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.

The Global Configuration of Visual Stimuli Alters Co-Fluctuations of Cross-Hemispheric Human Brain Activity.

We tested how a stimulus gestalt, defined by the neuronal interaction between local and global features of a stimulus, is represented within human primary visual cortex (V1). We used high-resolution fMRI, which serves as a surrogate of neuronal activation, to measure co-fluctuations within subregions of V1 as (male and female) subjects were presented with peripheral stimuli, each with different global configurations. We found stronger cross-hemisphere correlations when fine-scale V1 cortical subregions represented parts of the same object compared with different objects. This result was consistent with the vertical bias in global processing and, critically, was independent of the task and local discontinuities within objects. Thus, despite the relatively small receptive fields of neurons within V1, global stimulus configuration affects neuronal processing via correlated fluctuations between regions that represent different sectors of the visual field.SIGNIFICANCE STATEMENT We provide the first evidence for the impact of global stimulus configuration on cross-hemispheric fMRI fluctuations, measured in human primary visual cortex. Our results are consistent with changes in the level of γ-band synchrony, which has been shown to be affected by global stimulus configuration, being reflected in the level fMRI co-fluctuations. These data help narrow the gap between knowledge of global stimulus configuration encoding at the single-neuron level versus at the behavioral level.

Critical factors in achieving fine-scale functional MRI: Removing sources of inadvertent spatial smoothing.

Ultra-high Field (≥7T) functional magnetic resonance imaging (UHF-fMRI) provides opportunities to resolve fine-scale features of functional architecture such as cerebral cortical columns and layers, in vivo. While the nominal resolution of modern fMRI acquisitions may appear to be sufficient to resolve these features, several common data preprocessing steps can introduce unwanted spatial blurring, especially those that require interpolation of the data. These resolution losses can impede the detection of the fine-scale features of interest. To examine quantitatively and systematically the sources of spatial resolution losses occurring during preprocessing, we used synthetic fMRI data and real fMRI data from the human visual cortex-the spatially interdigitated human V2 "thin" and "thick" stripes. The pattern of these cortical columns lies along the cortical surface and thus can be best appreciated using surface-based fMRI analysis. We used this as a testbed for evaluating strategies that can reduce spatial blurring of fMRI data. Our results show that resolution losses can be mitigated at multiple points in preprocessing pathway. We show that unwanted blur is introduced at each step of volume transformation and surface projection, and can be ameliorated by replacing multi-step transformations with equivalent single-step transformations. Surprisingly, the simple approaches of volume upsampling and of cortical mesh refinement also helped to reduce resolution losses caused by interpolation. Volume upsampling also serves to improve motion estimation accuracy, which helps to reduce blur. Moreover, we demonstrate that the level of spatial blurring is nonuniform over the brain-knowledge which is critical for interpreting data in high-resolution fMRI studies. Importantly, our study provides recommendations for reducing unwanted blurring during preprocessing as well as methods that enable quantitative comparisons between preprocessing strategies. These findings highlight several underappreciated sources of a spatial blur. Individually, the factors that contribute to spatial blur may appear to be minor, but in combination, the cumulative effects can hinder the interpretation of fine-scale fMRI and the detectability of these fine-scale features of functional architecture.

A 31-channel integrated "AC/DC" B0 shim and radiofrequency receive array coil for improved 7T MRI.

PURPOSE: To test an integrated "AC/DC" array approach at 7T, where B0 inhomogeneity poses an obstacle for functional imaging, diffusion-weighted MRI, MR spectroscopy, and other applications. METHODS: A close-fitting 7T 31-channel (31-ch) brain array was constructed and tested using combined Rx and ΔB0 shim channels driven by a set of rapidly switchable current amplifiers. The coil was compared to a shape-matched 31-ch reference receive-only array for RF safety, signal-to-noise ratio (SNR), and inter-element noise correlation. We characterize the coil array's ability to provide global and dynamic (slice-optimized) shimming using ΔB0 field maps and echo planar imaging (EPI) acquisitions. RESULTS: The SNR and average noise correlation were similar to the 31-ch reference array. Global and slice-optimized shimming provide 11% and 40% improvements respectively compared to baseline second-order spherical harmonic shimming. Birdcage transmit coil efficiency was similar for the reference and AC/DC array setups. CONCLUSION: Adding ΔB0 shim capability to a 31-ch 7T receive array can significantly boost 7T brain B0 homogeneity without sacrificing the array's rdiofrequency performance, potentially improving ultra-high field neuroimaging applications that are vulnerable to off-resonance effects.

Altered temporal, but intact spatial, features of transient network dynamics in psychosis.

