However, the effects of stimulus repetition on functional selectivity are largely unknown. We investigated the effects of short-lagged (SL, immediate) and long-lagged (LL, many intervening stimuli) repetitions on category selectivity in VTC using high-resolution fMRI. We asked whether repetition produces scaling or sharpening of fMRI responses both within category-selective regions as well as in the distributed response pattern across VTC. Results illustrate that repetition effects across time scales vary quantitatively along an anterior-posterior axis and qualitatively along a lateral-medial axis. In lateral VTC, both SL and LL repetitions produce proportional fMRI-A with no change in either selectivity or distributed responses as predicted by a scaling model. Further, there is larger fMRI-A in anterior subregions irrespective of category selectivity. Medial VTC exhibits similar scaling effects during SL repetitions. However, for LL repetitions, both the selectivity and distributed pattern of responses vary with category in medial VTC as predicted by a sharpening model. Specifically, there is larger fMRI-A for nonpreferred categories compared with the preferred category, and category selectivity does not predict fMRI-A across the pattern of distributed response. Finally, simulations indicate that different neural mechanisms likely underlie fMRI-A in medial compared to lateral VTC. These results have important implications for future fMRI-A experiments because they suggest that fMRI-A does not reflect a universal neural mechanism and that results of fMRI-A experiments will likely be paradigm independent in lateral VTC but paradigm dependent in medial VTC.

    View details for DOI 10.1152/jn.01108.2009

    View details for Web of Science ID 000278493900035

    View details for PubMedID 20375251

  • The Fusiform Face Area is Enlarged in Williams Syndrome JOURNAL OF NEUROSCIENCE Golarai, G., Hong, S., Haas, B. W., Galaburda, A. M., Mills, D. L., Bellugi, U., Grill-Spector, K., Reiss, A. L. 2010; 30 (19): 6700-6712


    Williams syndrome (WS) is a genetic condition characterized by atypical brain structure, cognitive deficits, and a life-long fascination with faces. Face recognition is relatively spared in WS, despite abnormalities in aspects of face processing and structural alterations in the fusiform gyrus, part of the ventral visual stream. Thus, face recognition in WS may be subserved by abnormal neural substrates in the ventral stream. To test this hypothesis, we used functional magnetic resonance imaging and examined the fusiform face area (FFA), which is implicated in face recognition in typically developed (TD) individuals, but its role in WS is not well understood. We found that the FFA was approximately two times larger among WS than TD participants (both absolutely and relative to the fusiform gyrus), despite apparently normal levels of face recognition performance on a Benton face recognition test. Thus, a larger FFA may play a role in face recognition proficiency among WS.

    View details for DOI 10.1523/JNEUROSCI.4268-09.2010

    View details for Web of Science ID 000277653600023

    View details for PubMedID 20463232

    View details for PubMedCentralID PMC3670816

  • Differential development of the ventral visual cortex extends through adolescence. Frontiers in human neuroscience Golarai, G., Liberman, A., Yoon, J. M., Grill-Spector, K. 2010; 3: 80-?


    The ventral temporal cortex (VTC) in humans includes functionally defined regions that preferentially respond to objects, faces, and places. Recent developmental studies suggest that the face selective region in the fusiform gyrus ('fusiform face area', FFA) undergoes a prolonged development involving substantial increases in its volume after 7 years of age. However, the endpoint of this development is not known. Here we used functional magnetic resonance imaging (fMRI) to examine the development of face-, object- and place selective regions in the VTC of adolescents (12-16 year olds) and adults (18-40 year olds). We found that the volume of face selective activations in the right fusiform gyrus was substantially larger in adults than in adolescents, and was positively correlated with age. This development was associated with higher response amplitudes and selectivity for faces in face selective regions of VTC and increased differentiation of the distributed response patterns to faces versus non-face stimuli across the entire VTC. Furthermore, right FFA size was positively correlated with face recognition memory performance, but not with recognition memory of objects or places. In contrast, the volume of object- and place selective cortical regions or their response amplitudes did not change across these age groups. Thus, we found a striking and prolonged development of face selectivity across the VTC during adolescence that was specifically associated with proficiency in face recognition memory. These findings have important implications for theories of development and functional specialization in VTC.

