Interdigitated Columnar Representation of Personal Space and Visual Space in Human Parietal Cortex.

Personal space (PS) is the space around the body that people prefer to maintain between themselves and unfamiliar others. Intrusion into personal space evokes discomfort and an urge to move away. Physiologic studies in nonhuman primates suggest that defensive responses to intruding stimuli involve the parietal cortex. We hypothesized that the spatial encoding of interpersonal distance is initially transformed from purely sensory to more egocentric mapping within human parietal cortex. This hypothesis was tested using 7 Tesla (7T) fMRI at high spatial resolution (1.1 mm isotropic), in seven subjects (four females, three males). In response to visual stimuli presented at a range of virtual distances, we found two categories of distance encoding in two corresponding radially-extending columns of activity within parietal cortex. One set of columns (P columns) responded selectively to moving and stationary face images presented at virtual distances that were nearer (but not farther) than each subject's behaviorally-defined personal space boundary. In most P columns, BOLD response amplitudes increased monotonically and nonlinearly with increasing virtual face proximity. In the remaining P columns, BOLD responses decreased with increasing proximity. A second set of parietal columns (D columns) responded selectively to disparity-based distance cues (near or far) in random dot stimuli, similar to disparity-selective columns described previously in occipital cortex. Critically, in parietal cortex, P columns were topographically interdigitated (nonoverlapping) with D columns. These results suggest that visual spatial information is transformed from visual to body-centered (or person-centered) dimensions in multiple local sites within human parietal cortex.SIGNIFICANCE STATEMENT Recent COVID-related social distancing practices highlight the need to better understand brain mechanisms which regulate "personal space" (PS), which is defined by the closest interpersonal distance that is comfortable for an individual. Using high spatial resolution brain imaging, we tested whether a map of external space is transformed from purely visual (3D-based) information to a more egocentric map (related to personal space) in human parietal cortex. We confirmed this transformation and further showed that it was mediated by two mutually segregated sets of columns: one which encoded interpersonal distance and another that encoded visual distance. These results suggest that the cortical transformation of sensory-centered to person-centered encoding of space near the body involves short-range communication across interdigitated columns within parietal cortex.

Scotopic Vision Is Selectively Processed in Thick-Type Columns in Human Extrastriate Cortex.

In humans, visual stimuli can be perceived across an enormous range of light levels. Evidence suggests that different neural mechanisms process different subdivisions of this range. For instance, in the retina, stimuli presented at very low (scotopic) light levels activate rod photoreceptors, whereas cone photoreceptors are activated relatively more at higher (photopic) light levels. Similarly, different retinal ganglion cells are activated by scotopic versus photopic stimuli. However, in the brain, it remains unknown whether scotopic versus photopic information is: 1) processed in distinct channels, or 2) neurally merged. Using high-resolution functional magnetic resonance imaging at 7 T, we confirmed the first hypothesis. We first localized thick versus thin-type columns within areas V2, V3, and V4, based on photopic selectivity to motion versus color, respectively. Next, we found that scotopic stimuli selectively activated thick- (compared to thin-) type columns in V2 and V3 (in measurements of both overlap and amplitude) and V4 (based on overlap). Finally, we found stronger resting-state functional connections between scotopically dominated area MT with thick- (compared to thin-) type columns in areas V2, V3, and V4. We conclude that scotopic stimuli are processed in partially segregated parallel streams, emphasizing magnocellular influence, from retina through middle stages of visual cortex.

Asymmetries in Global Perception Are Represented in Near- versus Far-Preferring Clusters in Human Visual Cortex.

