Functionally Separable Font-invariant and Font-sensitive Neural Populations in Occipitotemporal Cortex.

Reading relies on the rapid visual recognition of words viewed in a wide variety of fonts. We used fMRI to identify neural populations showing reduced fMRI responses to repeated words displayed in different fonts ("font-invariant" repetition suppression). We also identified neural populations showing greater fMRI responses to words repeated in a changing font as compared with words repeated in the same font ("font-sensitive" release from repetition suppression). We observed font-invariant repetition suppression in two anatomically distinct regions of the left occipitotemporal cortex (OT), a "visual word form area" in mid-fusiform cortex, and a more posterior region in the middle occipital gyrus. In contrast, bilateral shape-selective lateral occipital cortex and posterior fusiform showed considerable sensitivity to font changes during the viewing of repeated words. Although the visual word form area and the left middle occipital gyrus showed some evidence of font sensitivity, both regions showed a relatively greater degree of font invariance than font sensitivity. Our results show that the neural mechanisms in the left OT involved in font-invariant word recognition are anatomically distinct from those sensitive to font-related shape changes. We conclude that font-invariant representation of visual word form is instantiated at multiple levels by anatomically distinct neural mechanisms within the left OT.

Sex differences in the human visual system.

This Mini-Review summarizes a wide range of sex differences in the human visual system, with a primary focus on sex differences in visual perception and its neural basis. We highlight sex differences in both basic and high-level visual processing, with evidence from behavioral, neurophysiological, and neuroimaging studies. We argue that sex differences in human visual processing, no matter how small or subtle, support the view that females and males truly see the world differently. We acknowledge some of the controversy regarding sex differences in human vision and propose that such controversy should be interpreted as a source of motivation for continued efforts to assess the validity and reliability of published sex differences and for continued research on sex differences in human vision and the nervous system in general. © 2016 Wiley Periodicals, Inc.

Distinct effects of contour smoothness and observer bias on visual persistence.

Stable object perception relies on persistent yet temporary neural representations under constantly fluctuating stimulus conditions. The mechanisms by which such representations are formed and maintained are not fully understood but presumably involve interplay between early and higher tier visual cortical mechanisms. Some neurophysiological models of feature binding in early visual cortex predict persistent contour perception under certain stimulus conditions. Here we show that the duration of contour persistence reflects the persistent operation of visual mechanisms sensitive to contour smoothness, which also influences contour visibility more generally under highly camouflaging stimulus conditions. We distinguish the effect of contour smoothness on contour persistence from observer bias, which also contributes to the surprisingly long duration of contour persistence. We conclude that the strong modulatory effects of contour smoothness on persistence are due to the sustained reverberation of local and global contour-binding mechanisms in visual cortex, which form an important basis of perceptual continuity and stable object perception.

Visual Cortical Representation of Whole Words and Hemifield-split Word Parts.

Reading requires the neural integration of visual word form information that is split between our retinal hemifields. We examined multiple visual cortical areas involved in this process by measuring fMRI responses while observers viewed words that changed or repeated in one or both hemifields. We were specifically interested in identifying brain areas that exhibit decreased fMRI responses as a result of repeated versus changing visual word form information in each visual hemifield. Our method yielded highly significant effects of word repetition in a previously reported visual word form area (VWFA) in occipitotemporal cortex, which represents hemifield-split words as whole units. We also identified a more posterior occipital word form area (OWFA), which represents word form information in the right and left hemifields independently and is thus both functionally and anatomically distinct from the VWFA. Both the VWFA and the OWFA were left-lateralized in our study and strikingly symmetric in anatomical location relative to known face-selective visual cortical areas in the right hemisphere. Our findings are consistent with the observation that category-selective visual areas come in pairs and support the view that neural mechanisms in left visual cortex--especially those that evolved to support the visual processing of faces--are developmentally malleable and become incorporated into a left-lateralized visual word form network that supports rapid word recognition and reading.

