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The readings listed below are the foundation of this course. Where available, journal article abstracts from PubMed (an online database providing access to citations from biomedical literature) are included.

Course Textbook

Farah, M. The Cognitive Neuroscience of Vision. Blackwell Publishers, 2000.

Introduction to fMRI and Visual Cognition, Modularity
Gazzaniga, Ivry, and Mangun. Cognitive Neuroscience. Chap. 4. W. W. Norton & Company, 2002.
Functional Organization Of The Visual System
Farah, M. The Cognitive Neuroscience of Vision. Chap. 1 and 2. Blackwell Publishers, 2000.
Visual Recognition I: Theoretical / Behavioral Background on Shape Perception
Farah, M. The Cognitive Neuroscience of Vision. Chap. 2 and 3. Blackwell Publishers, 2000.
Visual Recognition II: Imaging Work on Shape Perception, Focusing especially on the Lateral Occipital Complex (LOC)
Grill-Spector, Kourtzi, and Kanwisher. Vision Research 41 (2001): 1409-1422.

Malach, R., I. Levy, and U. Hasson. "The Topography of High Order Human Object Areas." Trends Cogn Sci. 6 (2002): 176-184.

PubMed abstract:  Cortical topography is one of the most fundamental organizing principles of cortical areas. One such topography - eccentricity mapping - is present even in high-order, ventral stream visual areas. Within these areas, different object categories have specific eccentricity biases. In particular, faces, letters and words appear to be associated with central visual-field bias, whereas buildings are associated with a peripheral one. We propose that resolution needs are an important factor in organizing object representations: objects whose recognition depends on analysis of fine detail will be associated with central-biased representations, whereas objects whose recognition entails large-scale integration will be more peripherally biased.

And at least one of these:

James, T., G. Humphrey, J. Gati, R. Menon, and M. Goodale. "Differential Effects of Viewpoint on Object-driven Activation in Dorsal and Ventral Streams." Neuron 35 (2002): 793-801.

PubMed abstract:  Using fMRI, we showed that an area in the ventral temporo-occipital cortex (area vTO), which is part of the human homolog of the ventral stream of visual processing, exhibited priming for both identical and depth-rotated images of objects. This pattern of activation in area vTO corresponded to performance in a behavioral matching task. An area in the caudal part of the intraparietal sulcus (area cIPS) also showed priming, but only with identical images of objects. This dorsal-stream area treated rotated images as new objects. The difference in the pattern of priming-related activation in the two areas may reflect the respective roles of the ventral and dorsal streams in object recognition and object-directed action.

Vuilleumier, P., R. N. Henson, J. Driver, and R. J. Dolan. "Multiple Levels of Visual Object Constancy Revealed by Event-related fMRI of Repetition Priming." Nat Neurosci. 5 (2002): 491-9.

PubMed abstract:  We conducted two event-related functional magnetic resonance imaging (fMRI) experiments to investigate the neural substrates of visual object recognition in humans. We used a repetition-priming method with visual stimuli recurring at unpredictable intervals, either with the same appearance or with changes in size, viewpoint or exemplar. Lateral occipital and posterior inferior temporal cortex showed lower activity for repetitions of both real and non-sense objects; fusiform and left inferior frontal regions showed decreases for repetitions of only real objects. Repetition of different exemplars with the same name affected only the left inferior frontal cortex. Crucially, priming-induced decreases in activity of the right fusiform cortex depended on whether the three-dimensional objects were repeated with the same viewpoint, regardless of whether retinal image size changed; left fusiform decreases were independent of both viewpoint and size. These data show that dissociable subsystems in ventral visual cortex maintain distinct view-dependent and view-invariant object representations.

Yin, C., S. Shimojo, C. Moore, and S. Engel. "Dynamic Shape Integration in Extrastriate Cortex." Curr Biol. 20 (2002): 1379-1385.

PubMed abstract:  BACKGROUND: In anorthoscopic viewing conditions, observers can perceive a moving object through a narrow slit even when only portions of its contour are visible at any time. We used fMRI to examine the contribution of early and later visual cortical areas to dynamic shape integration. Observers' success at integrating the shape of the slit-viewed object was manipulated by varying the degree to which the stimulus was dynamically distorted. Line drawings of common objects were either moderately distorted, strongly distorted, or shown undistorted. Phenomenologically, increasing the stimulus distortion made both object shape and motion more difficult to perceive.RESULTS: We found that bilateral cortical activity in portions of the ventral occipital cortex, corresponding to known object areas within the lateral occipital complex (LOC), was inversely correlated with the degree of stimulus distortion. We found that activity in left MT+, the human cortical area specialized for motion, showed a similar pattern as the ventral occipital region. The LOC also showed greater activity to a fully visible moving object than to the undistorted slit-viewed object. Area MT+, however, showed more equivalent activity to both the slit-viewed and fully visible moving objects.CONCLUSIONS: In early retinotopic cortex, the distorted and undistorted stimuli elicited the same amount of activity. Higher visual areas, however, were correlated with the percept of the coherent object, and this correlation suggests that the shape integration is mediated by later visual cortical areas. Motion information from the dorsal stream may project to the LOC to produce the shape percept.

