Spatial stereoresolution for depth corrugations may be set in primary visual cortex.
ABSTRACT: Stereo "3D" depth perception requires the visual system to extract binocular disparities between the two eyes' images. Several current models of this process, based on the known physiology of primary visual cortex (V1), do this by computing a piecewise-frontoparallel local cross-correlation between the left and right eye's images. The size of the "window" within which detectors examine the local cross-correlation corresponds to the receptive field size of V1 neurons. This basic model has successfully captured many aspects of human depth perception. In particular, it accounts for the low human stereoresolution for sinusoidal depth corrugations, suggesting that the limit on stereoresolution may be set in primary visual cortex. An important feature of the model, reflecting a key property of V1 neurons, is that the initial disparity encoding is performed by detectors tuned to locally uniform patches of disparity. Such detectors respond better to square-wave depth corrugations, since these are locally flat, than to sinusoidal corrugations which are slanted almost everywhere. Consequently, for any given window size, current models predict better performance for square-wave disparity corrugations than for sine-wave corrugations at high amplitudes. We have recently shown that this prediction is not borne out: humans perform no better with square-wave than with sine-wave corrugations, even at high amplitudes. The failure of this prediction raised the question of whether stereoresolution may actually be set at later stages of cortical processing, perhaps involving neurons tuned to disparity slant or curvature. Here we extend the local cross-correlation model to include existing physiological and psychophysical evidence indicating that larger disparities are detected by neurons with larger receptive fields (a size/disparity correlation). We show that this simple modification succeeds in reconciling the model with human results, confirming that stereoresolution for disparity gratings may indeed be limited by the size of receptive fields in primary visual cortex.
Project description:For the important task of binocular depth perception from complex natural-image stimuli, the neurophysiological basis for disambiguating multiple matches between the eyes across similar features has remained a long-standing problem. Recurrent interactions among binocular disparity-tuned neurons in the primary visual cortex (V1) could play a role in stereoscopic computations by altering responses to favor the most likely depth interpretation for a given image pair. Psychophysical research has shown that binocular disparity stimuli displayed in 1 region of the visual field can be extrapolated into neighboring regions that contain ambiguous depth information. We tested whether neurons in macaque V1 interact in a similar manner and found that unambiguous binocular disparity stimuli displayed in the surrounding visual fields of disparity-selective V1 neurons indeed modified their responses when either bistable stereoscopic or uniform featureless stimuli were presented within their receptive field centers. The delayed timing of the response behavior compared with the timing of classical surround suppression and multiple control experiments suggests that these modulations are carried out by slower disparity-specific recurrent connections among V1 neurons. These results provide explicit evidence that the spatial interactions that are predicted by cooperative algorithms play an important role in solving the stereo correspondence problem.
Project description:One of the fundamental challenges of binocular vision is that objects project to different positions on the two retinas (binocular disparity). Neurons in visual cortex show two distinct types of tuning to disparity, position and phase disparity, which are the results of differences in receptive field location and profile, respectively. Here, we point out that phase disparity does not occur in natural images. Why, then, should the brain encode it? We propose that phase-disparity detectors help to work out which feature in the left eye corresponds to a given feature in the right. This correspondence problem is plagued by false matches: regions of the image that look similar, but do not correspond to the same object. We show that phase-disparity neurons tend to be more strongly activated by false matches. Thus, they may act as 'lie detectors', enabling the true correspondence to be deduced by a process of elimination.
Project description:The stereo correspondence problem poses a challenge to visual neurons because localized receptive fields potentially cause false responses. Neurons in the primary visual cortex (V1) partially resolve this problem by combining excitatory and suppressive responses to encode binocular disparity. We explored the time course of this combination in awake, monkey V1 neurons using subspace mapping of receptive fields. The stimulus was a binocular noise pattern constructed from discrete spatial frequency components. We forward correlated the firing of the V1 neuron with the occurrence of binocular presentations of each spatial frequency component. The forward correlation yielded a complete set of response time courses to every combination of spatial frequency and interocular phase difference. Some combinations produced suppressive responses. Typically, if an interocular phase difference for a given spatial frequency produced strong excitation, we saw suppression in response to the opposite interocular phase difference at lower spatial frequencies. The suppression was delayed relative to the excitation, with a median difference in latency of 7 ms. We found that the suppressive mechanism explains a well-known mismatch of monocular and binocular signals. The suppressive components increased power at low spatial frequencies in disparity tuning, whereas they reduced the monocular response to low spatial frequencies. This long-recognized mismatch of binocular and monocular signals reflects a suppressive mechanism that helps reduce the response to false matches.