    View details for DOI 10.3389/neuro.09.080.2009

    View details for PubMedID 20204140

  • Controlling stimulus variability reveals stronger face-selective responses near the average face 17th Annual Meeting on Object Perception, Attention and Memory Davidenko, N., Grill-Spector, K. PSYCHOLOGY PRESS. 2010: 122–26
  • The representation of object viewpoint in human visual cortex NEUROIMAGE Andresen, D. R., Vinberg, J., Grill-Spector, K. 2009; 45 (2): 522-536


    Understanding the nature of object representations in the human brain is critical for understanding the neural basis of invariant object recognition. However, the degree to which object representations are sensitive to object viewpoint is unknown. Using fMRI we employed a parametric approach to examine the sensitivity to object view as a function of rotation (0 degrees-180 degrees ), category (animal/vehicle) and fMRI-adaptation paradigm (short or long-lagged). For both categories and fMRI-adaptation paradigms, object-selective regions recovered from adaptation when a rotated view of an object was shown after adaptation to a specific view of that object, suggesting that representations are sensitive to object rotation. However, we found evidence for differential representations across categories and ventral stream regions. Rotation cross-adaptation was larger for animals than vehicles, suggesting higher sensitivity to vehicle than animal rotation, and was largest in the left fusiform/occipito-temporal sulcus (pFUS/OTS), suggesting that this region has low sensitivity to rotation. Moreover, right pFUS/OTS and FFA responded more strongly to front than back views of animals (without adaptation) and rotation cross-adaptation depended both on the level of rotation and the adapting view. This result suggests a prevalence of neurons that prefer frontal views of animals in fusiform regions. Using a computational model of view-tuned neurons, we demonstrate that differential neural view tuning widths and relative distributions of neural-tuned populations in fMRI voxels can explain the fMRI results. Overall, our findings underscore the utility of parametric approaches for studying the neural basis of object invariance and suggest that there is no complete invariance to object view in the human ventral stream.

    View details for DOI 10.1016/j.neuroimage.2008.11.009

    View details for Web of Science ID 000263863000031

    View details for PubMedID 19100844

  • Fine-Scale Spatial Organization of Face and Object Selectivity in the Temporal Lobe: Do Functional Magnetic Resonance Imaging, Optical Imaging, and Electrophysiology Agree? JOURNAL OF NEUROSCIENCE Op de Beeck, H. P., DiCarlo, J. J., Goense, J. B., Grill-Spector, K., Papanastassiou, A., Tanifuji, M., Tsao, D. Y. 2008; 28 (46): 11796-11801


    The spatial organization of the brain's object and face representations in the temporal lobe is critical for understanding high-level vision and cognition but is poorly understood. Recently, exciting progress has been made using advanced imaging and physiology methods in humans and nonhuman primates, and the combination of such methods may be particularly powerful. Studies applying these methods help us to understand how neuronal activity, optical imaging, and functional magnetic resonance imaging signals are related within the temporal lobe, and to uncover the fine-grained and large-scale spatial organization of object and face representations in the primate brain.

    View details for DOI 10.1523/JNEUROSCI.3799-08.2008

    View details for Web of Science ID 000260827600009

    View details for PubMedID 19005042

  • Relating retinotopic and object-selective responses in human lateral occipital cortex JOURNAL OF NEUROPHYSIOLOGY Sayres, R., Grill-Spector, K. 2008; 100 (1): 249-267