Contemporary models of psychosis suggest that a continuum of severity of psychotic symptoms exists, with subthreshold psychotic experiences (PEs) potentially reflecting some genetic and environmental risk factors shared with clinical psychosis. Thus, identifying abnormalities in brain activity that manifest across this continuum can shed new light on the pathophysiology of psychosis. Here, we investigated the moment-to-moment engagement of brain networks ("states") in individuals with schizophrenia (SCZ) and PEs and identified features of these states that are associated with psychosis-spectrum symptoms. Transient brain states were defined by clustering "single snapshots" of blood oxygen level-dependent images, based on spatial similarity of the images. We found that individuals with SCZ (n = 35) demonstrated reduced recruitment of three brain states compared to demographically matched healthy controls (n = 35). Of these three illness-related states, one specific state, involving primarily the visual and salience networks, also occurred at a lower rate in individuals with persistent PEs (n = 22), compared to demographically matched healthy youth (n = 22). Moreover, the occurrence rate of this marker brain state was negatively correlated with the severity of PEs (r = -0.26, p = 0.003, n = 130). In contrast, the spatial map of this state appeared to be unaffected in the SCZ or PE groups. Thus, reduced engagement of a brain state involving the visual and salience networks was demonstrated across the psychosis continuum, suggesting that early disruptions of perceptual and affective function may underlie some of the core symptoms of the illness.

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.

Ultra-high spatial resolution BOLD fMRI in humans using combined segmented-accelerated VFA-FLEET with a recursive RF pulse design.

PURPOSE: To alleviate the spatial encoding limitations of single-shot echo-planar imaging (EPI) by developing multi-shot segmented EPI for ultra-high-resolution functional MRI (fMRI) with reduced ghosting artifacts from subject motion and respiration. THEORY AND METHODS: Segmented EPI can reduce readout duration and reduce acceleration factors, however, the time elapsed between segment acquisitions (on the order of seconds) can result in intermittent ghosting, limiting its use for fMRI. Here, "FLEET" segment ordering, where segments are looped over before slices, was combined with a variable flip angle progression (VFA-FLEET) to improve inter-segment fidelity and maximize signal for fMRI. Scaling a sinc pulse's flip angle for each segment (VFA-FLEET-Sinc) produced inconsistent slice profiles and ghosting, therefore, a recursive Shinnar-Le Roux (SLR) radiofrequency (RF) pulse design was developed (VFA-FLEET-SLR) to generate unique pulses for every segment that together produce consistent slice profiles and signals. RESULTS: The temporal stability of VFA-FLEET-SLR was compared against conventional-segmented EPI and VFA-FLEET-Sinc at 3T and 7T. VFA-FLEET-SLR showed reductions in both intermittent and stable ghosting compared to conventional-segmented and VFA-FLEET-Sinc, resulting in improved image quality with a minor trade-off in temporal SNR. Combining VFA-FLEET-SLR with acceleration, we achieved a 0.6-mm isotropic acquisition at 7T, without zoomed imaging or partial Fourier, demonstrating reliable detection of blood oxygenation level-dependent (BOLD) responses to a visual stimulus. To counteract the increased repetition time from segmentation, simultaneous multi-slice VFA-FLEET-SLR was demonstrated using RF-encoded controlled aliasing. CONCLUSIONS: VFA-FLEET with a recursive RF pulse design supports acquisitions with low levels of artifact and spatial blur, enabling fMRI at previously inaccessible spatial resolutions with a "full-brain" field of view.

In vivo functional localization of the temporal monocular crescent representation in human primary visual cortex.

The temporal monocular crescent (TMC) is the most peripheral portion of the visual field whose perception relies solely on input from the ipsilateral eye. According to a handful of post-mortem histological studies in humans and non-human primates, the TMC is represented visuotopically within the most anterior portion of the primary visual cortical area (V1). However, functional evidence of the TMC visuotopic representation in human visual cortex is rare, mostly due to the small size of the TMC representation (~6% of V1) and due to the technical challenges of stimulating the most peripheral portion of the visual field inside the MRI scanner. In this study, by taking advantage of custom-built MRI-compatible visual stimulation goggles with curved displays, we successfully stimulated the TMC region of the visual field in eight human subjects, half of them right-eye dominant, inside a 3 â€‹T MRI scanner. This enabled us to localize the representation of TMC, along with the blind spot representation (another visuotopic landmark in V1), in all volunteers, which match the expected spatial pattern based on prior anatomical studies. In all hemispheres, the TMC visuotopic representation was localized along the peripheral border of V1, within the most anterior portion of the calcarine sulcus, without any apparent extension into the second visual area (V2). We further demonstrate the reliability of this localization within/across experimental sessions, and consistency in the spatial location of TMC across individuals after accounting for inter-subject structural differences.

The relationship of perceptual discrimination to neural mechanisms of fear generalization.

