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Selectivity for direction of motion is a key feature of primary visual cortical neurons. Visual experience is required for direction selectivity in carnivore and primate visual cortex, but the circuit mechanisms of its formation remain incompletely understood. Here, we examined how developing lateral geniculate nucleus (LGN) neurons may contribute to cortical direction selectivity. Using in vivo electrophysiology techniques, we examined LGN receptive field properties of visually naive female ferrets before and after exposure to 6 h of motion stimuli to assess the effect of acute visual experience on LGN cell development. We found that acute experience with motion stimuli did not significantly affect the weak orientation or direction selectivity of LGN neurons. In addition, we found that neither latency nor sustainedness or transience of LGN neurons significantly changed with acute experience. These results suggest that the direction selectivity that emerges in cortex after acute experience is computed in cortex and cannot be explained by changes in LGN cells. [ABSTRACT FROM AUTHOR]

How do cortical responses to local image elements combine to form a spatial pattern of population activity in primate VI? Here, we used voltage-sensitive dye imaging, which measures summed membrane potential activity, to examine the rules that govern lateral interactions between the representations of two small local-oriented elements in macaque (Macaca mulatta) VI. We find strong subadditive and mostly orientation-independent interactions for nearby elements [2- 4 mm interelement cortical distance (IED)] that gradually become linear at larger separations (>6 mm IED). These results are consistent with a population gain control model describing nonlinear VI population responses to single oriented elements. However, because of the membrane potential-to-spiking accelerating nonlinearity, the model predicts supra-additive lateral interactions of spiking responses for intermediate separations at a range of locations between the two elements, consistent with some prior facilitatory effects observed in electrophysiology and psychophysics. Overall, our results suggest that population-level lateral interactions in V1 are primarily explained by a simple orientation-independent contrast gain control mechanism. [ABSTRACT FROM AUTHOR]

Serotonin, an important neuromodulator in the brain, is implicated in affective and cognitive functions. However, its role even for basic cortical processes is controversial. For example, in the mammalian primary visual cortex (VI), heterogenous serotonergic modulation has been observed in anesthetized animals. Here, we combined extracellular single-unit recordings with iontophoresis in awake animals. We examined the role of serotonin on well-defined tuning properties (orientation, spatial frequency, contrast, and size) in V1 of two male macaque monkeys. We find that in the awake macaque the modulatory effect of serotonin is surprisingly uniform: it causes a mainly multiplicative decrease of the visual responses and a slight increase in the stimulus-selective response latency. Moreover, serotonin neither systematically changes the selectivity or variability of the response, nor the interneuronal correlation unexplained by the stimulus ("noise-correlation"). The modulation by serotonin has qualitative similarities with that for a decrease in stimulus contrast, but differs quantitatively from decreasing contrast. It can be captured by a simple additive change to a threshold-linear spiking nonlinearity. Together, our results show that serotonin is well suited to control the response gain of neurons in VI depending on the animal's behavioral or motivational context, complementing other known state-dependent gain-control mechanisms. [ABSTRACT FROM AUTHOR]

Sensory experience is necessary for the development of some receptive field properties of neurons in primary sensory cortical areas. However, it remains unclear whether the parameters of an individual animal's experience play an instructive role and influence the tuning parameters of cortical sensory neurons as selectivity emerges, or rather whether experience merely permits the completion of processes that are fully seeded at the onset of experience. Here we have examined whether the speed of visual stimuli that are presented to visually naive ferrets can influence the parameters of speed tuning and direction selectivity in cortical neurons. Visual experience is necessary for the development of direction selectivity in carnivores. If, during development, cortical neurons had the flexibility to choose from among different inputs with a range of spatial positions and temporal delays, then correlation-based plasticity mechanisms could instruct the precise spatiotemporal selectivity that underlies speed tuning and direction selectivity, and the parameters of an individual animal's experience would influence the tuning that emerges. Alternatively, the tuning parameters of these neurons may already be established at the onset of visual experience, and experience may merely permit the expression of this tuning. We found that providing different groups of animals with either slow (12.5 deg/s) or fast (50 deg/s) visual stimuli resulted in emergence of direction selectivity, but that speed tuning and direction selectivity were similar in the two groups. These results are more consistent with a permissive role for experience in the development of direction selectivity. [ABSTRACT FROM AUTHOR]

