Mark W. Pettet

Collinear stimuli regulate visual responses depending on cell's contrast threshold

Neurons in the primary visual cortex are selective for the size, orientation and direction of motion of patterns falling within a restricted region of visual space known as the receptive field. The response to stimuli presented within the receptive field can be facilitated or suppressed by other stimuli falling outside the receptive field which, when presented in isolation, fail to activate the cell. Whether this interaction is facilitative,,, or suppressive,,,, depends on the relative orientation of pattern elements inside and outside the receptive field. Here we show that neuronal facilitation preferentially occurs when a near-threshold stimulus inside the receptive field is flanked by higher-contrast, collinear elements located in surrounding regions of visual space. Collinear flanks and orthogonally oriented flanks, however, both act to reduce the response to high-contrast stimuli presented within the receptive field. The observed pattern of facilitation and suppression may be the cellular basis for the observation in humans that the detectability of an oriented pattern is enhanced by collinear flanking elements. Modulation of neuronal responses by stimuli falling outside their receptive fields may thus represent an early neural mechanism for encoding objects and enhancing their perceptual saliency.

Constraints on long range interactions mediating contour detection

Contour detection may be mediated by lateral interactions between neighboring cortical neurons whose receptive fields have collinear axes of preferred orientation. This hypothesis was tested in psychophysical experiments and computer simulations using a contour detection task in which observers searched for groups of Gabor patches that followed spatially extended contour paths embedded in noise consisting of several hundred Gabor patches with random positions and orientations. The orientation-selective units in the simulated neural network were linked by facilitatory interconnections whose strength depended on the geometry (distance, curvature, change in curvature) of smooth curves connecting the orientation axes of units in a pairwise fashion. Psychophysical detection performance was much higher for contour signal groups that followed closed rather than open-ended paths. However, just two sudden changes in orientation of neighboring Gabor patch elements in closed-path contours reduced detection performance to the same levels obtained with open-ended contours. These psychophysical data agreed with the results of the neural network simulations. Furthermore, the simulations also accounted for previous findings that removal of a single Gabor patch element from a closed-path contour group significantly degraded detection performance. We conclude that closure alone is not sufficient to enhance the visibility of a contour. However, if a closed contour meets certain geometric constraints, then lateral interactions based on these constraints can generate facilitation that reverberates around the closed path, thereby enhancing the contours visibility.

Cue-invariant networks for figure and background processing in human visual cortex

Lateral occipital cortical areas are involved in the perception of objects, but it is not clear how these areas interact with first tier visual areas. Using synthetic images portraying a simple texture-defined figure and an electrophysiological paradigm that allows us to monitor cortical responses to figure and background regions separately, we found distinct neuronal networks responsible for the processing of each region. The figure region of our displays was tagged with one temporal frequency (3.0 Hz) and the background region with another (3.6 Hz). Spectral analysis was used to separate the responses to the two regions during their simultaneous presentation. Distributed source reconstructions were made by using the minimum norm method, and cortical current density was measured in a set of visual areas defined on retinotopic and functional criteria with the use of functional magnetic resonance imaging. The results of the main experiments, combined with a set of control experiments, indicate that the figure region, but not the background, was routed preferentially to lateral cortex. A separate network extending from first tier through more dorsal areas responded preferentially to the background region. The figure-related responses were mostly invariant with respect to the texture types used to define the figure, did not depend on its spatial location or size, and mostly were unaffected by attentional instructions. Because of the emergent nature of a segmented figure in our displays, feedback from higher cortical areas is a likely candidate for the selection mechanism by which the figure region is routed to lateral occipital cortex.

Temporal dynamics of the human response to symmetry

Symmetry is a highly salient feature of animals, plants, and the constructed environment. Although the perceptual phenomenology of symmetry processing is well understood, little is known about the underlying neural mechanisms. Here we use visual evoked potentials to measure the time course of neural events associated with the extraction of symmetry in random dot fields. We presented sparse random dot patterns that were symmetric about both the vertical and horizontal axes. Symmetric patterns were alternated with random patterns of the same density every 500 msec, using new exemplars of symmetric and random patterns on each image update. Random/random exchanges were used as a control. The response to updates of random patterns was multiphasic, consisting of P65, N90, P110, N140 and P220 peaks. The response to symmetric/random sequences was indistinguishable from that for random/random sequences up to about 220 msec, after which the response to symmetric patterns became relatively more negative. Symmetry in random dot patterns thus appears to be extracted after an initial response phase that is indifferent to configuration. These results are consistent with the hypothesis (Lee, Mumford, Romero, & Lamme, 1998; Tyler & Baseler, 1998) that the symmetry property is extracted by processing in extrastriate cortex.