Human perception is more "global" when stimuli are viewed within the lower (rather than the upper) visual field. This phenomenon is typically considered as a 2-D phenomenon, likely due to differential neural processing within dorsal versus ventral cortical areas that represent lower versus upper visual fields, respectively. Here we test a novel hypothesis that this vertical asymmetry in global processing is a 3-D phenomenon associated with (1) higher ecological relevance of low-spatial frequency (SF) components in encoding near (compared with far) visual objects and (2) the fact that near objects are more frequently found in lower rather than upper visual fields. Using high-resolution fMRI, collected within an ultra-high-field (7 T) scanner, we found that the extent of vertical asymmetry in global visual processing in human subjects (n = 10) was correlated with the fMRI response evoked by disparity-varying stimuli in human cortical area V3A. We also found that near-preferring clusters in V3A, located within stereoselective cortical columns, responded more selectively than far-preferring clusters, to low-SF features. These findings support the hypothesis that vertical asymmetry in global processing is a 3-D (not a 2-D) phenomenon, associated with the function of the stereoselective columns within visual cortex, especially those located within visual area V3A.SIGNIFICANCE STATEMENT Here we test and confirm a new hypothesis: fine-scale neural mechanisms underlying the vertical asymmetry in global visual processing. According to this hypothesis, the asymmetry in global visual processing is a 3-D (rather than a 2-D) phenomenon, reflected in the function of fine-scale cortical structures (clusters and columns) underlying depth perception. Our findings highlight the importance of considering these structures, as regions of interest, in clarifying the neural mechanisms underlying visual perception. The results also highlight the importance of statistics of natural scenes in shaping human visual perception.

Columnar Segregation of Magnocellular and Parvocellular Streams in Human Extrastriate Cortex.

Magnocellular versus parvocellular (M-P) streams are fundamental to the organization of macaque visual cortex. Segregated, paired M-P streams extend from retina through LGN into V1. The M stream extends further into area V5/MT, and parts of V2. However, elsewhere in visual cortex, it remains unclear whether M-P-derived information (1) becomes intermixed or (2) remains segregated in M-P-dominated columns and neurons. Here we tested whether M-P streams exist in extrastriate cortical columns, in 8 human subjects (4 female). We acquired high-resolution fMRI at high field (7T), testing for M- and P-influenced columns within each of four cortical areas (V2, V3, V3A, and V4), based on known functional distinctions in M-P streams in macaque: (1) color versus luminance, (2) binocular disparity, (3) luminance contrast sensitivity, (4) peak spatial frequency, and (5) color/spatial interactions. Additional measurements of resting state activity (eyes closed) tested for segregated functional connections between these columns. We found M- and P-like functions and connections within and between segregated cortical columns in V2, V3, and (in most experiments) area V4. Area V3A was dominated by the M stream, without significant influence from the P stream. These results suggest that M-P streams exist, and extend through, specific columns in early/middle stages of human extrastriate cortex.SIGNIFICANCE STATEMENT The magnocellular and parvocellular (M-P) streams are fundamental components of primate visual cortical organization. These streams segregate both anatomical and functional properties in parallel, from retina through primary visual cortex. However, in most higher-order cortical sites, it is unknown whether such M-P streams exist and/or what form those streams would take. Moreover, it is unknown whether M-P streams exist in human cortex. Here, fMRI evidence measured at high field (7T) and high resolution revealed segregated M-P streams in four areas of human extrastriate cortex. These results suggest that M-P information is processed in segregated parallel channels throughout much of human visual cortex; the M-P streams are more than a convenient sorting property in earlier stages of the visual system.

Columnar organization of mid-spectral and end-spectral hue preferences in human visual cortex.