    What is the relationship between retinotopy and object selectivity in human lateral occipital (LO) cortex? We used functional magnetic resonance imaging (fMRI) to examine sensitivity to retinal position and category in LO, an object-selective region positioned posterior to MT along the lateral cortical surface. Six subjects participated in phase-encoded retinotopic mapping experiments as well as block-design experiments in which objects from six different categories were presented at six distinct positions in the visual field. We found substantial position modulation in LO using standard nonobject retinotopic mapping stimuli; this modulation extended beyond the boundaries of visual field maps LO-1 and LO-2. Further, LO showed a pronounced lower visual field bias: more LO voxels represented the lower contralateral visual field, and the mean LO response was higher to objects presented below fixation than above fixation. However, eccentricity effects produced by retinotopic mapping stimuli and objects differed. Whereas LO voxels preferred a range of eccentricities lying mostly outside the fovea in the retinotopic mapping experiment, LO responses were strongest to foveally presented objects. Finally, we found a stronger effect of position than category on both the mean LO response, as well as the distributed response across voxels. Overall these results demonstrate that retinal position exhibits strong effects on neural response in LO and indicates that these position effects may be explained by retinotopic organization.

    View details for DOI 10.1152/jn.01383.2007

    View details for Web of Science ID 000257635400023

    View details for PubMedID 18463186

  • Developmental neuroimaging of the human ventral visual cortex TRENDS IN COGNITIVE SCIENCES Grill-Spector, K., Golarai, G., Gabrieli, J. 2008; 12 (4): 152-162


    Here, we review recent results that investigate the development of the human ventral stream from childhood, through adolescence and into adulthood. Converging evidence suggests a differential developmental trajectory across ventral stream regions, in which face-selective regions show a particularly long developmental time course, taking more than a decade to become adult-like. We discuss the implications of these recent findings, how they relate to age-dependent improvements in recognition memory performance and propose possible neural mechanisms that might underlie this development. These results have important implications regarding the role of experience in shaping the ventral stream and the nature of the underlying representations.

    View details for Web of Science ID 000255469200008

    View details for PubMedID 18359267

  • Object recognition: Insights from advances in fMRI methods CURRENT DIRECTIONS IN PSYCHOLOGICAL SCIENCE Grill-Spector, K., Sayres, R. 2008; 17 (2): 73-79
  • Representation of shapes, edges, and surfaces across multiple cues in the human visual cortex JOURNAL OF NEUROPHYSIOLOGY Vinberg, J., Grill-Spector, K. 2008; 99 (3): 1380-1393


    The lateral occipital complex (LOC) responds preferentially to objects compared with random stimuli or textures independent of the visual cue. However, it is unknown whether the LOC (or other cortical regions) are involved in the processing of edges or global surfaces without shape information. Here, we examined processing of 1) global shape, 2) disconnected edges without a global shape, and 3) global surfaces without edges versus random stimuli across motion and stereo cues. The LOC responded more strongly to global shapes than to edges, surfaces, or random stimuli, for both motion and stereo cues. However, its responses to local edges or global surfaces were not different from random stimuli. This suggests that the LOC processes shapes, not edges or surfaces. LOC also responded more strongly to objects than to holes with the same shape, suggesting sensitivity to border ownership. V7 responded more strongly to edges than to surfaces or random stimuli for both motion and stereo cues, whereas V3a and V4 preferred motion edges. Finally, a region in the caudal intraparietal sulcus (cIPS) responded more strongly to both stereo versus motion and to stereo surfaces versus random stereo (but not to motion surfaces vs. random motion). Thus we found evidence for cue-specific responses to surfaces in the cIPS, both cue-specific and cue-independent responses to edges in intermediate visual areas, and shape-selective responses across multiple cues in the LOC. Overall, these data suggest that integration of visual information across multiple cues is mainly achieved at the level of shape and underscore LOC's role in shape computations.