Project description:Decades of experimental studies are available on disparity selective cells in visual cortex of macaque and cat. Recently, local disparity map for iso-orientation sites for near-vertical edge preference is reported in area 18 of cat visual cortex. No experiment is yet reported on complete disparity map in V1. Disparity map for layer IV in V1 can provide insight into how disparity selective complex cell receptive field is organized from simple cell subunits. Though substantial amounts of experimental data on disparity selective cells is available, no model on receptive field development of such cells or disparity map development exists in literature. We model disparity selectivity in layer IV of cat V1 using a reaction-diffusion two-eye paradigm. In this model, the wiring between LGN and cortical layer IV is determined by resource an LGN cell has for supporting connections to cortical cells and competition for target space in layer IV. While competing for target space, the same type of LGN cells, irrespective of whether it belongs to left-eye-specific or right-eye-specific LGN layer, cooperate with each other while trying to push off the other type. Our model captures realistic 2D disparity selective simple cell receptive fields, their response properties and disparity map along with orientation and ocular dominance maps. There is lack of correlation between ocular dominance and disparity selectivity at the cell population level. At the map level, disparity selectivity topography is not random but weakly clustered for similar preferred disparities. This is similar to the experimental result reported for macaque. The details of weakly clustered disparity selectivity map in V1 indicate two types of complex cell receptive field organization.
Project description:Stimuli that project the same retinal visual angle can appear to occupy very different proportions of the visual field if they are perceived to be at different distances [1-8]. Previous research shows that perceived angular size alters the spatial distribution of activity in early retinotopic visual cortex [7, 9-11]. For example, a sphere superimposed on the far end of a corridor scene appears to occupy a larger visual angle and activates a larger region of primary visual cortex (V1) compared with the same sphere superimposed on the near end of the corridor . These previous results, however, were obtained from human subjects using psychophysics and fMRI, a fact that fundamentally limits our understanding of the underlying neuronal mechanisms. Here, we present an animal model that allows for a finer examination of size perception at the level of single neurons. We first show that macaque monkeys perceive a size-distance illusion similarly to humans. Then, using extracellular recordings, we test the specific hypothesis  that neurons in V1 shift the position of their receptive fields (RFs) in response to complex monocular depth cues. Consistent with this hypothesis, we found that when ring-shaped stimuli appeared at the back of the corridor, RFs of V1 neurons shifted toward the center of the rings. When the same stimuli appeared at the front of the corridor, RFs shifted outward. Thus, our results show for the first time that V1 RFs can shift, potentially serving as the neural basis for the perception of angular size.
Project description:Accurate estimation of neuronal receptive fields is essential for understanding sensory processing in the early visual system. Yet a full characterization of receptive fields is still incomplete, especially with regard to natural visual stimuli and in complete populations of cortical neurons. While previous work has incorporated known structural properties of the early visual system, such as lateral connectivity, or imposing simple-cell-like receptive field structure, no study has exploited the fact that nearby V1 neurons share common feed-forward input from thalamus and other upstream cortical neurons. We introduce a new method for estimating receptive fields simultaneously for a population of V1 neurons, using a model-based analysis incorporating knowledge of the feed-forward visual hierarchy. We assume that a population of V1 neurons shares a common pool of thalamic inputs, and consists of two layers of simple and complex-like V1 neurons. When fit to recordings of a local population of mouse layer 2/3 V1 neurons, our model offers an accurate description of their response to natural images and significant improvement of prediction power over the current state-of-the-art methods. We show that the responses of a large local population of V1 neurons with locally diverse receptive fields can be described with surprisingly limited number of thalamic inputs, consistent with recent experimental findings. Our structural model not only offers an improved functional characterization of V1 neurons, but also provides a framework for studying the relationship between connectivity and function in visual cortical areas.