Multiple color-selective areas have been described in visual cortex, in both humans and non-human primates. In macaques, hue-selective columns have been reported in several areas. In V2, it has been proposed that such hue-selective columns are mapped so as to mirror the order of wavelength through the visible spectrum, within thin-type stripes. Other studies have suggested a neural segregation of mid-spectral vs. end-spectral hue preferences (e.g. red and blue vs. green and yellow), within thin- and thick-type stripes, respectively. This latter segregation could reduce the spatial 'blur' due to chromatic aberration in the encoding of fine spatial details in the thick-type stripes. To distinguish between these and related models, we tested the organization of hue preferences in human visual cortex using fMRI at high spatial resolution. We used a high field (7T) scanner in humans (n = 7), measuring responses to four independent hues, including end-spectral (i.e. red-gray and blue-gray) and mid-spectral (i.e. green-gray and yellow-gray) isoluminant gratings, and also relative to achromatic luminance-varying (control) stimuli. In each subject, thin- and thick-type columns in V2 and V3 were localized using an independent set of stimuli and scans. We found distinct hue-selective differences along the dimension of mid-vs. end-spectral hues, in striate and early extrastriate visual cortex. First, as reported previously in macaques, V1 responded more strongly to end-spectral hues, compared to mid-spectral hues. Second, the color-selective thin-type stripes in V2 and V3 showed a greater response to end- and mid-spectral hues, relative to luminance-varying gratings. Third, thick-type stripes in V2/V3 showed a significantly stronger response to mid-spectral (compared to end-spectral) hues. Fourth, in the higher-tier color-selective area in occipital temporal cortex (n = 4), responses to all four hues were statistically equivalent to each other. These results suggest that early visual cortex segregates the processing of mid-vs. end-spectral hues, perhaps to counter the challenging optical constraint of chromatic aberration.

Visual field biases for near and far stimuli in disparity selective columns in human visual cortex.

When visual objects are located in the lower visual field, human observers perceive objects to be nearer than their real physical location. Conversely, objects in the upper visual field are viewed farther than their physical location. This bias may be linked to the statistics of natural scenes, and perhaps the ecological relevance of objects in the upper and lower visual fields (Previc, 1990; Yang and Purves, 2003). However, the neural mechanisms underlying such perceptual distortions have remained unknown. To test for underlying brain mechanisms, we presented visual stimuli at different perceptual distances, while measuring high-resolution fMRI in human subjects. First, we localized disparity-selective thick stripes and thick-type columns in secondary and third visual cortical areas, respectively. Consistent with the perceptual bias, we found that the thick stripe/columns that represent the lower visual field also responded more selectively to near rather than far visual stimuli. Conversely, thick stripe/columns that represent the upper visual field show a complementary bias, i.e. selectively higher activity to far rather than near stimuli. Thus, the statistics of natural scenes may play a significant role in the organization of near- and far-selective neurons within V2 thick stripes and V3 thick-type columns.

Interdigitated Color- and Disparity-Selective Columns within Human Visual Cortical Areas V2 and V3.

In nonhuman primates (NHPs), secondary visual cortex (V2) is composed of repeating columnar stripes, which are evident in histological variations of cytochrome oxidase (CO) levels. Distinctive "thin" and "thick" stripes of dark CO staining reportedly respond selectively to stimulus variations in color and binocular disparity, respectively. Here, we first tested whether similar color-selective or disparity-selective stripes exist in human V2. If so, available evidence predicts that such stripes should (1) radiate "outward" from the V1-V2 border, (2) interdigitate, (3) differ from each other in both thickness and length, (4) be spaced ∼3.5-4 mm apart (center-to-center), and, perhaps, (5) have segregated functional connections. Second, we tested whether analogous segregated columns exist in a "next-higher" tier area, V3. To answer these questions, we used high-resolution fMRI (1 × 1 × 1 mm(3)) at high field (7 T), presenting color-selective or disparity-selective stimuli, plus extensive signal averaging across multiple scan sessions and cortical surface-based analysis. All hypotheses were confirmed. V2 stripes and V3 columns were reliably localized in all subjects. The two stripe/column types were largely interdigitated (e.g., nonoverlapping) in both V2 and V3. Color-selective stripes differed from disparity-selective stripes in both width (thickness) and length. Analysis of resting-state functional connections (eyes closed) showed a stronger correlation between functionally alike (compared with functionally unlike) stripes/columns in V2 and V3. These results revealed a fine-scale segregation of color-selective or disparity-selective streams within human areas V2 and V3. Together with prior evidence from NHPs, this suggests that two parallel processing streams extend from visual subcortical regions through V1, V2, and V3. SIGNIFICANCE STATEMENT: In current textbooks and reviews, diagrams of cortical visual processing highlight two distinct neural-processing streams within the first and second cortical areas in monkeys. Two major streams consist of segregated cortical columns that are selectively activated by either color or ocular interactions. Because such cortical columns are so small, they were not revealed previously by conventional imaging techniques in humans. Here we demonstrate that such segregated columnar systems exist in humans. We find that, in humans, color versus binocular disparity columns extend one full area further, into the third visual area. Our approach can be extended to reveal and study additional types of columns in human cortex, perhaps including columns underlying more cognitive functions.

