A J Parker

Binocular depth perception and the cerebral cortex

Our ability to coordinate the use of our left and right eyes and to make use of subtle differences between the images received by each eye allows us to perceive stereoscopic depth, which is important for the visual perception of three-dimensional space. Binocular neurons in the visual cortex combine signals from the left and right eyes. Probing the roles of binocular neurons in different perceptual tasks has advanced our understanding of the stages within the visual cortex that lead to binocular depth perception.

Integration of depth modules: Stereopsis and texture

Global shape judgements were employed to examine the combination of stereopsis and shape-from-texture in the determination of three-dimensional shape. Adding textural variations to stereograms increased perceived depth. Thus, texture was not simply vetoed by the strong stereo cue. In experiments where the depth specified by texture was incongruent with that specified by stereo, the data were well described by a weighted linear combination rule. Although only a small weight was assigned to texture, this weight was somewhat greater at a farther viewing distance. This could be a consequence of the decreased reliability of stereopsis at far viewing distances.

Contrast sensitivity and orientation selectivity in lamina IV of the striate cortex of Old World monkeys.

SummaryContrast sensitivity and orientation selectivity were measured for neurons in lamina IV of macaque striate cortex. Contrast sensitivity was determined for a range of spatial frequencies, using a staircase method. The stimuli were at the optimal orientation, direction and speed of drift for each neuron. The assignment of each recording site to a subdivision of lamina IV was made by histological reconstruction of each electrode penetration from sections reacted for cytochrome oxidase and stained for Nissl substance. Neurons in the magnocellular recipient zone of IVc (IVcα) have high contrast sensitivities, while those in the parvocellular recipient zone (IVcβ) have low contrast sensitivities. Both of the upper subdivisions of lamina IV (IVa and IVb) contain a mixture of neurons with high and low contrast sensitivities. There were orientation selective neurons within all subdivisions of lamina IV, even in IVc, whereas non-oriented neurons were found only in those subdivisions that receive a direct parvocellular geniculate input (IVa and IVcβ).

Local disparity not perceived depth is signaled by binocular neurons in cortical area V1 of the Macaque.

Binocular neurons that are closely related to depth perception should respond selectively for stimuli eliciting an appropriate depth sensation. To separate perceived depth from local disparity within the receptive field, sinusoidal luminance gratings were presented within a circular aperture. The disparity of the aperture was coupled to that of the grating, thereby rendering unambiguous the psychophysical matching between repeating cycles of the grating. In cases in which the stimulus disparity differs by one horizontal period of the grating, the portion of the grating that locally covers a receptive field is binocularly identical, but the depth sensation is very different because of the aperture. For 117 disparity-selective V1 neurons tested in two monkeys, the overwhelming majority responded equally well to configurations that were locally identical but led to different perceptions of depth. Because the psychophysical sensation is not reflected in the firing rate of V1 neurons, the signals that make stereo matches explicit are most likely elaborated in extrastriate cortex.

Binocular mechanisms for detecting motion-in-depth

There are in principle at least two binocular sources of information that could be used to determine the motion of an object towards or away from an observer: such motion produces changes in binocular disparities over time and also generates different image velocities in the two eyes. Existing psychophysical and physiological evidence is reviewed. It is concluded that these data are inconclusive concerning whether one or both of these sources of information are used in primate vision. Thresholds were measured for disparity modulations in dynamic (temporally uncorrelated) random dot stereograms (RDS), and for RDS in which the same random dot pattern was used throughout (temporally correlated). Although the first stimulus contains no consistent inter-ocular velocity differences, thresholds were generally slightly lower for this stimulus than for temporally correlated stimuli. Sensitivity to the temporal derivative of disparity is therefore adequate to account for human stereomotion detection. A stimulus was devised in which monocular motion was clearly visible to each eye (with opposite velocities) but in which all disparity changes were beyond the temporal resolution of stereopsis. This produced no sensation of motion-in-depth. Similarly, stimuli beyond the spatial resolution of stereopsis did not support stereomotion detection. These data strongly suggest that stereomotion is primarily detected by means of temporal changes in binocular disparity. We argue that there is no experimental evidence that supports the existence of a mechanism sensitive to inter-ocular velocity differences.

Capabilities of monkey cortical cells in spatial-resolution tasks

The performance of individual neurons in monkey striate cortex has been examined in three spatial-resolution tasks by making microelectrode recordings from single cells in anaesthetized, paralyzed animals. The statistical reliability of responses from cells was used to estimate threshold levels of performance. For each task (resolution acuity for high-contrast gratings, discrimination of gratings varying in spatial frequency, and localization ability, i.e., discrimination of spatial phase), performance approaching psychophysical thresholds was obtained from single cortical cells. The receptive-field organization underlying localization performance was examined in detail by the use of a linear model that relates localization ability to the sensitivity of the receptive field to luminance contrast. Calculations from this model agree well with direct measurements of localization performance and are comparable with psychophysical measurements of hyperacuity. Though it has been suggested that cells with nonoriented receptive fields in cortical layer IVc beta may be responsible for recovering fine-grain spatial information, our calculations indicate that these cells are poorer at localization than many other cells in the cortex.