Two incongruent images viewed by the two eyes cause binocular rivalry, during which observers perceive continuous alternations between these two visual images. Previous studies in both humans and monkeys have shown that the primary visual cortex (V1) plays a critical role in the rivalry perception. However, it is unclear whether the rivalry activity observed in VI relies on conscious influences. Here, we examine the responses of VI in monkeys under general anesthesia. With intrinsic signal optical imaging and single-trial analysis, alternating activation of ocular dominance columns in V1 was observed during binocularly incongruent stimulation. Left- and right-eye columns exhibited counterphase activation, which were modulated by stimulus features in ways similar to those found in conscious human observers. These observations indicated that binocular rivalry occurs in V1 without consciousness, suggesting that the low-level automatic mechanisms play a more important role than previously believed in handling visual ambiguities [ABSTRACT FROM AUTHOR]

Many studies have reported correlations between the activity of sensory neurons and animals' judgments in discrimination tasks. Here, we suggest that such neuron-behavior correlations may require a cortical map for the task relevant features. This would explain why studies using discrimination tasks based on disparity in area V1 have not found these correlations: V1 contains no map for disparity. This scheme predicts that activity of V1 neurons correlates with decisions in an orientation-discrimination task. To test this prediction, we trained two macaque monkeys in a coarse orientation discrimination task using band-pass-filtered dynamic noise. The two orientations were always 90° apart and task difficulty was controlled by varying the orientation bandwidth of the filter. While the trained animals performed this task, we recorded from orientation-selective V1 neurons (n=82, n=31 for Monkey 1,n=51 for Monkey2). For both monkeys, we observed significant correlation (quantified as "choice probabilities") of the V1 activity with the monkeys' perceptual judgments (mean choice probability 0.54, p= 10–5). In one of these animals, we had previously measured choice probabilities in a disparity discrimination task in V1, which had been at chance (0.49, not significantly different from 0.5). The choice probabilities in this monkey for the orientation discrimination task were significantly larger than those for the disparity discrimination task (p=0.032). These results are predicted by our suggestion that choice probabilities are only observed for cortical sensory neurons that are organized in maps for the task-relevant feature. [ABSTRACT FROM AUTHOR]

Stereo vision relies on cortical signals that encode binocular disparity. In V1, the disparity energy model explains many features of binocular interaction, but it overestimates the responses to anticorrelated images. Combining the outputs of two, or more, energy model-like subunits [two-subunit (2SU) model] can resolve this discrepancy and provides an alternative explanation for disparity signals previously thought to indicate phase disparity between the receptive fields (RFs) of each eye. The 2SU model naturally explains how "near/far" (odd-symmetric) tuning becomes dominant in extrastriate cortex. To compare the energy and the 2SU models, we used a broadband compound grating and applied a common interocular phase difference to all spatial frequency components (a stimulus phase disparity), combined with a common spatial displacement (a stimulus position disparity). This produces binocular images that never occur in natural viewing, for which the 2SU model and the energy model make distinctively different predictions. Responses of neurons recorded from both V1 and V2 of awake rhesus macaques systematically deviated from the predictions of the energy model, in accordance with the 2SU model. These deviations correlated with the symmetry of the tuning curve, indicating that the 2SU mechanism is exploited to produce odd symmetry. Nonetheless, individual subunits also contain RF phase disparity that contributes to odd symmetry. The results suggest that neurons in V2 probably inherit phase disparity signals from V1 neurons, but systematically combine input from V1 neurons with different position disparities, in a way that elaborates odd-symmetric tuning and extends the range of disparities encoded by single neurons. [ABSTRACT FROM AUTHOR]