    View details for DOI 10.1152/jn.01223.2007

    View details for Web of Science ID 000253969300030

    View details for PubMedID 18171705

  • Differential development of high-level visual cortex correlates with category-specific recognition memory NATURE NEUROSCIENCE Golarai, G., Ghahremani, D. G., Whitfield-Gabrieli, S., Reiss, A., Eberhardt, J. L., Gabrieli, J. D., Grill-Spector, K. 2007; 10 (4): 512-522


    High-level visual cortex in humans includes functionally defined regions that preferentially respond to objects, faces and places. It is unknown how these regions develop and whether their development relates to recognition memory. We used functional magnetic resonance imaging to examine the development of several functionally defined regions including object (lateral occipital complex, LOC)-, face ('fusiform face area', FFA; superior temporal sulcus, STS)- and place ('parahippocampal place area', PPA)-selective cortices in children (ages 7-11), adolescents (12-16) and adults. Right FFA and left PPA volumes were substantially larger in adults than in children. This development occurred by expansion of FFA and PPA into surrounding cortex and was correlated with improved recognition memory for faces and places, respectively. In contrast, LOC and STS volumes and object-recognition memory remained constant across ages. Thus, the ventral stream undergoes a prolonged maturation that varies temporally across functional regions, is determined by brain region rather than stimulus category, and is correlated with the development of category-specific recognition memory.

    View details for DOI 10.1038/nn1865

    View details for Web of Science ID 000245228600023

    View details for PubMedID 17351637

    View details for PubMedCentralID PMC3660101

  • Autism and the development of face processing 85th Annual Conference of the Association-for-Reseach-in-Nervous-and-Mental-Disease Golarai, G., Grill-Spector, K., Reiss, A. L. ELSEVIER SCI LTD. 2006: 145–60


    Autism is a pervasive developmental condition, characterized by impairments in non-verbal communication, social relationships and stereotypical patterns of behavior. A large body of evidence suggests that several aspects of face processing are impaired in autism, including anomalies in gaze processing, memory for facial identity and recognition of facial expressions of emotion. In search of neural markers of anomalous face processing in autism, much interest has focused on a network of brain regions that are implicated in social cognition and face processing. In this review, we will focus on three such regions, namely the STS for its role in processing gaze and facial movements, the FFA in face detection and identification and the amygdala in processing facial expressions of emotion. Much evidence suggests that a better understanding of the normal development of these specialized regions is essential for discovering the neural bases of face processing anomalies in autism. Thus, we will also examine the available literature on the normal development of face processing. Key unknowns in this research area are the neuro-developmental processes, the role of experience and the interactions among components of the face processing system in shaping each of the specialized regions for processing faces during normal development and in autism.

    View details for DOI 10.1016/j.cnr.2006.08.001

    View details for Web of Science ID 000242486200006

    View details for PubMedCentralID PMC2174902

  • Autism and the development of face processing. Clinical neuroscience research Golarai, G., Grill-Spector, K., Reiss, A. L. 2006; 6 (3): 145-160


    Autism is a pervasive developmental condition, characterized by impairments in non-verbal communication, social relationships and stereotypical patterns of behavior. A large body of evidence suggests that several aspects of face processing are impaired in autism, including anomalies in gaze processing, memory for facial identity and recognition of facial expressions of emotion. In search of neural markers of anomalous face processing in autism, much interest has focused on a network of brain regions that are implicated in social cognition and face processing. In this review, we will focus on three such regions, namely the STS for its role in processing gaze and facial movements, the FFA in face detection and identification and the amygdala in processing facial expressions of emotion. Much evidence suggests that a better understanding of the normal development of these specialized regions is essential for discovering the neural bases of face processing anomalies in autism. Thus, we will also examine the available literature on the normal development of face processing. Key unknowns in this research area are the neuro-developmental processes, the role of experience and the interactions among components of the face processing system in shaping each of the specialized regions for processing faces during normal development and in autism.