Curvature-processing network in macaque visual cortex.

Our visual environment abounds with curved features. Thus, the goal of understanding visual processing should include the processing of curved features. Using functional magnetic resonance imaging in behaving monkeys, we demonstrated a network of cortical areas selective for the processing of curved features. This network includes three distinct hierarchically organized regions within the ventral visual pathway: a posterior curvature-biased patch (PCP) located in the near-foveal representation of dorsal V4, a middle curvature-biased patch (MCP) located on the ventral lip of the posterior superior temporal sulcus (STS) in area TEO, and an anterior curvature-biased patch (ACP) located just below the STS in anterior area TE. Our results further indicate that the processing of curvature becomes increasingly complex from PCP to ACP. The proximity of the curvature-processing network to the well-known face-processing network suggests a possible functional link between them.

Increased Visual Stimulation Systematically Decreases Activity in Lateral Intermediate Cortex.

Previous studies have attributed multiple diverse roles to the posterior superior temporal cortex (STC), both visually driven and cognitive, including part of the default mode network (DMN). Here, we demonstrate a unifying property across this multimodal region. Specifically, the lateral intermediate (LIM) portion of STC showed an unexpected feature: a progressively decreasing fMRI response to increases in visual stimulus size (or number). Such responses are reversed in sign, relative to well-known responses in classic occipital temporal visual cortex. In LIM, this "reversed" size function was present across multiple object categories and retinotopic eccentricities. Moreover, we found a significant interaction between the LIM size function and the distribution of subjects' attention. These findings suggest that LIM serves as a part of the DMN. Further analysis of functional connectivity, plus a meta-analysis of previous fMRI results, suggests that LIM is a heterogeneous area including different subdivisions. Surprisingly, analogous fMRI tests in macaque monkeys did not reveal a clear homolog of LIM. This interspecies discrepancy supports the idea that self-referential thinking and theory of mind are more prominent in humans, compared with monkeys.

Thinking outside the box: rectilinear shapes selectively activate scene-selective cortex.

Fifteen years ago, an intriguing area was found in human visual cortex. This area (the parahippocampal place area [PPA]) was initially interpreted as responding selectively to images of places. However, subsequent studies reported that PPA also responds strongly to a much wider range of image categories, including inanimate objects, tools, spatial context, landmarks, objectively large objects, indoor scenes, and/or isolated buildings. Here, we hypothesized that PPA responds selectively to a lower-level stimulus property (rectilinear features), which are common to many of the above higher-order categories. Using a novel wavelet image filter, we first demonstrated that rectangular features are common in these diverse stimulus categories. Then we tested whether PPA is selectively activated by rectangular features in six independent fMRI experiments using progressively simplified stimuli, from complex real-world images, through 3D/2D computer-generated shapes, through simple line stimuli. We found that PPA was consistently activated by rectilinear features, compared with curved and nonrectangular features. This rectilinear preference was (1) comparable in amplitude and selectivity, relative to the preference for category (scenes vs faces), (2) independent of known biases for specific orientations and spatial frequency, and (3) not predictable from V1 activity. Two additional scene-responsive areas were sensitive to a subset of rectilinear features. Thus, rectilinear selectivity may serve as a crucial building block for category-selective responses in PPA and functionally related areas.

Neural correlates of personal space intrusion.