We used spike-triggered current source-density analysis to examine axonal and postsynaptic currents generated in the visual cortex of awake rabbits by spontaneous spikes of individual sustained and transient dorsal lateral geniculate nucleus (LGNd) neurons. Using these data, we asked whether sustained/transient sensory responses are related to short-term synaptic dynamics at the thalamocortical synapse. Most sustained (34 of 40) and transient (24 of 25) neurons generated axonal and monosynaptic responses in layer 4 and/or 6 of the aligned cortical domain, with input from transient neurons arriving ~0.3 ms earlier and 100 -200 µm deeper. Postsynaptic cortical responses generated by both thalamic cell classes were reduced in amplitude after a preceding impulse and slowly recovered over a period of >750 ms. We interpret this to reflect interval-dependent recovery from chronic depression at the thalamocortical synapse, caused by significant spontaneous firing of LGNd cells (~8 Hz). Surprisingly, postsynaptic cortical responses generated by spontaneous spikes of sustained thalamic neurons were more depressed than those of transient neurons. This difference was seen both in layers 4 and 6. The depression saturated rapidly with multiple preceding impulses, and postsynaptic responses generated by sustained neurons during maintained visual stimulation remained sufficiently robust to allow a sustained flow of information to the cortex. Our results indicate a relationship between the sensory response properties of thalamic neurons and the short-term dynamics of their synapses, and suggest that cortical recipients of sustained and transient thalamic inputs will differ considerably in their response modulation by prior impulse activity. [ABSTRACT FROM AUTHOR]

Prolonged viewing of a chromatically modulated stimulus usually leads to changes in its appearance, and that of similar stimuli. These aftereffects of habituation have been thought to reflect the activity of two populations of neurons in visual cortex that have particular importance in color vision, one sensitive to red-green modulation, the other to blue-yellow, but they have not been identified. We show here, in recordings from macaque primary visual cortex (V1), that prolonged exposure to chromatic modulation reveals two fundamental mechanisms with distinctive chromatic signatures that match those of the mechanisms identified by perceptual observations. In nearly all neurons, these mechanisms contribute to both excitation and to regulatory gain controls, and as a result their habituation can have paradoxical effects on response. The mechanisms must be located near the input layers of V1, before their distinct chromatic signatures diffuse. Our observations suggest that the fundamental mechanisms do not give rise to two distinct L-M and S chromatic pathways. Rather, the mechanisms are better understood as stages in the elaboration of chromatic tuning, expressed in varying proportions in all cells in V1 (and beyond), and made accessible to physiological and perceptual investigation only through habituation. [ABSTRACT FROM AUTHOR]

Contrary to the conventional assumption that humans perceive shapes of rigid objects as constant despite retinal-image variations caused by changes in orientation and position, we show that the depths of three-dimensional (3-D) textured shapes appear to vary when the image is rotated. In paired comparisons of static stimuli, depth amplitude was perceived to be greater at vertical than at oblique orientations. A similar oblique bias was found for simple two-dimensional (2-D) obtuse angles. Using projective geometry to link angle magnitude to the orientation flows that convey 3-D shape from texture cues, we show quantitatively that the 2-D bias predicts the 3-D bias. Our finding that perception of angular magnitude is dependent on orientation has broad implications for shape constancy because orientation flows have also been implicated in 3-D perception from reflections and shading, and contour curvature is fundamental to uncovering depth and part-structure of shapes. We examined the roles played in the observed biases by anisotropies in numbers and tuning widths of orientation-tuned cells in striate cortex as well as the distribution of oriented energy in natural scenes. An optimal stimulus decoding model for 2-D angles revealed that the narrower tuning of cells for horizontal orientations combined with cross-orientation inhibition explains the orientation-dependent angle distortion and hence the 3-D shape inconstancy. Variations in properties within neural populations can thus have direct effects on visual perceptions and need to be included in neural decoding models. [ABSTRACT FROM AUTHOR]