    View details for PubMedID 18176635

  • High-resolution imaging reveals highly selective nonface clusters in the fusiform face area NATURE NEUROSCIENCE Grill-Spector, K., Sayres, R., Ress, D. 2006; 9 (9): 1177-1185


    A region in ventral human cortex (fusiform face area, FFA) thought to be important for face perception responds strongly to faces and less strongly to nonface objects. This pattern of response may reflect a uniform face-selective neural population or activity averaged across populations with heterogeneous selectivity. Using high-resolution functional magnetic resonance imaging (MRI), we found that the FFA has a reliable heterogeneous structure: localized subregions within the FFA highly selective to faces are spatially interdigitated with localized subregions highly selective to different object categories. We found a preponderance of face-selective responses in the FFA, but no difference in selectivity to faces compared to nonfaces. Thus, standard fMRI of the FFA reflects averaging of heterogeneous highly selective neural populations of differing sizes, rather than higher selectivity to faces. These results suggest that visual processing in this region is not exclusive to faces. Overall, our approach provides a framework for understanding the fine-scale structure of neural representations in the human brain.

    View details for DOI 10.1038/nn1745

    View details for Web of Science ID 000240080800020

    View details for PubMedID 16892057

  • Object-selective cortex exhibits performance-independent repetition suppression JOURNAL OF NEUROPHYSIOLOGY Sayres, R., Grill-Spector, K. 2006; 95 (2): 995-1007


    Object-selective cortical regions exhibit a decreased response when an object stimulus is repeated [repetition suppression (RS)]. RS is often associated with priming: reduced response times and increased accuracy for repeated stimuli. It is unknown whether RS reflects stimulus-specific repetition, the associated changes in response time, or the combination of the two. To address this question, we performed a rapid event-related functional MRI (fMRI) study in which we measured BOLD signal in object-selective cortex, as well as object recognition performance, while we manipulated stimulus repetition. Our design allowed us to examine separately the roles of response time and repetition in explaining RS. We found that repetition played a robust role in explaining RS: repeated trials produced weaker BOLD responses than nonrepeated trials, even when comparing trials with matched response times. In contrast, response time played a weak role in explaining RS when repetition was controlled for: it explained BOLD responses only for one region of interest (ROI) and one experimental condition. Thus repetition suppression seems to be mostly driven by repetition rather than performance changes. We further examined whether RS reflects processes occurring at the same time as recognition or after recognition by manipulating stimulus presentation duration. In one experiment, durations were longer than required for recognition (2 s), whereas in a second experiment, durations were close to the minimum time required for recognition (85-101 ms). We found significant RS for brief presentations (albeit with a reduced magnitude), which again persisted when controlling for performance. This suggests a substantial amount of RS occurs during recognition.

    View details for DOI 10.1152/jn.00500.2005

    View details for Web of Science ID 000234759600040

    View details for PubMedID 16236787

  • Selectivity of adaptation in single units: Implications for fMRI experiments NEURON Grill-Spector, K. 2006; 49 (2): 170-171


    Understanding the neural basis of adaptation (repetition suppression) is critical for interpreting fMRI-adaptation experiments. Sawamura and colleagues provide a critical stepping-stone by elucidating the relation between neural adaptation and response selectivity. They find some cross-adaptation by two different stimuli that activate the same neuron.

    View details for DOI 10.1016/j.neuron.2006.01.004

    View details for Web of Science ID 000234979900003

    View details for PubMedID 16423690

  • Repetition and the brain: neural models of stimulus-specific effects TRENDS IN COGNITIVE SCIENCES Grill-Spector, K., Henson, R., Martin, A. 2006; 10 (1): 14-23


    One of the most robust experience-related cortical dynamics is reduced neural activity when stimuli are repeated. This reduction has been linked to performance improvements due to repetition and also used to probe functional characteristics of neural populations. However, the underlying neural mechanisms are as yet unknown. Here, we consider three models that have been proposed to account for repetition-related reductions in neural activity, and evaluate them in terms of their ability to account for the main properties of this phenomenon as measured with single-cell recordings and neuroimaging techniques. We also discuss future directions for distinguishing between these models, which will be important for understanding the neural consequences of repetition and for interpreting repetition-related effects in neuroimaging data.