Shape and contour detection

Detectability of contours may be affected by long-range interactions between neurons in early stages of visual cortex. Specifically, neurons with receptive fields arrayed along the length of a contour may facilitate each other in a position- and orientation-dependent manner. Accordingly, the overall geometry of a contour should significantly influence both the strength of these long-range interactions and the contours detectability. Psychophysical experiments measuring the detectability of sampled, curvilinear contours hidden by randomly-oriented and -positioned noise elements revealed two main findings. First, changes in direction of curvature degraded contour detectability. Second, the effect of changes in magnitude of curvature were predicted by the average of local curvature along the length of the contour. While the first result emphasizes the importance of uniform direction of curvature, the second result rules out penalties for deviation from circularity.

Colinear facilitation promotes reliability of single-cell responses in cat striate cortex

Abstract. Behavior is controlled by neural activity in the brain. The final outcome of this neural control may critically depend on the firing reliability of individual neurons. A nearly constant, proportional relationship is usually found between the response mean and response variance. Here we asked whether lateral interactions within striate cortex that modulate response magnitude also proportionately modify the response variance of cortical neurons. In many cases, response variability depended on stimulus organization: discrete flankers colinearly placed well outside the neurons receptive field increased response magnitude without a proportional increase in variance, thus improving the neurons response reliability. Since colinear flanker facilitation is often seen near the neurons firing threshold, increased response reliability for weak stimuli may contribute to enhancing perceptual saliency.

Figure-ground interaction in the human visual cortex

Discontinuities in feature maps serve as important cues for the location of object boundaries. Here we used multi-input nonlinear analysis methods and EEG source imaging to assess the role of several different boundary cues in visual scene segmentation. Synthetic figure/ground displays portraying a circular figure region were defined solely by differences in the temporal frequency of the figure and background regions in the limiting case and by the addition of orientation or relative alignment cues in other cases. The use of distinct temporal frequencies made it possible to separately record responses arising from each region and to characterize the nature of nonlinear interactions between the two regions as measured in a set of retinotopically and functionally defined cortical areas. Figure/background interactions were prominent in retinotopic areas, and in an extra-striate region lying dorsal and anterior to area MT+. Figure/background interaction was greatly diminished by the elimination of orientation cues, the introduction of small gaps between the two regions, or by the presence of a constant second-order border between regions. Nonlinear figure/background interactions therefore carry spatially precise, time-locked information about the continuity/discontinuity of oriented texture fields. This information is widely distributed throughout occipital areas, including areas that do not display strong retinotopy.

Spatio-temporal tuning of coherent motion evoked responses in 4-6 month old infants and adults

Motion cues provide a rich source of information about translations of the observer through the environment as well as the movements of objects and surfaces. While the direction of motion can be extracted locally these local measurements are, in general, insufficient for determining object and surface motions. To study the development of local and global motion processing mechanisms, we recorded Visual Evoked Potentials (VEPs) in response to dynamic random dot displays that alternated between coherent rotational motion and random motion at 0.8 Hz. We compared the spatio-temporal tuning of the evoked response in 4-6 months old infants to that of adults by recording over a range of dot displacements and temporal update rates. Responses recorded at the frequency of the coherent motion modulation were tuned for displacement at the occipital midline in both adults in infants. Responses at lateral electrodes were tuned for speed in adults, but not in infants. Infant responses were maximal at a larger range of spatial displacement than that of adults. In contrast, responses recorded at the dot-update rate showed a more similar parametric displacement tuning and scalp topography in infants and adults. Taken together, our results suggest that while local motion processing is relatively mature at 4-6 months, global integration mechanisms exhibit significant immaturities at this age.

Development of Sensitivity to Texture and Contour Information in the Human Infant

Texture discrimination and bounding contour extraction are essential parts of the object segmentation and shape discrimination process. As such, successful texture and contour processing are key components underlying the development of the perception of both objects and surfaces. By recording visual-evoked potentials, we investigate whether young infants can detect orientation-defined textures and contours. We measured responses to an organized texture comprised of many Gabor patches of the same orientation, alternated with images containing the same number of patches, but all of random orientation. These responses were compared with a control condition consisting of the alternation between two independently random configurations. Significant difference potentials were found as early as 25 months, as were significant odd harmonics in the test conditions. Responses were also measured to Gabor patches organized either as circles (all patches tangent to an imaginary circular path) alternated with pinwheels (all patches having a fixed orientation offset from the path). Infants between 6 and 13 months also showed sensitivity to the global organization of the elements along contours. Differential responses to our texture and contour stimuli and their controls could only have been generated by mechanisms that are capable of comparing the relative orientation of 2 or more patches, as no local information at a single patch distinguished the random and organized textures or the circle and pinwheel configurations.

Event-related potentials show configural specificity of global form processing

Glass patterns are a type of moiré created when a random-dot field is overlaid with a rotated, translated or dilated copy. The overall form of the moiré cannot be detected using local processing mechanisms, and because of this, Glass patterns are useful probes of global form processing. Here, we use event-related potentials to show that certain global organizations (concentric structure created by rotation and radial structure produced by dilation) produce much larger brain responses than others (linear structure created by translation). The results are consistent with the existence of specialized form processing mechanisms in the extrastriate cortex.