Jascha D. Swisher

Visual Topography of Human Intraparietal Sulcus

Human parietal cortex is implicated in a wide variety of sensory and cognitive functions, yet its precise organization remains unclear. Visual field maps provide a potential structural basis for descriptions of functional organization. Here, we detail the topography of a series of five maps of the contralateral visual hemifield within human posterior parietal cortex. These maps are located along the medial bank of the intraparietal sulcus (IPS) and are revealed by direct visual stimulation during functional magnetic resonance imaging, allowing these parietal regions to be routinely and reliably identified simultaneously with occipital visual areas. Two of these maps (IPS3 and IPS4) are novel, whereas two others (IPS1 and IPS2) have previously been revealed only by higher-order cognitive tasks. Area V7, a previously identified visual map, is observed to lie within posterior IPS and to share a foveal representation with IPS1. These parietal maps are reliably observed across scan sessions; however, their precise topography varies between individuals. The multimodal organization of posterior IPS mirrors this variability in visual topography, with complementary tactile activations found immediately adjacent to the visual maps both medially and laterally. These visual maps may provide a practical framework in which to characterize the functional organization of human IPS.

Multiscale Pattern Analysis of Orientation-Selective Activity in the Primary Visual Cortex

Although orientation columns are less than a millimeter in width, recent neuroimaging studies indicate that viewed orientations can be decoded from cortical activity patterns sampled at relatively coarse resolutions of several millimeters. One proposal is that these differential signals arise from random spatial irregularities in the columnar map. However, direct support for this hypothesis has yet to be obtained. Here, we used high-field, high-resolution functional magnetic resonance imaging (fMRI) and multivariate pattern analysis to determine the spatial scales at which orientation-selective information can be found in the primary visual cortex (V1) of cats and humans. We applied a multiscale pattern analysis approach in which fine- and coarse-scale signals were first removed by ideal spatial lowpass and highpass filters, and the residual activity patterns then analyzed by linear classifiers. Cat visual cortex, imaged at 0.3125 mm resolution, showed a strong orientation signal at the scale of individual columns. Nonetheless, reliable orientation bias could still be found at spatial scales of several millimeters. In the human visual cortex, imaged at 1 mm resolution, a majority of orientation information was found on scales of millimeters, with small contributions from global spatial biases exceeding ∼1 cm. Our high-resolution imaging results demonstrate a reliable millimeters-scale orientation signal, likely emerging from irregular spatial arrangements of orientation columns and their supporting vasculature. fMRI pattern analysis methods are thus likely to be sensitive to signals originating from other irregular columnar structures elsewhere in the brain.

Rapid and reversible recruitment of early visual cortex for touch.

Background The loss of vision has been associated with enhanced performance in non-visual tasks such as tactile discrimination and sound localization. Current evidence suggests that these functional gains are linked to the recruitment of the occipital visual cortex for non-visual processing, but the neurophysiological mechanisms underlying these crossmodal changes remain uncertain. One possible explanation is that visual deprivation is associated with an unmasking of non-visual input into visual cortex. Methodology/Principal Findings We investigated the effect of sudden, complete and prolonged visual deprivation (five days) in normally sighted adult individuals while they were immersed in an intensive tactile training program. Following the five-day period, blindfolded subjects performed better on a Braille character discrimination task. In the blindfold group, serial fMRI scans revealed an increase in BOLD signal within the occipital cortex in response to tactile stimulation after five days of complete visual deprivation. This increase in signal was no longer present 24 hours after blindfold removal. Finally, reversible disruption of occipital cortex function on the fifth day (by repetitive transcranial magnetic stimulation; rTMS) impaired Braille character recognition ability in the blindfold group but not in non-blindfolded controls. This disruptive effect was no longer evident once the blindfold had been removed for 24 hours. Conclusions/Significance Overall, our findings suggest that sudden and complete visual deprivation in normally sighted individuals can lead to profound, but rapidly reversible, neuroplastic changes by which the occipital cortex becomes engaged in processing of non-visual information. The speed and dynamic nature of the observed changes suggests that normally inhibited or masked functions in the sighted are revealed by visual loss. The unmasking of pre-existing connections and shifts in connectivity represent rapid, early plastic changes, which presumably can lead, if sustained and reinforced, to slower developing, but more permanent structural changes, such as the establishment of new neural connections in the blind.

