Academic Appointments

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.

Stanford Advisees

All Publications

  • 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

  • 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 Web of Science ID 000348295100020

    View details for PubMedID 25569351

  • 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
  • Where Is Human V4? Predicting the Location of hV4 and VO1 from Cortical Folding CEREBRAL CORTEX Witthoft, N., Mai Lin 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 Web of Science ID 000343407500012

  • 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 Web of Science ID 000339392400009

  • 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

  • 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

  • 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

  • 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

  • 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

  • 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

  • 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

  • Controlling stimulus variability reveals stronger face-selective responses near the average face VISUAL COGNITION Davidenko, N., Grill-Spector, K. 2010; 18 (1): 122-126
  • 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

  • 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

  • Autism and the development of face processing CLINICAL NEUROSCIENCE RESEARCH Golarai, G., Grill-Spector, K., Reiss, A. L. 2006; 6 (3-4): 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 DOI 10.1016/j.cnr.2006.08.001

    View details for Web of Science ID 000242486200006

  • 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