Project description:The dynamic nature of the brain is critical for the success of treatments aimed at restoring vision at the retinal level. The success of these treatments relies highly on the functionality of the surviving neurons along the entire visual pathway. Electrophysiological properties at the retina level have been investigated during the progression of retinal degeneration; however, little is known about the changes in electrophysiological properties that occur in the primary visual cortex (V1) during the course of retinal degeneration. By conducting extracellular recording, we examined the electrophysiological properties of V1 in S334ter-line-3 rats (a transgenic model of retinal degeneration developed to express a rhodopsin mutation similar to that found in human retinitis pigmentosa patients). We measured the orientation tuning, spatial and temporal frequency tunings and the receptive field (RF) size for 127 V1 neurons from 11 S334ter-3 rats and 10 Long-Evans (LE) rats. V1 neurons in the S334ter-3 rats showed weaker orientation selectivity, lower optimal spatial and temporal frequency values and a smaller receptive field size compared to the LE rats. These results suggest that the visual cognitive ability significantly changes during retinal degeneration.
Project description:The primary visual cortex (V1) receives its driving input from the eyes via the lateral geniculate nucleus (LGN) of the thalamus. The lateral pulvinar nucleus of the thalamus also projects to V1, but this input is not well understood. We manipulated lateral pulvinar neural activity in prosimian primates and assessed the effect on supra-granular layers of V1 that project to higher visual cortex. Reversibly inactivating lateral pulvinar prevented supra-granular V1 neurons from responding to visual stimulation. Reversible, focal excitation of lateral pulvinar receptive fields increased the visual responses in coincident V1 receptive fields fourfold and shifted partially overlapping V1 receptive fields toward the center of excitation. V1 responses to regions surrounding the excited lateral pulvinar receptive fields were suppressed. LGN responses were unaffected by these lateral pulvinar manipulations. Excitation of lateral pulvinar after LGN lesion activated supra-granular layer V1 neurons. Thus, lateral pulvinar is able to powerfully control and gate information outflow from V1.
Project description:Inferring depth from binocular disparities is a difficult problem for the visual system because local features in the left- and right-eye images must be matched correctly to solve this "stereo correspondence problem." Cortical architecture and computational studies suggest that lateral interactions among neurons could help resolve local uncertainty about disparity encoded in individual neurons by incorporating contextual constraints. We found that correlated activity among pairs of neurons in primary visual cortex depended both on disparity-tuning relationships and the stimuli displayed within the receptive fields of the neurons. Nearby pairs of neurons with distinct disparity tuning exhibited a decrease in spike correlation at competing disparities soon after response onset. Distant neuronal pairs of similar disparity tuning exhibited an increase in spike correlation at mutually preferred disparities. The observed correlated activity and response dynamics suggests that local competitive and distant cooperative interactions improve disparity tuning of individual neurons over time. Such interactions could represent a neural substrate for the principal constraints underlying cooperative stereo algorithms.
Project description:The mapping of the topographic representation of the visual field onto cortical areas changes throughout the hierarchy of cortical visual areas. The changes are believed to reflect the establishment of modules with different spatial processing emphasis. The receptive fields (RFs) of neurons within these modules, however, may not be governed by the same spatial topographic map parameters. Here it is shown that the RFs of area V4 neurons (centered 1-12 degrees in eccentricity) are based on a circularly symmetric sampling of the primary visual cortical retinotopic map. No eccentricity dependent magnification beyond that observed in V1 is apparent in the V4 neurons. The size and shape of V4 RFs can be explained by a simple, constant sized, two-dimensional Gaussian sample of visual input from the retinotopic map laid out across the surface of V1. Inferences about the spatial scale of interactions within the receptive fields of neurons cannot be based on a visual area's apparent cortical magnification derived from topographic mapping.