A parietal-frontal network in primates is thought to support many behaviors occurring in the space around the body, including interpersonal interactions and maintenance of a particular "comfort zone" or distance from other people ("personal space"). To better understand this network in humans, we used functional MRI to measure the responses to moving objects (faces, cars, simple spheres) and the functional connectivity of two regions in this network, the dorsal intraparietal sulcus (DIPS) and the ventral premotor cortex (PMv). We found that both areas responded more strongly to faces that were moving toward (vs away from) subjects, but did not show this bias in response to comparable motion in control stimuli (cars or spheres). Moreover, these two regions were functionally interconnected. Tests of activity-behavior associations revealed that the strength of DIPS-PMv connectivity was correlated with the preferred distance that subjects chose to stand from an unfamiliar person (personal space size). In addition, the magnitude of DIPS and PMv responses was correlated with the preferred level of social activity. Together, these findings suggest that this parietal-frontal network plays a role in everyday interactions with others.

Amygdala lesions disrupt modulation of functional MRI activity evoked by facial expression in the monkey inferior temporal cortex.

We previously showed that facial expressions modulate functional MRI activity in the face-processing regions of the macaque monkey's amygdala and inferior temporal (IT) cortex. Specifically, we showed that faces expressing emotion yield greater activation than neutral faces; we term this difference the "valence effect." We hypothesized that amygdala lesions would disrupt the valence effect by eliminating the modulatory feedback from the amygdala to the IT cortex. We compared the valence effects within the IT cortex in monkeys with excitotoxic amygdala lesions (n = 3) with those in intact control animals (n = 3) using contrast agent-based functional MRI at 3 T. Images of four distinct monkey facial expressions--neutral, aggressive (open mouth threat), fearful (fear grin), and appeasing (lip smack)--were presented to the subjects in a blocked design. Our results showed that in monkeys with amygdala lesions the valence effects were strongly disrupted within the IT cortex, whereas face responsivity (neutral faces > scrambled faces) and face selectivity (neutral faces > non-face objects) were unaffected. Furthermore, sparing of the anterior amygdala led to intact valence effects in the anterior IT cortex (which included the anterior face-selective regions), whereas sparing of the posterior amygdala led to intact valence effects in the posterior IT cortex (which included the posterior face-selective regions). Overall, our data demonstrate that the feedback projections from the amygdala to the IT cortex mediate the valence effect found there. Moreover, these modulatory effects are consistent with an anterior-to-posterior gradient of projections, as suggested by classical tracer studies.

Intrinsic structure of visual exemplar and category representations in macaque brain.

One of the most remarkable properties of the visual system is the ability to identify and categorize a wide variety of objects effortlessly. However, the underlying neural mechanisms remain elusive. Specifically, the question of how individual object information is represented and intrinsically organized is still poorly understood. To address this question, we presented images of isolated real-world objects spanning a wide range of categories to awake monkeys using a rapid event-related functional magnetic resonance imaging (fMRI) design and analyzed the responses of multiple areas involved in object processing. We found that the multivoxel response patterns to individual exemplars in the inferior temporal (IT) cortex, especially area TE, encoded the animate-inanimate categorical division, with a subordinate cluster of faces within the animate category. In contrast, the individual exemplar representations in V4, the amygdala, and prefrontal cortex showed either no categorical structure, or a categorical structure different from that in IT cortex. Moreover, in the IT face-selective regions ("face patches"), especially the anterior face patches, (1) the multivoxel response patterns to individual exemplars showed a categorical distinction between faces and nonface objects (i.e., body parts and inanimate objects), and (2) the regionally averaged activations to individual exemplars showed face-selectivity and within-face exemplar-selectivity. Our findings demonstrate that, at both the single-exemplar and the population level, intrinsic object representation and categorization are organized hierarchically as one moves anteriorly along the ventral pathway, reflecting both modular and distributed processing.

Retinotopy versus face selectivity in macaque visual cortex.

Retinotopic organization is a ubiquitous property of lower-tier visual cortical areas in human and nonhuman primates. In macaque visual cortex, the retinotopic maps extend to higher-order areas in the ventral visual pathway, including area TEO in the inferior temporal (IT) cortex. Distinct regions within IT cortex are also selective to specific object categories such as faces. Here we tested the topographic relationship between retinotopic maps and face-selective patches in macaque visual cortex using high-resolution fMRI and retinotopic face stimuli. Distinct subregions within face-selective patches showed either (1) a coarse retinotopic map of eccentricity and polar angle, (2) a retinotopic bias to a specific location of visual field, or (3) nonretinotopic selectivity. In general, regions along the lateral convexity of IT cortex showed more overlap between retinotopic maps and face selectivity, compared with regions within the STS. Thus, face patches in macaques can be subdivided into smaller patches with distinguishable retinotopic properties.