While studying the visual response dynamics of neurons in the macaque primary visual cortex (V1), we found a nonlinearity of temporal response that influences the visual functions of V1 neurons. Simple cells were recorded in all layers of V1; the nonlinearity was strongest in neurons located in layer 2/3. We recorded the spike responses to optimal sinusoidal gratings that were displayed for 100 ms, a temporal step response. The step responses were measured at many spatial phases of the grating stimulus. To judge whether simple cell behavior was consistent with linear temporal integration, the decay of the 100 ms step response at the preferred spatial phase was used to predict the step response at the opposite spatial phase. Responses in layers 4B and 4C were mostly consistent with a linear-plus-static-nonlinearity cascade model. However, this was not true in layer 2/3 where most cells had little or no step responses at the opposite spatial phase. Many layer 2/3 cells had transient preferred-phase responses but did not respond at the offset of the opposite-phase stimuli, indicating a dynamic nonlinearity. A different stimulus sequence, rapidly presented random sinusoids, also produced the same effect, with layer 2/3 simple cells exhibiting elevated spike rates in response to stimuli at one spatial phase but not 180° away. The presence of a dynamic nonlinearity in the responses of V1 simple cells indicates that first-order analyses often capture only a fraction of neuronal behavior. The visual implication of our results is that simple cells in layer 2/3 are spatial phase-sensitive detectors that respond to contrast boundaries of one sign but not the opposite. [ABSTRACT FROM AUTHOR]

Saccadic eye movements are responsible for bringing relevant parts of the visual field onto the fovea for detailed analysis. Because the retina is physiologically unable to deliver sharp images at very high transsaccadic speeds, the visual system minimizes the repercussion of the blurry images we would otherwise perceive during transsaccadic vision by reducing general visual sensitivity and increasing the detection threshold for visual stimuli. Ruling out a pure retinal origin, the effects of saccadic suppression can be already observed some 75 ms before the onset of a saccadic eye movement and are maximal at the onset of motion. The perception of a briefly presented stimulus immediately before the onset of any retinal motion is thus impaired despite the fact that this stimulus is projected onto the stationary retina and is, therefore, physically identical to that presented when no saccadic programming is in course. In this functional magnetic resonance imaging event-related study, we flashed Gabor patches at different times before the onset of a horizontal saccade and measured blood oxygen level-dependent responses at their encoding regions in primary visual cortex (V1) while subjects judged the relative orientation of the stimuli. Closely matching the significant reduction in behavioral performance, the amplitude of the responses in V1 consistently decreased as the stimuli were presented closer to the saccadic onset. These results demonstrate that the neural processes underlying saccade programming transiently modulate cortical responses to briefly presented visual stimuli in areas as early a V1, providing additional evidence for the existence of an active saccadic suppression mechanism in humans. [ABSTRACT FROM AUTHOR]

The primate visual system is arranged hierarchically, starting from the retina and continuing through a series of extrastriate visual areas. Selectivity for motion is first found in individual neurons in the primate visual cortex (V1), in which many simple cells respond selectively to the direction and speed of moving stimuli. Beyond simple cells, most studies of direction selectivity have focused on either V1 complex cells or neurons in the middle temporal area (MT/V5). To understand how visual information is transferred along this pathway, we have studied all three types of neurons, using a reverse correlation procedure to obtain high spatial and temporal resolution maps of activity for different motion stimuli. Most complex and MT cells showed strong second-order interactions, indicating that they were tuned for particular displacements of an apparent motion stimulus. The spatiotemporal structure of these interactions showed a high degree of similarity between the populations of V1 complex cells and MT cells, in terms of the spatiotemporal limits and preferences for motion and their two-dimensional spatial structure. Much of the structure in the V1 and MT second-order kernels could be accounted for on the basis of the first-order responses of V1 simple cells, under the assumption of a Reichardt or motion-energy type of computation. [ABSTRACT FROM AUTHOR]