    View details for DOI 10.1016/j.tics.2005.11.006

    View details for Web of Science ID 000234910400007

    View details for PubMedID 16321563

  • Visual recognition - As soon as you know it is there, you know what it is PSYCHOLOGICAL SCIENCE Grill-Spector, K., Kanwisher, N. 2005; 16 (2): 152-160


    What is the sequence of processing steps involved in visual object recognition? We varied the exposure duration of natural images and measured subjects' performance on three different tasks, each designed to tap a different candidate component process of object recognition. For each exposure duration, accuracy was lower and reaction time longer on a within-category identification task (e.g., distinguishing pigeons from other birds) than on a perceptual categorization task (e.g., birds vs. cars). However, strikingly, at each exposure duration, subjects performed just as quickly and accurately on the categorization task as they did on a task requiring only object detection: By the time subjects knew an image contained an object at all, they already knew its category. These findings place powerful constraints on theories of object recognition.

    View details for Web of Science ID 000226656000011

    View details for PubMedID 15686582

  • The fusiform face area subserves face perception, not generic within-category identification NATURE NEUROSCIENCE Grill-Spector, K., Knouf, N., Kanwisher, N. 2004; 7 (5): 555-562


    The function of the fusiform face area (FFA), a face-selective region in human extrastriate cortex, is a matter of active debate. Here we measured the correlation between FFA activity measured by functional magnetic resonance imaging (fMRI) and behavioral outcomes in perceptual tasks to determine the role of the FFA in the detection and within-category identification of faces and objects. Our data show that FFA activation is correlated on a trial-by-trial basis with both detecting the presence of faces and identifying specific faces. However, for most non-face objects (including cars seen by car experts), within-category identification performance was correlated with activation in other regions of the ventral occipitotemporal cortex, not the FFA. These results indicate that the FFA is involved in both detection and identification of faces, but that it has little involvement in within-category identification of non-face objects (including objects of expertise).

    View details for DOI 10.1038/nn1224

    View details for Web of Science ID 000221101300034

    View details for PubMedID 15077112

  • The human visual cortex ANNUAL REVIEW OF NEUROSCIENCE Grill-Spector, K., Malach, R. 2004; 27: 649-677


    The discovery and analysis of cortical visual areas is a major accomplishment of visual neuroscience. In the past decade the use of noninvasive functional imaging, particularly functional magnetic resonance imaging (fMRI), has dramatically increased our detailed knowledge of the functional organization of the human visual cortex and its relation to visual perception. The fMRI method offers a major advantage over other techniques applied in neuroscience by providing a large-scale neuroanatomical perspective that stems from its ability to image the entire brain essentially at once. This bird's eye view has the potential to reveal large-scale principles within the very complex plethora of visual areas. Thus, it could arrange the entire constellation of human visual areas in a unified functional organizational framework. Here we review recent findings and methods employed to uncover the functional properties of the human visual cortex focusing on two themes: functional specialization and hierarchical processing.

    View details for DOI 10.1146/annurev.neuro.27.070203.144220

    View details for Web of Science ID 000223246300023

    View details for PubMedID 15217346

  • The neural basis of object perception CURRENT OPINION IN NEUROBIOLOGY Grill-Spector, K. 2003; 13 (2): 159-166


    Humans can recognize an object within a fraction of a second, even if there are no clues about what kind of object it might be. Recent findings have identified functional properties of extrastriate regions in the ventral visual pathway that are involved in the representation and perception of objects and faces. The functional properties of these regions, and the correlation between the activation of these regions and visual recognition, indicate that the lateral and ventral occipito-temporal areas are important in perceiving and recognizing objects and faces.