The generalization of conditioned fear responses has been shown to decrease as a function of perceptual similarity. However, generalization may also extend beyond the perceptual discrimination threshold, ostensibly due to contributions from processes other than perception. Currently the neural mechanisms that mediate perceptual and non-perceptual aspects of fear generalization are unclear. To investigate this question, we conducted a Pavlovian fear conditioning and generalization experiment, collecting functional magnetic resonance imaging (fMRI), skin conductance and explicit shock likelihood ratings, in 37 healthy subjects. Face stimuli were initially paired (CS+) or not paired (CS) with an electrical shock. During the generalization phase, responses were measured to the CS+, CS and a range of CS + -toCS morphs (generalization stimuli), selected for each participant based on that participant's discrimination ability. Across multiple measurements, we found that fear generalization responses were limited to stimuli that could not be distinguished from the CS + stimulus, thus following a gradient closely linked to perceptual discriminability. These measurements, which were correlated with one another, included skin conductance responses, behavioral ratings, and fMRI responses of anterior insula and superior frontal gyrus. In contrast, responses in areas of the default network, including the posterior cingulate gyrus, angular gyrus and hippocampus, showed a negative generalization function extending to stimuli that were more likely to be distinguished from the CS+. In addition, the generalization gradients of the anterior insula and the behavioral ratings showed some evidence for extension beyond perceptual limits. Taken together, these results suggest that distinct brain areas are involved in perceptual and non-perceptual components of fear generalization.

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.

Impact of Visual Corticostriatal Loop Disruption on Neural Processing within the Parahippocampal Place Area.

The caudate nucleus is a part of the visual corticostriatal loop (VCSL), receiving input from different visual areas and projecting back to the same cortical areas via globus pallidus, substantia nigra, and thalamus. Despite perceptual and navigation impairments in patients with VCSL disruption due to caudate atrophy (e.g., Huntington's disease, HD), the relevance of the caudate nucleus and VCSL on cortical visual processing is not fully understood. In a series of fMRI experiments, we found that the caudate showed a stronger functional connection to parahippocampal place area (PPA) compared with adjacent regions (e.g., fusiform face area, FFA) within the temporal visual cortex. Consistent with this functional link, the caudate showed a higher response to scenes compared with faces, similar to the PPA. Testing the impact of VCSL disruption on neural processes within PPA, HD patients showed reduced scene-selective activity within PPA compared with healthy matched controls. In contrast, the level of selective activity in adjacent cortical and subcortical face-selective areas (i.e., FFA and amygdala) remained intact. These results show some of the first evidence for the direct impact and potential clinical significance of VCSL on the generation of "selective" activity within PPA. SIGNIFICANCE STATEMENT: Visual perception is often considered the product of a multistage feedforward neural processing between visual cortical areas, ignoring the likely impact of corticosubcortical loops on this process. Here, we provide evidence for the contribution of visual corticostriatal loop and the caudate nucleus on generating selective response within parahippocampal place area (PPA). Our results show that disruption of this loop in Huntington's disease patients reduces the level of selective activity within PPA, which may lead to related perceptual impairments in these patients.

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.

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.

A problem of overlap.

Here we propose that earlier-demonstrated details in the primate visual cortical map may account for an otherwise puzzling (and problematic) finding in the current human fMRI literature. Specifically, the well-known regions LO and MT(+) reportedly overlap in the human cortical visual map, when those two regions are localized using standard stimulus comparisons in conventional fMRI experiments. Here we describe evidence supporting the idea that the apparent functional overlap between LO and MT arises from a third area (the MT crescent: "MTc"), which is well known to surround posterior MT based on earlier histological, neuroanatomical, and electrophysiological studies in nonhuman primates. If we assume that MTc also exists in human visual cortex, and that it has a location and functional properties intermediate to those in LO and MT, simplistic modeling confirmed that this arrangement could produce apparent overlap between localizers for LO and MT in conventional fMRI maps in human visual cortex.

Selective Functional Connectivity between Ocular Dominance Columns in the Primary Visual Cortex.

The primary visual cortex (V1) in humans and many animals is comprised of fine-scale neuronal ensembles that respond preferentially to the stimulation of one eye over the other, also known as the ocular dominance columns (ODCs). Despite its importance in shaping our perception, to date, the nature of the functional interactions between ODCs has remained poorly understood. In this work, we aimed to improve our understanding of the interaction mechanisms between fine-scale neuronal structures distributed within V1. To that end, we applied high-resolution functional MRI to study mechanisms of functional connectivity between ODCs. Using this technique, we quantified the level of functional connectivity between ODCs as a function of the ocular preference of ODCs, showing that alike ODCs are functionally more connected compared to unalike ones. Through these experiments, we aspired to contribute to filling the gap in our knowledge of the functional connectivity of ODCs in humans as compared to animals.