The response of a neuron in striate cortex to an optimally configured visual stimulus is generally reduced when the stimulus is enlarged to encroach on a suppressive region that surrounds its classical receptive field (CRF). To characterize the mechanism that gives rise to this suppression, we measured its spatiotemporal tuning, its susceptibility to contrast adaptation, and its capacity for interocular transfer. Responses to an optimally configured grating confined to the CRF were strongly suppressed by annular surrounding gratings drifting at a wide range of temporal and spatial frequencies (including spatially uniform fields) that extended from well below to well above the range that drives most cortical neurons. Suppression from gratings capable of driving cortical CRFs was profoundly reduced by contrast adaptation and showed substantial interocular transfer. Suppression from stimuli that lay outside the spatiotemporal passband of most cortical CRFs was relatively stronger when the stimulus on the CRF was of low contrast, was generally insusceptible to contrast adaptation, and showed little interocular transfer. Our findings point to the existence of two mechanisms of surround suppression: one that is prominent when high-contrast stimuli drive the CRF, is orientation selective, has relatively sharp spatiotemporal tuning, is binocularly driven, and can be substantially desensitized by adaptation; the other is relatively more prominent when low-contrast stimuli drive the CRF, has very broad spatiotemporal tuning, is monocularly driven, and is insusceptible to adaptation. Its character suggests an origin in the input layers of primary visual cortex, or earlier. [ABSTRACT FROM AUTHOR]

The human ability to detect modulation of binocular disparity over time is poor compared with detection of luminance modulation. We examined the physiological origin of this limitation by analyzing neuronal responses to temporal modulation of binocular disparity in striate cortex of awake monkeys. When neurons were presented with random-dot stereograms in which disparity varied sinusoidally over time, their responses modulated at the stimulus temporal frequency, with little change in mean firing rate. We calculated modulation amplitude as a function of temporal frequency and compared this with the psychophysical performance of four human observers. Neuronal and psychophysical functions showed similar peak frequencies (2 Hz) and comparable high-cut frequencies (10 and 5.5 Hz, respectively). Thus, V1 (primary visual cortex) neurons appear to limit psychophysical performance. The temporal resolution of the same neurons for contrast modulation was ∼2.5 times greater, which parallels the superior psychophysical performance for contrast. There is a simple mathematical explanation for this difference: it results from calculating cross-correlation between temporally broadband monocular images that are bandpass filtered before measuring correlation. The limit on temporal resolution is a direct consequence of the binocular energy model that adds to the list of properties of human stereoscopic performance that are explained by this simple model of disparity encoding in V1: the same neurons can account for the performance of psychophysical tasks that result in either high (contrast) or low (disparity) temporal resolution. Because this principle holds whenever a broadband input is bandpass filtered before computing correlation, it may limit the resolution of other neuronal systems. [ABSTRACT FROM AUTHOR]

Models of perceptual decision making often assume that sensory evidence is accumulated over time in favor of the various possible decisions, until the evidence in favor of one of them outweighs the evidence for the others. Saccadic eye movements are among the most frequent perceptual decisions that the human brain performs. We used stochastic visual stimuli to identify the temporal impulse response underlying saccadic eye movement decisions. Observers performed a contrast search task, with temporal variability in the visual signals. In experiment 1, we derived the temporal filter observers used to integrate the visual information. The integration window was restricted to the first ∼100 ms after display onset. In experiment 2, we showed that observers cannot perform the task if there is no useful information to distinguish the target from the distractor within this time epoch. We conclude that (1) observers did not integrate sensory evidence up to a criterion level, (2) observers did not integrate visual information up to the start of the saccadic dead time, and (3) variability in saccade latency does not correspond to variability in the visual integration period. Instead, our results support a temporal filter model of saccadic decision making. The temporal impulse response identified by our methods corresponds well with estimates of integration times of V1 output neurons. [ABSTRACT FROM AUTHOR]