Visual Recognition III: Behavioral / Theoretical Background on Face Recognition and Category Specificity
Farah, M. The Cognitive Neuroscience of Vision. Chap. 5. Blackwell Publishers, 2000.
Visual Recognition IV: Imaging Work on Category Specificity
Chao, L. L., J. V. Haxby, and A. Martin. "Attribute-based Neural Substrates in Temporal Cortex for Perceiving and Knowing About Objects." Nat Neurosci. 2 (1999): 913-9.

PubMed abstract:  The cognitive and neural mechanisms underlying category-specific knowledge remain controversial. Here we report that, across multiple tasks (viewing, delayed match to sample, naming), pictures of animals and tools were associated with highly consistent, category-related patterns of activation in ventral (fusiform gyrus) and lateral (superior and middle temporal gyri) regions of the posterior temporal lobes. In addition, similar patterns of category-related activity occurred when subjects read the names of, and answered questions about, animals and tools. These findings suggest that semantic object information is represented in distributed networks that include sites for storing information about specific object attributes such as form (ventral temporal cortex) and motion (lateral temporal cortex).

Downing, P. E., Y. Jiang, M. Shuman, and N. Kanwisher. "A Cortical Area Selective for Visual Processing of the Human Body." Science 293 (2001): 2470-3.

PubMed abstract:  Despite extensive evidence for regions of human visual cortex that respond selectively to faces, few studies have considered the cortical representation of the appearance of the rest of the human body. We present a series of functional magnetic resonance imaging (fMRI) studies revealing substantial evidence for a distinct cortical region in humans that responds selectively to images of the human body, as compared with a wide range of control stimuli. This region was found in the lateral occipitotemporal cortex in all subjects tested and apparently reflects a specialized neural system for the visual perception of the human body.

Kanwisher. "The Ventral Visual Object Pathway in Humans: Evidence from fMRI." The Visual Neurosciences. Edited by Chalupa, and Werner. (In press)

Pierce, K., R. A. Muller, J. Ambrose, G. Allen, and E. Courchesne. "Face Processing Occurs Outside the Fusiform 'face area' in Autism: Evidence from Functional MRI." Brain 124 (2001): 2059-73.

PubMed abstract:  Processing the human face is at the focal point of most social interactions, yet this simple perceptual task is difficult for individuals with autism, a population that spends limited amounts of time engaged in face-to-face eye contact or social interactions in general. Thus, the study of face processing in autism is not only important because it may be integral to understanding the social deficits of this disorder, but also, because it provides a unique opportunity to study experiential factors related to the functional specialization of normal face processing. In short, autism may be one of the only disorders where affected individuals spend reduced amounts of time engaged in face processing from birth. Using functional MRI, haemodynamic responses during a face perception task were compared between adults with autism and normal control subjects. Four regions of interest (ROIs), the fusiform gyrus (FG), inferior temporal gyrus, middle temporal gyrus and amygdala were manually traced on non-spatially normalized images and the percentage ROI active was calculated for each subject. Analyses in Talairach space were also performed. Overall results revealed either abnormally weak or no activation in FG in autistic patients, as well as significantly reduced activation in the inferior occipital gyrus, superior temporal sulcus and amygdala. Anatomical abnormalities, in contrast, were present only in the amygdala in autistic patients, whose mean volume was significantly reduced as compared with normals. Reaction time and accuracy measures were not different between groups. Thus, while autistic subjects could perform the face perception task, none of the regions supporting face processing in normals were found to be significantly active in the autistic subjects. Instead, in every autistic patient, faces maximally activated aberrant and individual-specific neural sites (e.g. frontal cortex, primary visual cortex, etc.), which was in contrast to the 100% consistency of maximal activation within the traditional fusiform face area (FFA) for every normal subject. It appears that, as compared with normal individuals, autistic individuals 'see' faces utilizing different neural systems, with each patient doing so via a unique neural circuitry. Such a pattern of individual-specific, scattered activation seen in autistic patients in contrast to the highly consistent FG activation seen in normals, suggests that experiential factors do indeed play a role in the normal development of the FFA.


Haxby, J. V., M. I. Gobbini, M. L. Furey, A. Ishai, J. L. Schouten, and P. Pietrini. "Distributed and Overlapping Representations of Faces and Objects in Ventral Temporal Cortex." Science 293 (2001): 2425-2430.