Interaction of frontal and perirhinal cortices in visual object recognition memory in monkeys.

Monkeys were trained preoperatively in visual object recognition memory. The task was delayed matching‐to‐sample with lists of trial‐unique randomly generated visual stimuli in an automated apparatus, and the stimuli were 2D visual objects made from randomly generated coloured shapes. We then examined the effect of either: (i) disconnecting the frontal cortex in one hemisphere from the perirhinal cortex in the contralateral hemisphere by crossed unilateral ablations; (ii) disconnecting the magnocellular portion of the mediodorsal (MDmc) thalamic nucleus in one hemisphere from the perirhinal cortex in the contralateral hemisphere; or (iii) bilaterally ablating first the amygdala, then adding fornix transection, then finally perirhinal cortex ablation. We found that both frontal/perirhinal and MDmc/perirhinal disconnection had a large effect on visual object recognition memory, whereas both amygdalectomy and the addition of fornix transection had only a mild effect. We conclude that the frontal lobe needs to interact with the perirhinal cortex within the same hemisphere for visual object recognition memory, but that routes through the amygdala and hippocampus are not of primary importance.

Effects of different texture cues on curved surfaces viewed stereoscopically.

Stereoscopic shape judgements can be modified by the addition of texture cues. This paper examines the properties of texture that are responsible for this effect. When a three-dimensional curved surface is projected onto a two-dimensional image, changes in surface orientation result in gradients of texture element size (or area), shape (compression) and density in the image. Manipulating each of these gradients independently we found that 97% of the variance in the results could be accounted for by the compression gradient. When the texture pattern corresponds to a highly anisotropic texture on the objects surface, shape-from-texture becomes ineffective. These results suggest that human shape-from-texture proceeds under the assumption that textures are statistically isotropic, and not that they are homogeneous.

Independent anatomical and functional measures of the V1/V2 boundary in human visual cortex

The cerebral cortex has both anatomical and functional specialization, but the level of correspondence between the two in the human brain has remained largely elusive. Recent successes in high-resolution magnetic resonance imaging of myeloarchitecture patterns in the cortex suggest that it may now be possible to compare directly human anatomy and function in vivo. We independently investigated the anatomical and functional borders between primary and secondary human visual areas (V1 and V2) in vivo. Functional borders were mapped with functional magnetic resonance imaging (fMRI) using a narrow, vertical black and white contrast-reversing wedge. In three separate scanning sessions, anatomical images were collected at three different slice orientations (300 microm x 300 microm, slice thickness, 1.5 mm). The anatomical signature of V1 was determined by the presence of a hypointense band in the middle of the cortical gray matter. The band was identified in between 81% and 33% (mean 57%) of V1 defined using fMRI, and less than 5% of the identified band was in cortex outside V1. Intensity profiles taken through the gray matter on the V1 and V2 sides of the functional border indicate a measurable difference in the size of the hypointense band for all subjects. This is the first demonstration that the definition of V1 by fMRI closely matches the anatomically defined striate cortex in the human brain. The development of very high-resolution structural MRI may permit the definition of cortical areas based on myeloarchitecture when functional definition is not possible.

A simple model accounts for the response of disparity-tuned V1 neurons to anticorrelated images.

Disparity-tuned cells in primary visual cortex (VI) are thought to play a significant role in the processing of stereoscopic depth. The disparity-specific responses of these neurons have been previously described by an energy model based on local, feedforward interactions. This model fails to predict the response to binocularly anticorrelated stimuli, in which images presented to left and right eyes have opposite contrasts. The original energy model predicts that anticorrelation should invert the disparity tuning curve (phase difference pi), with no change in the amplitude of the response. Experimentally, the amplitude tends to be reduced with anticorrelated stimuli and a spread of phase differences is observed, although phase differences near pi are the most common. These experimental observations could potentially reflect a modulation of the V1 signals by feedback from higher visual areas (because anticorrelated stimuli create a weaker or nonexistent stereoscopic depth sensation). This hypothesis could explain the effects on amplitude, but the spread of phase differences is harder to understand. Here, we demonstrate that changes in both amplitude and phase can be explained by a straightforward modification of the energy model that involves only local processing. Input from each eye is passed through a monocular simple cell, incorporating a threshold, before being combined at a binocular simple cell that feeds into the energy computation. Since this local feedforward model can explain the responses of complex cells to both correlated and anticorrelated stimuli, there is no need to invoke any influence of global stereoscopic matching.