Haptic shape processing in visual cortex.

Humans typically rely upon vision to identify object shape, but we can also recognize shape via touch (haptics). Our haptic shape recognition ability raises an intriguing question: To what extent do visual cortical shape recognition mechanisms support haptic object recognition? We addressed this question using a haptic fMRI repetition design, which allowed us to identify neuronal populations sensitive to the shape of objects that were touched but not seen. In addition to the expected shape-selective fMRI responses in dorsal frontoparietal areas, we observed widespread shape-selective responses in the ventral visual cortical pathway, including primary visual cortex. Our results indicate that shape processing via touch engages many of the same neural mechanisms as visual object recognition. The shape-specific repetition effects we observed in primary visual cortex show that visual sensory areas are engaged during the haptic exploration of object shape, even in the absence of concurrent shape-related visual input. Our results complement related findings in visually deprived individuals and highlight the fundamental role of the visual system in the processing of object shape.

Figure-ground representation and its decay in primary visual cortex.

We used fMRI to study figure-ground representation and its decay in primary visual cortex (V1). Human observers viewed a motion-defined figure that gradually became camouflaged by a cluttered background after it stopped moving. V1 showed positive fMRI responses corresponding to the moving figure and negative fMRI responses corresponding to the static background. This positive-negative delineation of V1 "figure" and "background" fMRI responses defined a retinotopically organized figure-ground representation that persisted after the figure stopped moving but eventually decayed. The temporal dynamics of V1 "figure" and "background" fMRI responses differed substantially. Positive "figure" responses continued to increase for several seconds after the figure stopped moving and remained elevated after the figure had disappeared. We propose that the sustained positive V1 "figure" fMRI responses reflected both persistent figure-ground representation and sustained attention to the location of the figure after its disappearance, as did subjects' reports of persistence. The decreasing "background" fMRI responses were relatively shorter-lived and less biased by spatial attention. Our results show that the transition from a vivid figure-ground percept to its disappearance corresponds to the concurrent decay of figure enhancement and background suppression in V1, both of which play a role in form-based perceptual memory.

Double representation of the wrist and elbow in human motor cortex.

Movements of the fingers, hand and arm involve overlapping neural representations in primary motor cortex (M1). Monkey M1 exhibits a core-surround organisation in which cortical representation of the hand and fingers is surrounded by representations of the wrist, elbow and shoulder. A potentially homologous organisation in human M1 has only been observed in a single study, a functional MRI (fMRI) study by [J.D. Meier, T.N. Aflalo, S. Kastner & M.S. Graziano.(2008) J Neurophysiol, 100(4), 1800-1812]. The results of their study suggested a double representation of the wrist in human M1, an unprecedented finding. Our purpose was to document and simultaneously provide evidence that would extend the presence of double representation of the wrist to that of the elbow. Using fMRI, we observed somatotopic maps in M1 and the supplementary motor area (SMA), the only other cortical area that showed robust within-limb somatotopy during self-timed finger, wrist and elbow movements. We observed double wrist and elbow representation that bracketed finger fMRI responses in M1 and the SMA. Our results show that the cortical locations of these double representations are well predicted by local cortical anatomy. Double representation of the wrist and elbow is important because it violates the traditional somatotopic progression in M1 but it is consistent with the representation of synergistic movements involving adjacent effectors.

Does face-selective cortex show a left visual field bias for centrally-viewed faces?