PubMed abstract:  The functional architecture of the object vision pathway in the human brain was investigated using functional magnetic resonance imaging to measure patterns of response in ventral temporal cortex while subjects viewed faces, cats, five categories of man-made objects, and nonsense pictures. A distinct pattern of response was found for each stimulus category. The distinctiveness of the response to a given category was not due simply to the regions that responded maximally to that category, because the category being viewed also could be identified on the basis of the pattern of response when those regions were excluded from the analysis. Patterns of response that discriminated among all categories were found even within cortical regions that responded maximally to only one category. These results indicate that the representations of faces and objects in ventral temporal cortex are widely distributed and overlapping.

Visual Attention I: Theory
Driver, J. "A Selective Review of Selective Attention Research from the Past Century." Br J Psychol. 92 Part 1 (2001): 53-78.

PubMed abstract:    Research on attention is concerned with selective processing of incoming sensory information. To some extent, our awareness of the world depends on what we choose to attend, not merely on the stimulation entering our senses. British psychologists have made substantial contributions to this topic in the past century. Celebrated examples include Donald Broadbent's filter theory of attention, which set the agenda for most subsequent work; and Anne Treisman's revisions of this account, and her later feature-integration theory. More recent contributions include Alan Allport's prescient emphasis on the relevance of neuroscience data, and John Duncan's integration of such data with psychological theory. An idiosyncratic but roughly chronological review of developments is presented, some practical and clinical implications are briefly sketched, and future directions suggested. One of the biggest changes in the field has been the increasing interplay between psychology and neuroscience, which promises much for the future. A related change has been the realization that selection attention is best thought of as a broad topic, encompassing a range of selective issues, rather than as a single explanatory process.

Farah, M. The Cognitive Neuroscience of Vision. Chap. 7 and 8. Blackwell Publishers, 2000.

Visual Attention II: Imaging 
Kanwisher, N., and E. Wojciulik. "Visual Attention: Insights from Brain Imaging." Nature Reviews Neuroscience 1 (2000): 91-100.

PubMed abstract:  We are not passive recipients of the information that impinges on our retinae, but active participants in our own perceptual processes. Visual experience depends critically on attention. We select particular aspects of a visual scene for detailed analysis and control of subsequent behaviour, but ignore other aspects so completely that moments after they disappear from view we cannot report anything about them. Here we show that functional neuroimaging is revealing much more than where attention happens in the brain; it is beginning to answer some of the oldest and deepest questions about what visual attention is and how it works.

And at least one of these:

Corbetta, M., and G. L. Shulman. "Control of Goal-directed and Stimulus-driven Attention in the Brain." Nat Rev Neurosci. 3 (2002): 201-15.

PubMed abstract:  We review evidence for partially segregated networks of brain areas that carry out different attentional functions. One system, which includes parts of the intraparietal cortex and superior frontal cortex, is involved in preparing and applying goal-directed (top-down) selection for stimuli and responses. This system is also modulated by the detection of stimuli. The other system, which includes the temporoparietal cortex and inferior frontal cortex, and is largely lateralized to the right hemisphere, is not involved in top-down selection. Instead, this system is specialized for the detection of behaviourally relevant stimuli, particularly when they are salient or unexpected. This ventral frontoparietal network works as a 'circuit breaker' for the dorsal system, directing attention to salient events. Both attentional systems interact during normal vision, and both are disrupted in unilateral spatial neglect.

Noesselt, T., S. Hillyard, M. Woldorff, A. Schoenfeld, T. Hagner, L. Jancke, C . Tempelmann, H. Hinrichs, and H. Heinze. "Delayed Striate Cortical Activation during Spatial Attention." Neuron 35 (2002): 575.

PubMed abstract:  Recordings of event-related potentials (ERPs) and event-related magnetic fields (ERMFs) were combined with functional magnetic resonance imaging (fMRI) to study visual cortical activity in humans during spatial attention. While subjects attended selectively to stimulus arrays in one visual field, fMRI revealed stimulus-related activations in the contralateral primary visual cortex and in multiple extrastriate areas. ERP and ERMF recordings showed that attention did not affect the initial evoked response at 60-90 ms poststimulus that was localized to primary cortex, but a similarly localized late response at 140-250 ms was enhanced to attended stimuli. These findings provide evidence that the primary visual cortex participates in the selective processing of attended stimuli by means of delayed feedback from higher visual-cortical areas.

Shafritz, K. M., J. C. Gore, and R. Marois. "The Role of the Parietal Cortex in Visual Feature Binding." Proc Natl Acad Sci USA 99 (2002): 10917-22.