The "parahippocampal place area" responds preferentially to high spatial frequencies in humans and monkeys.

Defining the exact mechanisms by which the brain processes visual objects and scenes remains an unresolved challenge. Valuable clues to this process have emerged from the demonstration that clusters of neurons ("modules") in inferior temporal cortex apparently respond selectively to specific categories of visual stimuli, such as places/scenes. However, the higher-order "category-selective" response could also reflect specific lower-level spatial factors. Here we tested this idea in multiple functional MRI experiments, in humans and macaque monkeys, by systematically manipulating the spatial content of geometrical shapes and natural images. These tests revealed that visual spatial discontinuities (as reflected by an increased response to high spatial frequencies) selectively activate a well-known place-selective region of visual cortex (the "parahippocampal place area") in humans. In macaques, we demonstrate a homologous cortical area, and show that it also responds selectively to higher spatial frequencies. The parahippocampal place area may use such information for detecting object borders and scene details during spatial perception and navigation.

A previously undescribed scene-selective site is the key to encoding ego-motion in naturalistic environments.

Current models of scene processing in the human brain include three scene-selective areas: the Parahippocampal Place Area (or the temporal place areas; PPA/TPA), the restrosplenial cortex (or the medial place area; RSC/MPA) and the transverse occipital sulcus (or the occipital place area; TOS/OPA). Here, we challenged this model by showing that at least one other scene-selective site can also be detected within the human posterior intraparietal gyrus. Despite the smaller size of this site compared to the other scene-selective areas, the posterior intraparietal gyrus scene-selective (PIGS) site was detected consistently in a large pool of subjects (n=59; 33 females). The reproducibility of this finding was tested based on multiple criteria, including comparing the results across sessions, utilizing different scanners (3T and 7T) and stimulus sets. Furthermore, we found that this site (but not the other three scene-selective areas) is significantly sensitive to ego-motion in scenes, thus distinguishing the role of PIGS in scene perception relative to other scene-selective areas. These results highlight the importance of including finer scale scene-selective sites in models of scene processing - a crucial step toward a more comprehensive understanding of how scenes are encoded under dynamic conditions.

A previously undescribed scene-selective site is the key to encoding ego-motion in naturalistic environments.

Current models of scene processing in the human brain include three scene-selective areas: the parahippocampal place area (or the temporal place areas), the restrosplenial cortex (or the medial place area), and the transverse occipital sulcus (or the occipital place area). Here, we challenged this model by showing that at least one other scene-selective site can also be detected within the human posterior intraparietal gyrus. Despite the smaller size of this site compared to the other scene-selective areas, the posterior intraparietal gyrus scene-selective (PIGS) site was detected consistently in a large pool of subjects (n = 59; 33 females). The reproducibility of this finding was tested based on multiple criteria, including comparing the results across sessions, utilizing different scanners (3T and 7T) and stimulus sets. Furthermore, we found that this site (but not the other three scene-selective areas) is significantly sensitive to ego-motion in scenes, thus distinguishing the role of PIGS in scene perception relative to other scene-selective areas. These results highlight the importance of including finer scale scene-selective sites in models of scene processing - a crucial step toward a more comprehensive understanding of how scenes are encoded under dynamic conditions.

Decreased scene-selective activity within the posterior intraparietal cortex in amblyopic adults.