The left half of a centrally-viewed face contributes more strongly to recognition performance than the right. This left visual field (LVF) advantage is typically attributed to an untested assumption that face-selective cortex in the right hemisphere (RH) exhibits a contralateral bias, even for centrally-viewed faces. We tested the validity of this assumption using a behavioral measure of the LVF advantage and an fMRI experiment that measured laterality of face-selective cortex and neural contralateral bias. In the behavioral experiment, participants performed a chimeric face-matching task (Harrison and Strother, 2019). In the fMRI experiment, participants viewed chimeric faces comprised of face halves that either repeated or changed simultaneously in both hemifields, or repeated in one hemifield and changed in the other. This enabled us to measure lateralization of fMRI face-repetition suppression and hemifield-specific half-face sensitivity in face-selective cortex. We found that LVF bias in the fusiform face area (FFA) and right-lateralization of the FFA for changing versus repeated faces were both positively correlated with a behavioral measure of the LVF advantage for upright (but not inverted) faces. Results from regression analyses showed that LVF bias in the right FFA and FFA laterality make separable contributions to the prediction of our behavioral measure of the LVF bias for upright faces. Our results confirm a ubiquitous but previously untested assumption that RH superiority combined with contralateral bias in face-selective cortex explains the LVF advantage in face recognition. Specifically, our results show that neural LVF bias in the right FFA is sufficient to explain the relationship between FFA laterality and the perceptual LVF bias for centrally-viewed faces.

Equal degrees of object selectivity for upper and lower visual field stimuli.

Functional MRI (fMRI) studies of the human object recognition system commonly identify object-selective cortical regions by comparing blood oxygen level-dependent (BOLD) responses to objects versus those to scrambled objects. Object selectivity distinguishes human lateral occipital cortex (LO) from earlier visual areas. Recent studies suggest that, in addition to being object selective, LO is retinotopically organized; LO represents both object and location information. Although LO responses to objects have been shown to depend on location, it is not known whether responses to scrambled objects vary similarly. This is important because it would suggest that the degree of object selectivity in LO does not vary with retinal stimulus position. We used a conventional functional localizer to identify human visual area LO by comparing BOLD responses to objects versus scrambled objects presented to either the upper (UVF) or lower (LVF) visual field. In agreement with recent findings, we found evidence of position-dependent responses to objects. However, we observed the same degree of position dependence for scrambled objects and thus object selectivity did not differ for UVF and LVF stimuli. We conclude that, in terms of BOLD response, LO discriminates objects from non-objects equally well in either visual field location, despite stronger responses to objects in the LVF.

Subjective agency and awareness of shared actions.

Voluntary actions and their distal effects are intimately related in conscious awareness. When an expected effect follows a voluntary action, the experience of the interval between these events is compressed in time, a phenomenon known as 'intentional binding' (IB). Current accounts of IB suggest that it serves to reinforce associations between our goals and our intention to attain these goals via action, and that IB only occurs for self-generated (i.e., intentional) actions. We used a novel approach to study IB in the context of shared intentions and actions. Pairs of participants judged the time of occurrence of actions and events attributed either to oneself or to another agent. We found that IB and subjective agency are not mutually predictive when an action can be attributed to only one of two 'co-intending' agents. Our results pose a complication for the prevailing view that IB and subjective agency reflect a common mechanism.

Does right hemisphere superiority sufficiently explain the left visual field advantage in face recognition?

The tendency to perceive the identity of the left half of a centrally viewed face more strongly than that of the right half is associated with visual processing of faces in the right hemisphere (RH). Here we investigate conditions under which this well-known left visual field (LVF) half-face advantage fails to occur. Our findings challenge the sufficiency of its explanation as a function of RH specialization for face processing coupled with LVF-RH correspondence. In two experiments we show that the LVF half-face advantage occurs for normal faces and chimeric faces composed of different half-face identities. In a third experiment, we show that face inversion disrupts the LVF half-face advantage. In two additional experiments we show that half-faces viewed in isolation or paired with inverted half-faces fail to show the LVF advantage. Consistent with previous explanations of the LVF half-face advantage, our findings suggest that the LVF half-face advantage reflects RH superiority for processing faces and direct transfer of LVF face information to visual cortex in the RH. Critically, however, our findings also suggest the operation of a third factor, which involves the prioritization of face-processing resources to the LVF, but only when two upright face-halves compete for these resources. We therefore conclude that RH superiority alone does not suffice to explain the LVF advantage in face recognition. We also discuss the implications of our findings for specialized visual processing of faces by the right hemisphere, and we distinguish LVF advantages for faces viewed centrally and peripherally in divided field studies.