Perceptual Learning Selectively Refines Orientation Representations in Early Visual Cortex

Although practice has long been known to improve perceptual performance, the neural basis of this improvement in humans remains unclear. Using fMRI in conjunction with a novel signal detection-based analysis, we show that extensive practice selectively enhances the neural representation of trained orientations in the human visual cortex. Twelve observers practiced discriminating small changes in the orientation of a laterally presented grating over 20 or more daily 1 h training sessions. Training on average led to a twofold improvement in discrimination sensitivity, specific to the trained orientation and the trained location, with minimal improvement found for untrained orthogonal orientations or for orientations presented in the untrained hemifield. We measured the strength of orientation-selective responses in individual voxels in early visual areas (V1–V4) using signal detection measures, both before and after training. Although the overall amplitude of the BOLD response was no greater after training, practice nonetheless specifically enhanced the neural representation of the trained orientation at the trained location. This training-specific enhancement of orientation-selective responses was observed in the primary visual cortex (V1) as well as higher extrastriate visual areas V2–V4, and moreover, reliably predicted individual differences in the behavioral effects of perceptual learning. These results demonstrate that extensive training can lead to targeted functional reorganization of the human visual cortex, refining the cortical representation of behaviorally relevant information.

Working memory for social cues recruits orbitofrontal cortex and amygdala: A functional magnetic resonance imaging study of delayed matching to sample for emotional expressions

During everyday interactions, we continuously monitor and maintain information about different individuals and their changing emotions in memory. Yet to date, working memory (WM) studies have primarily focused on mechanisms for maintaining face identity, but not emotional expression, and studies investigating the neural basis of emotion have focused on transient activity, not delay related activity. The goal of this functional magnetic resonance imaging study was to investigate WM for two critical social cues: identity and emotion. Subjects performed a delayed match-to-sample task that required them to match either the emotional expression or the identity of a face after a 10 s delay. Neuroanatomically, our predictions focused on the orbitofrontal cortex (OFC) and the amygdala, as these regions have previously been implicated in emotional processing and long-term memory, and studies have demonstrated sustained OFC and medial temporal lobe activity during visual WM. Consistent with previous studies, transient activity during the sample period representing emotion and identity was found in the superior temporal sulcus and inferior occipital cortex, respectively. Sustained delay-period activity was evident in OFC, amygdala, and hippocampus, for both emotion and identity trials. These results suggest that, although initial processing of emotion and identity is accomplished in anatomically segregated temporal and occipital regions, sustained delay related memory for these two critical features is held by the OFC, amygdala and hippocampus. These regions share rich connections, and have been shown previously to be necessary for binding features together in long-term memory. Our results suggest a role for these regions in active maintenance as well.

Auditory Spatial Attention Representations in the Human Cerebral Cortex

Auditory spatial attention serves important functions in auditory source separation and selection. Although auditory spatial attention mechanisms have been generally investigated, the neural substrates encoding spatial information acted on by attention have not been identified in the human neocortex. We performed functional magnetic resonance imaging experiments to identify cortical regions that support auditory spatial attention and to test 2 hypotheses regarding the coding of auditory spatial attention: 1) auditory spatial attention might recruit the visuospatial maps of the intraparietal sulcus (IPS) to create multimodal spatial attention maps; 2) auditory spatial information might be encoded without explicit cortical maps. We mapped visuotopic IPS regions in individual subjects and measured auditory spatial attention effects within these regions of interest. Contrary to the multimodal map hypothesis, we observed that auditory spatial attentional modulations spared the visuotopic maps of IPS; the parietal regions activated by auditory attention lacked map structure. However, multivoxel pattern analysis revealed that the superior temporal gyrus and the supramarginal gyrus contained significant information about the direction of spatial attention. These findings support the hypothesis that auditory spatial information is coded without a cortical map representation. Our findings suggest that audiospatial and visuospatial attention utilize distinctly different spatial coding schemes.