    View details for DOI 10.1016/S0959-4388(03)00040-0

    View details for Web of Science ID 000183092700004

    View details for PubMedID 12744968

  • fMR-adaptation: a tool for studying the functional properties of human cortical neurons ACTA PSYCHOLOGICA Grill-Spector, K., Malach, R. 2001; 107 (1-3): 293-321


    The invariant properties of human cortical neurons cannot be studied directly by fMRI due to its limited spatial resolution. One voxel obtained from a fMRI scan contains several hundred thousands neurons. Therefore, the fMRI signal may average out a heterogeneous group of highly selective neurons. Here, we present a novel experimental paradigm for fMRI, functional magnetic resonance-adaptation (fMR-A), that enables to tag specific neuronal populations within an area and investigate their functional properties. This approach contrasts with conventional mapping methods that measure the averaged activity of a region. The application of fMR-A to study the functional properties of cortical neurons proceeds in two stages: First, the neuronal population is adapted by repeated presentation of a single stimulus. Second, some property of the stimulus is varied and the recovery from adaptation is assessed. If the signal remains adapted, it will indicate that the neurons are invariant to that attribute. However, if the fMRI signal will recover from the adapted state it would imply that the neurons are sensitive to the property that was varied. Here, an application of fMR-A for studying the invariant properties of high-order object areas (lateral occipital complex--LOC) to changes in object size, position, illumination and rotation is presented. The results show that LOC is less sensitive to changes in object size and position compared to changes of illumination and viewpoint. fMR-A can be extended to other neuronal systems in which adaptation is manifested and can be used with event-related paradigms as well. By manipulating experimental parameters and testing recovery from adaptation it should be possible to gain insight into the functional properties of cortical neurons which are beyond the spatial resolution limits imposed by conventional fMRI.

    View details for Web of Science ID 000168792700013

    View details for PubMedID 11388140

  • The lateral occipital complex and its role in object recognition VISION RESEARCH Grill-Spector, K., Kourtzi, Z., Kanwisher, N. 2001; 41 (10-11): 1409-1422


    Here we review recent findings that reveal the functional properties of extra-striate regions in the human visual cortex that are involved in the representation and perception of objects. We characterize both the invariant and non-invariant properties of these regions and we discuss the correlation between activation of these regions and recognition. Overall, these results indicate that the lateral occipital complex plays an important role in human object recognition.

    View details for Web of Science ID 000168565200016

    View details for PubMedID 11322983

  • The dynamics of object-selective activation correlate with recognition performance in humans NATURE NEUROSCIENCE Grill-Spector, K., Kushnir, T., Hendler, T., Malach, R. 2000; 3 (8): 837-843


    To investigate the relationship between perceptual awareness and brain activity, we measured both recognition performance and fMRI signal from object-related areas in human cortex while images were presented briefly using a masking protocol. Our results suggest that recognition performance is correlated with selective activation in object areas. Selective activation was correlated to object naming when exposure duration was varied from 20 to 500 milliseconds. Subjects' recognition during identical visual stimulation could be enhanced by training, which also increased the fMRI signal. Overall, the correlation between recognition performance and fMRI signal was highest in occipitotemporal object areas (the lateral occipital complex).

    View details for Web of Science ID 000167177300021

    View details for PubMedID 10903579

  • Differential processing of objects under various viewing conditions in the human lateral occipital complex NEURON Grill-Spector, K., Kushnir, T., Edelman, S., Avidan, G., Itzchak, Y., Malach, R. 1999; 24 (1): 187-203


    The invariant properties of human cortical neurons cannot be studied directly by fMRI due to its limited spatial resolution. Here, we circumvented this limitation by using fMR adaptation, namely, reduction of the fMR signal due to repeated presentation of identical images. Object-selective regions (lateral occipital complex [LOC]) showed a monotonic signal decrease as repetition frequency increased. The invariant properties of fMR adaptation were studied by presenting the same object in different viewing conditions. LOC exhibited stronger fMR adaptation to changes in size and position (more invariance) compared to illumination and viewpoint. The effect revealed two putative subdivisions within LOC: caudal-dorsal (LO), which exhibited substantial recovery from adaptation under all transformations, and posterior fusiform (PF/LOa), which displayed stronger adaptation. This study demonstrates the utility of fMR adaptation for revealing functional characteristics of neurons in fMRI studies.