The conscious experience of action and intention.

The neural signals that give rise to our experience of the actions we perform are not well understood. Obhi et al. (Cognition 110:65-73, 2009) proposed that both efferent and re-afferent signals contribute to the conscious awareness of intentional actions. We further highlight the role of these signals in the awareness of our actions and the intention to perform these actions. We obtained temporal judgments of movement onset and the intention to execute finger and toe movements. This enabled us to compare our results with predictions corresponding to the conduction length of either effector. Our results confirm the findings of Obhi et al. (Cognition 110:65-73, 2009) that both efferent and re-afferent signals contribute to the awareness of planned actions and suggest that these signals may also play a role in our experience of our intention to perform an action.

Embedded word priming elicits enhanced fMRI responses in the visual word form area.

Lexical embedding is common in all languages and elicits mutual orthographic interference between an embedded word and its carrier. The neural basis of such interference remains unknown. We employed a novel fMRI prime-target embedded word paradigm to test for involvement of a visual word form area (VWFA) in left ventral occipitotemporal cortex in co-activation of embedded words and their carriers. Based on the results of related fMRI studies we predicted either enhancement or suppression of fMRI responses to embedded words initially viewed as primes, and repeated in the context of target carrier words. Our results clearly showed enhancement of fMRI responses in the VWFA to embedded-carrier word pairs as compared to unrelated prime-target pairs. In contrast to non-visual language-related areas (e.g., left inferior frontal gyrus), enhanced fMRI responses did not occur in the VWFA when embedded-carrier word pairs were restricted to the left visual hemifield. Our finding of fMRI enhancement in the VWFA is novel evidence of its involvement in representational rivalry between orthographically similar words, and the co-activation of embedded words and their carriers.

Visual recognition of mirrored letters and the right hemisphere advantage for mirror-invariant object recognition.

Unlike most objects, letter recognition is closely tied to orientation and mirroring, which in some cases (e.g., b and d), defines letter identity altogether. We combined a divided field paradigm with a negative priming procedure to examine the relationship between mirror generalization, its suppression during letter recognition, and language-related visual processing in the left hemisphere. In our main experiment, observers performed a centrally viewed letter-recognition task, followed by an object-recognition task performed in either the right or the left visual hemifield. The results show clear evidence of inhibition of mirror generalization for objects viewed in either hemifield but a right hemisphere advantage for visual recognition of mirrored and repeated objects. Our findings are consistent with an opponent relationship between symmetry-related visual processing in the right hemisphere and neurally recycled mechanisms in the left hemisphere used for visual processing of written language stimuli.

Object width modulates object-based attentional selection.

Visual input typically includes a myriad of objects, some of which are selected for further processing. While these objects vary in shape and size, most evidence supporting object-based guidance of attention is drawn from paradigms employing two identical objects. Importantly, object size is a readily perceived stimulus dimension, and whether it modulates the distribution of attention remains an open question. Across four experiments, the size of the objects in the display was manipulated in a modified version of the two-rectangle paradigm. In Experiment 1, two identical parallel rectangles of two sizes (thin or thick) were presented. Experiments 2-4 employed identical trapezoids (each having a thin and thick end), inverted in orientation. In the experiments, one end of an object was cued and participants performed either a T/L discrimination or a simple target-detection task. Combined results show that, in addition to the standard object-based attentional advantage, there was a further attentional benefit for processing information contained in the thick versus thin end of objects. Additionally, eye-tracking measures demonstrated increased saccade precision towards thick object ends, suggesting that Fitts's Law may play a role in object-based attentional shifts. Taken together, these results suggest that object-based attentional selection is modulated by object width.

An fMRI study of visual hemifield integration and cerebral lateralization.