Prevalence of Selectivity for Mirror-Symmetric Views of Faces in the Ventral and Dorsal Visual Pathways

Although the ability to recognize faces and objects from a variety of viewpoints is crucial to our everyday behavior, the underlying cortical mechanisms are not well understood. Recently, neurons in a face-selective region of the monkey temporal cortex were reported to be selective for mirror-symmetric viewing angles of faces as they were rotated in depth (Freiwald and Tsao, 2010). This property has been suggested to constitute a key computational step in achieving full view-invariance. Here, we measured functional magnetic resonance imaging activity in nine observers as they viewed upright or inverted faces presented at five different angles (−60, −30, 0, 30, and 60°). Using multivariate pattern analysis, we show that sensitivity to viewpoint mirror symmetry is widespread in the human visual system. The effect was observed in a large band of higher order visual areas, including the occipital face area, fusiform face area, lateral occipital cortex, mid fusiform, parahippocampal place area, and extending superiorly to encompass dorsal regions V3A/B and the posterior intraparietal sulcus. In contrast, early retinotopic regions V1–hV4 failed to exhibit sensitivity to viewpoint symmetry, as their responses could be largely explained by a computational model of low-level visual similarity. Our findings suggest that selectivity for mirror-symmetric viewing angles may constitute an intermediate-level processing step shared across multiple higher order areas of the ventral and dorsal streams, setting the stage for complete viewpoint-invariant representations at subsequent levels of visual processing.

Radial bias is not necessary for orientation decoding.

Multivariate pattern analysis can be used to decode the orientation of a viewed grating from fMRI signals in early visual areas. Although some studies have reported identifying multiple sources of the orientation information that make decoding possible, a recent study argued that orientation decoding is only possible because of a single source: a coarse-scale retinotopically organized preference for radial orientations. Here we aim to resolve these discrepant findings. We show that there were subtle, but critical, experimental design choices that led to the erroneous conclusion that a radial bias is the only source of orientation information in fMRI signals. In particular, we show that the reliance on a fast temporal-encoding paradigm for spatial mapping can be problematic, as effects of space and time become conflated and lead to distorted estimates of a voxels orientation or retinotopic preference. When we implement minor changes to the temporal paradigm or to the visual stimulus itself, by slowing the periodic rotation of the stimulus or by smoothing its contrast-energy profile, we find significant evidence of orientation information that does not originate from radial bias. In an additional block-paradigm experiment where space and time were not conflated, we apply a formal model comparison approach and find that many voxels exhibit more complex tuning properties than predicted by radial bias alone or in combination with other known coarse-scale biases. Our findings support the conclusion that radial bias is not necessary for orientation decoding. In addition, our study highlights potential limitations of using temporal phase-encoded fMRI designs for characterizing voxel tuning properties.

Phase-encoded fmri investigation of retinotopic remapping responses

We adapted the standard polar mapping presentation paradigm to contain back-and-forth saccades every TR (2s). In the mapping condition, wedge stimuli were presented at fixation after each saccade. In the remapping condition, wedge stimuli were presented in the periphere and saccades were made to the stimulus after it was extinguished34 A correspondence in phase between these conditions would indicate that the shape of a saccadic target is remapped across saccades. Phase-encoded fmri investigation of retinotopic remapping responses

Multishot versus single-shot pulse sequences in very high field fMRI: a comparison using retinotopic mapping.

High-resolution functional MRI is a leading application for very high field (7 Tesla) human MR imaging. Though higher field strengths promise improvements in signal-to-noise ratios (SNR) and BOLD contrast relative to fMRI at 3 Tesla, these benefits may be partially offset by accompanying increases in geometric distortion and other off-resonance effects. Such effects may be especially pronounced with the single-shot EPI pulse sequences typically used for fMRI at standard field strengths. As an alternative, one might consider multishot pulse sequences, which may lead to somewhat lower temporal SNR than standard EPI, but which are also often substantially less susceptible to off-resonance effects. Here we consider retinotopic mapping of human visual cortex as a practical test case by which to compare examples of these sequence types for high-resolution fMRI at 7 Tesla. We performed polar angle retinotopic mapping at each of 3 isotropic resolutions (2.0, 1.7, and 1.1 mm) using both accelerated single-shot 2D EPI and accelerated multishot 3D gradient-echo pulse sequences. We found that single-shot EPI indeed led to greater temporal SNR and contrast-to-noise ratios (CNR) than the multishot sequences. However, additional distortion correction in postprocessing was required in order to fully realize these advantages, particularly at higher resolutions. The retinotopic maps produced by both sequence types were qualitatively comparable, and showed equivalent test/retest reliability. Thus, when surface-based analyses are planned, or in other circumstances where geometric distortion is of particular concern, multishot pulse sequences could provide a viable alternative to single-shot EPI.