PubMed abstract:  When multiple objects are simultaneously present in a scene, the visual system must properly integrate the features associated with each object. It has been proposed that this "binding problem" is solved by selective attention to the locations of the objects [Treisman, A.M. & Gelade, E. (1980) Cogn. Psychol. 12, 97-136]. If spatial attention plays a role in feature integration, it should do so primarily when object location can serve as a binding cue. Using functional MRI (fMRI), we show that regions of the parietal cortex involved in spatial attention are more engaged in feature conjunction tasks than in single feature tasks when multiple objects are shown simultaneously at different locations but not when they are shown sequentially at the same location. These findings suggest that the spatial attention network of the parietal cortex is involved in feature binding but only when spatial information is available to resolve ambiguities about the relationships between object features.

Saenz, M., G. T. Buracas, and G. M. Boynton. "Global Effects of Feature-based Attention in Human Visual Cortex." Nat Neurosci. 5 (2002): 631-2.

PubMed abstract:  The content of visual experience depends on how selective attention is distributed in the visual field. We used functional magnetic resonance imaging (fMRI) in humans to test whether feature-based attention can globally influence visual cortical responses to stimuli outside the attended location. Attention to a stimulus feature (color or direction of motion) increased the response of cortical visual areas to a spatially distant, ignored stimulus that shared the same feature.

Visual Awareness
Dehaene, S., L. Naccache, L. Cohen, D. L. Bihan, J. F. Mangin, J. B. Poline, and D. Riviere. "Cerebral Mechanisms of Word Masking and Unconscious Repetition Priming." Nat Neurosci. 4 (2001): 752-8.

PubMed abstract:  We used functional magnetic resonance imaging (fMRI) and event-related potentials (ERPs) to visualize the cerebral processing of unseen masked words. Within the areas associated with conscious reading, masked words activated left extrastriate, fusiform and precentral areas. Furthermore, masked words reduced the amount of activation evoked by a subsequent conscious presentation of the same word. In the left fusiform gyrus, this repetition suppression phenomenon was independent of whether the prime and target shared the same case, indicating that case-independent information about letter strings was extracted unconsciously. In comparison to an unmasked situation, however, the activation evoked by masked words was drastically reduced and was undetectable in prefrontal and parietal areas, correlating with participants' inability to report the masked words.

Farah, M. The Cognitive Neuroscience of Vision. Chap. 10. Blackwell Publishers, 2000.

Kanwisher, N. "Neural Events and Perceptual Awareness." Cognition. 79 (2001): 89-113.

PubMed abstract:  Neural correlates of perceptual awareness, until very recently an elusive quarry, are now almost commonplace findings. This article first describes a variety of neural correlates of perceptual awareness based on fMRI, ERPs, and single-unit recordings. It is then argued that our quest should ultimately focus not on mere correlates of awareness, but rather on the neural events that are both necessary and sufficient for perceptual awareness. Indeed, preliminary evidence suggests that although many of the neural correlates already reported may be necessary for the corresponding state of awareness, it is unlikely that they are sufficient for it. The final section considers three hypotheses concerning the possible sufficiency conditions for perceptual awareness.

Moutoussis, and Zeki. "The Relationship between Cortical Activation and Perception Investigated with Invisible Stimuli." PNAS. 99 (2002): 9527-9532.

PubMed abstract:  The aim of this work was to study the relationship between cortical activity and visual perception. To do so, we developed a psychophysical technique that is able to dissociate the visual percept from the visual stimulus and thus distinguish brain activity reflecting the perceptual state from that reflecting other stages of stimulus processing. We used dichoptic color fusion to make identical monocular stimuli of opposite color contrast "disappear" at the binocular level and thus become "invisible" as far as conscious visual perception is concerned. By imaging brain activity in subjects during a discrimination task between face and house stimuli presented in this way, we found that house-specific and face-specific brain areas are always activated in a stimulus-specific way regardless of whether the stimuli are perceived. Absolute levels of cortical activation, however, were lower with invisible stimulation compared with visible stimulation. We conclude that there is no terminal "perceptual" area in the visual brain, but that the brain regions involved in processing a visual stimulus are also involved in its perception, the difference between the two being dictated by a higher level of activity in the specific brain region when the stimulus is perceived.


Baars. "The Conscious Access Hypothesis: Origins and Recent Evidence." Trends Cogn Sci. 6 (2002): 47-52.

PubMed abstract:  Consciousness might help to mobilize and integrate brain functions that are otherwise separate and independent. Evidence for this 'conscious access hypothesis' was described almost two decades ago, in a framework called global workspace theory. The theory had little impact at first, for three reasons: because consciousness was controversial; the evidence, though extensive, was indirect; and integrative theory was unfashionable. Recent neuroimaging evidence appears broadly to support the hypothesis, which has implications for perception, learning, working memory, voluntary control, attention and self systems in the brain.

Visual Working Memory and Imagery
Farah, M. The Cognitive Neuroscience of Vision. Chap. 9. Blackwell Publishers, 2000.
Monkey fMRI
Bandettini, P. A., and L. G. Ungerleider. "From Neuron to BOLD: New Connections." Nat Neurosci. 4 (2001): 864-6.