The human brain integrates hemifield-split visual information via interhemispheric transfer. The degree to which neural circuits involved in this process behave differently during word recognition as compared to object recognition is not known. Evidence from neuroimaging (fMRI) suggests that interhemispheric transfer during word viewing converges in the left hemisphere, in two distinct brain areas, an "occipital word form area" (OWFA) and a more anterior occipitotemporal "visual word form area" (VWFA). We used a novel fMRI half-field repetition technique to test whether or not these areas also integrate nonverbal hemifield-split string stimuli of similar visual complexity. We found that the fMRI responses of both the OWFA and VWFA while viewing nonverbal stimuli were strikingly different than those measured during word viewing, especially with respect to half-stimulus changes restricted to a single hemifield. We conclude that normal reading relies on left-lateralized neural mechanisms, which integrate hemifield-split visual information for words but not for nonverbal stimuli.

On the surprising salience of curvature in grouping by proximity.

The authors conducted 3 experiments to explore the roles of curvature, density, and relative proximity in the perceptual organization of ambiguous dot patterns. To this end, they developed a new family of regular dot patterns that tend to be perceptually grouped into parallel contours, dot-sampled structured grids (DSGs). DSGs are similar to the dot lattices used to study grouping by proximity, except that only one of the potential organizations is rectilinear; the others are curvilinear. The authors used the method of M. Kubovy and J. Wagemans (1995) to study grouping by proximity in DSGs. They found that in the competition between the most likely organizations, one rectilinear and the other curvilinear, the latter is more salient. This phenomenon cannot be explained by contemporary accounts of grouping by proximity or contour integration.

Atypical Asymmetry for Processing Human and Robot Faces in Autism Revealed by fNIRS.

Deficits in the visual processing of faces in autism spectrum disorder (ASD) individuals may be due to atypical brain organization and function. Studies assessing asymmetric brain function in ASD individuals have suggested that facial processing, which is known to be lateralized in neurotypical (NT) individuals, may be less lateralized in ASD. Here we used functional near-infrared spectroscopy (fNIRS) to first test this theory by comparing patterns of lateralized brain activity in homologous temporal-occipital facial processing regions during observation of faces in an ASD group and an NT group. As expected, the ASD participants showed reduced right hemisphere asymmetry for human faces, compared to the NT participants. Based on recent behavioral reports suggesting that robots can facilitate increased verbal interaction over human counterparts in ASD, we also measured responses to faces of robots to determine if these patterns of activation were lateralized in each group. In this exploratory test, both groups showed similar asymmetry patterns for the robot faces. Our findings confirm existing literature suggesting reduced asymmetry for human faces in ASD and provide a preliminary foundation for future testing of how the use of categorically different social stimuli in the clinical setting may be beneficial in this population.

Spatiotemporal Form Integration: sequentially presented inducers can lead to representations of stationary and rigidly rotating objects.

Objects in the world often are occluded and in motion. The visible fragments of such objects are revealed at different times and locations in space. To form coherent representations of the surfaces of these objects, the visual system must integrate local form information over space and time. We introduce a new illusion in which a rigidly rotating square is perceived on the basis of sequentially presented Pacman inducers. The illusion highlights two fundamental processes that allow us to perceive objects whose form features are revealed over time: Spatiotemporal Form Integration (STFI) and Position Updating. STFI refers to the spatial integration of persistent representations of local form features across time. Position updating of these persistent form representations allows them to be integrated into a rigid global motion percept. We describe three psychophysical experiments designed to identify spatial and temporal constraints that underlie these two processes and a fourth experiment that extends these findings to more ecologically valid stimuli. Our results indicate that although STFI can occur across relatively long delays between successive inducers (i.e., greater than 500 ms), position updating is limited to a more restricted temporal window (i.e., ~300 ms or less), and to a confined range of spatial (mis)alignment. These findings lend insight into the limits of mechanisms underlying the visual system's capacity to integrate transient, piecemeal form information, and support coherent object representations in the ever-changing environment.