Amblyopia is a developmental disorder associated with reduced performance in visually guided tasks, including binocular navigation within natural environments. To help understand the underlying neurological disorder, we used fMRI to test the impact of amblyopia on the functional organization of scene-selective cortical areas, including the posterior intraparietal gyrus scene-selective (PIGS) area, a recently discovered region that responds selectively to ego-motion within naturalistic environments (Kennedy et al., 2024). Nineteen amblyopic adults (10 female) and thirty age-matched controls (12 female) participated in this study. Amblyopic participants spanned a wide range of amblyopia severity, based on their interocular visual acuity difference and stereoacuity. The visual function questionnaire (VFQ-39) was used to assess the participants' perception of their visual capabilities. Compared to controls, we found weaker scene-selective activity within the PIGS area in amblyopic individuals. By contrast, the level of scene-selective activity across the occipital place area (OPA), parahippocampal place area (PPA), and retrosplenial cortex (RSC)) remained comparable between amblyopic and control participants. The subjects' scores on "general vision" (VFQ-39 subscale) correlated with the level of scene-selective activity in PIGS. These results provide novel and direct evidence for amblyopia-related changes in scene-processing networks, thus enabling future studies to potentially link these changes across the spectrum of documented disabilities in amblyopia.

Using high-resolution functional MRI to differentiate impacts of strabismic and anisometropic amblyopia on evoked ocular dominance activity in humans.

We employed high-resolution functional MRI (fMRI) to distinguish the impacts of anisometropia and strabismus (the two most frequent causes of amblyopia) on the evoked ocular dominance (OD) response. Sixteen amblyopic participants (8 females), comprising 8 individuals with strabismus, 7 with anisometropia, 1 with deprivational amblyopia, along with 8 individuals with normal visual acuity (1 female), participated in this study for whom, we measured the difference between the response to stimulation of the two eyes, across early visual areas (V1-V4). In controls, as expected from the organization of OD columns, the evoked OD response formed a striped pattern that was mostly confined to V1. Compared to controls, the OD response in amblyopic participants formed larger fused patches that extended into downstream visual areas. Moreover, both anisometropic and strabismic participants showed stronger OD responses in V1, as well as in downstream visual areas V2-V4. Although this increase was most pronounced in V1, the correlation between the OD response level and the interocular visual acuity difference (measured behaviorally) was stronger in higher-level visual areas (V2-V4). Beyond these common effects, and despite similar densities of amblyopia between the anisometropic and strabismic participants, we found a greater increase in the size of V1 portion that responded preferentially to fellow eye stimulation in anisometropic compared to strabismic individuals. We also found a greater difference between the amplitudes of the response to binocular stimulation, in those regions that responded preferentially to the fellow vs. amblyopic eye, in anisometropic compared to strabismic subjects. In contrast, strabismic subjects demonstrated increased correlation between the OD responses evoked within V1 superficial and deep cortical depths, whereas anisometropic subjects did not. These results provide some of the first direct functional evidence for distinct impacts of strabismus and anisometropia on the mesoscale functional organization of the human visual system, thus extending what was inferred previously about amblyopia from animal models.

A cardinal orientation bias in scene-selective visual cortex.

It has long been known that human vision is more sensitive to contours at cardinal (horizontal and vertical) orientations, compared with oblique orientations; this is the "oblique effect." However, the real-world relevance of the oblique effect is not well understood. Experiments here suggest that this effect is linked to scene perception, via a common bias in the image statistics of scenes. This statistical bias for cardinal orientations is found in many "carpentered environments" such as buildings and indoor scenes, and some natural scenes. In Experiment 1, we confirmed the presence of a perceptual oblique effect in a specific set of scene stimuli. Using those scenes, we found that a well known "scene-selective" visual cortical area (the parahippocampal place area; PPA) showed distinctively higher functional magnetic resonance imaging (fMRI) activity to cardinal versus oblique orientations. This fMRI-based oblique effect was not observed in other cortical areas (including scene-selective areas transverse occipital sulcus and retrosplenial cortex), although all three scene-selective areas showed the expected inversion effect to scenes. Experiments 2 and 3 tested for an analogous selectivity for cardinal orientations using computer-generated arrays of simple squares and line segments, respectively. The results confirmed the preference for cardinal orientations in PPA, thus demonstrating that the oblique effect can also be produced in PPA by simple geometrical images, with statistics similar to those in scenes. Thus, PPA shows distinctive fMRI selectivity for cardinal orientations across a broad range of stimuli, which may reflect a perceptual oblique effect.