    View details for Web of Science ID 000082893200019

    View details for PubMedID 10677037

  • Toward direct visualization of the internal shape representation space by fMRI PSYCHOBIOLOGY Edelman, S., Grill-Spector, K., Kushnir, T., Malach, R. 1998; 26 (4): 309-321
  • Cue-invariant activation in object-related areas of the human occipital lobe NEURON Grill-Spector, K., Kushnir, T., Edelman, S., Itzchak, Y., Malach, R. 1998; 21 (1): 191-202


    The extent to which primary visual cues such as motion or luminance are segregated in different cortical areas is a subject of controversy. To address this issue, we examined cortical activation in the human occipital lobe using functional magnetic resonance imaging (fMRI) while subjects performed a fixed visual task, object recognition, using three different primary visual cues: motion, texture, or luminance contrast. In the first experiment, a region located on the lateral aspect of the occipital lobe (LO complex) was preferentially activated in all 11 subjects both by luminance and motion-defined object silhouettes compared to full-field moving and stationary noise (ratios, 2.00+/-0.19 and 1.86+/-0.65, respectively). In the second experiment, all subjects showed enhanced activation in the LO complex to objects defined both by luminance and texture contrast compared to full-field texture patterns (ratios, 1.43+/-0.08 and 1.32+/-0.08, respectively). An additional smaller dorsal focus that exhibited convergence of object-related cues appeared to correspond to area V3a or a region slightly anterior to it. These results show convergence of visual cues in LO and provide strong evidence for its role in object processing.

    View details for Web of Science ID 000075061900018

    View details for PubMedID 9697863

  • A sequence of object-processing stages revealed by fMRI in the human occipital lobe HUMAN BRAIN MAPPING Grill-Spector, K., Kushnir, T., Hendler, T., Edelman, S., Itzchak, Y., Malach, R. 1998; 6 (4): 316-328


    Functional magnetic resonance imaging was used in combined functional selectivity and retinotopic mapping tests to reveal object-related visual areas in the human occipital lobe. Subjects were tested with right, left, up, or down hemivisual field stimuli which were composed of images of natural objects (faces, animals, man-made objects) or highly scrambled (1,024 elements) versions of the same images. In a similar fashion, the horizontal and vertical meridians were mapped to define the borders of these areas. Concurrently, the same cortical sites were tested for their sensitivity to image-scrambling by varying the number of scrambled picture fragments (from 16-1,024) while controlling for the Fourier power spectrum of the pictures and their order of presentation. Our results reveal a stagewise decrease in retinotopy and an increase in sensitivity to image-scrambling. Three main distinct foci were found in the human visual object recognition pathway (Ungerleider and Haxby [1994]: Curr Opin Neurobiol 4:157-165): 1) Retinotopic primary areas V1-3 did not exhibit significant reduction in activation to scrambled images. 2) Areas V4v (Sereno et al., [1995]: Science 268:889-893) and V3A (De Yoe et al., [1996]: Proc Natl Acad Sci USA 93:2382-2386; Tootell et al., [1997]: J Neurosci 71:7060-7078) manifested both retinotopy and decreased activation to highly scrambled images. 3) The essentially nonretinotopic lateral occipital complex (LO) (Malach et al., [1995]: Proc Natl Acad Sci USA 92:8135-8139; Tootell et al., [1996]: Trends Neurosci 19:481-489) exhibited the highest sensitivity to image scrambling, and appears to be homologous to macaque the infero-temporal (IT) cortex (Tanaka [1996]: Curr Opin Neurobiol 523-529). Breaking the images into 64, 256, or 1,024 randomly scrambled blocks reduced activation in LO voxels. However, many LO voxels remained significantly activated by mildly scrambled images (16 blocks). These results suggest the existence of object-fragment representation in LO.

    View details for Web of Science ID 000075169000009

    View details for PubMedID 9704268