Most people see movement in Figure 1, although the image is static. Motion is seen from black → blue → white → yellow → black. Many hypotheses for the illusory motion have been proposed, although none have been tested physiologically. We found that the illusion works well even if it is achromatic: yellow is replaced with light gray, and blue is replaced with dark gray. We show that the critical feature for inducing illusory motion is the luminance relationship of the static elements. Illusory motion is seen from black → dark gray → white light gray → black. In psychophysical experiments, we found that all four pairs of adjacent elements when presented alone each produced illusory motion consistent with the original illusion, a result not expected from any current models. We also show that direction-selective neurons in macaque visual cortex gave directional responses to the same static element pairs, also in a direction consistent with the illusory motion. This is the first demonstration of directional responses by single neurons to static displays and supports a model in which low-level, first-order motion detectors interpret contrast-dependent differences in response timing as motion. We demonstrate that this illusion is a static version of four-stroke apparent motion. [ABSTRACT FROM AUTHOR]

Although the response of a neuron in the visual cortex generally grows nonlinearly with contrast, the spatial tuning of the cell remains stable. This is thought to reflect the activity of a contrast gain control ("normalization") that has very broad tuning on the relevant stimulus dimension. Contrast invariant tuning on a particular dimension is probably necessary for reliable representation of stimuli on that dimension. In the lateral geniculate nucleus (LGN), V1, and V2 of anesthetized macaque, we measured chromatic tuning of neurons at several contrasts to characterize the gain controls and identify cells that might be important for representing color. We estimated separately the chromatic signature of the linear receptive field and that of the gain control. In the LGN, we found normalization in magnocellular cells and cells receiving excitatory S-cone input but not in parvocellular cells or those receiving inhibitory S-cone input. We found normalization in all types of cortical neurons. Among cells that preferred achromatic modulation, or modulation along intermediate directions in color space (making them responsive to both achromatic and chromatic stimuli), normalization was driven by mechanisms tuned to a restricted range of directions in color space, close to achromatic. As a result, chromatic tuning varied with contrast. Among the relatively few cells that strongly preferred chromatic modulation, normalization was driven by mechanisms sensitive to modulation along all directions in color space, especially isoluminant. As a result, chromatic tuning changed little with contrast. To the extent that contrast invariant tuning is important in representing chromaticity, relatively few cortical neurons are involved. [ABSTRACT FROM AUTHOR]

Cortical maps and feedback connections are ubiquitous features of the visual cerebral cortex. The role of the feedback connections, however, is unclear. This study was aimed at revealing possible organizational relationships between the feedback projections from area V2 and the functional maps of orientation and retinotopy in area V1. Optical imaging of intrinsic signals was combined with cytochrome oxidase histochemistry and connectional anatomy in owl monkeys. Tracer injections were administered at orientation-selective domains in regions of pale and thick cytochrome oxidase stripes adjacent to the border between these stripes. The feedback projections from V2 were found to be more diffuse than the intrinsic horizontal connections within V1, but they nevertheless demonstrated clustering. The clusters of feedback axons projected preferentially to interblob cytochrome oxidase regions. The distribution of preferred orientations of the recipient domains in V1 was broad but appeared biased toward values similar to the preferred orientation of the projecting cells in V2. The global spatial distribution of the feedback projections in V1 was anisotropic. The major axis of anisotropy was systematically parallel to a retinotopic axis in V1 corresponding to the preferred orientation of the cells of origin in V2. We conclude that the feedback connections from V2 to V1 might play a role in enhancing the response in V1 to collinear contour elements. [ABSTRACT FROM AUTHOR]

We investigated how V1 neurons integrate two stimulus features by presenting stimuli from a stimulus set made up of all combinations of eight different directions of motion and nine binocular disparities. We investigated the occurrence and shape of the resulting joint tuning function. Among V1 neurons, ∼ 80% were jointly tuned for disparity with orientation or direction. The joint tuning function of all jointly tuned neurons was separable into distinct tuning for disparity on the one hand, and orientation or direction tuning on the other. The degree of separability and the mutual information between the stimulus and the firing rates were strongly correlated. The mutual information of jointly tuned neurons when both features were decoded together was highly correlated with the mutual information when the two features were decoded separately, and the information was then summed. Jointly tuned neurons were just as good at representing information about single features as neurons tuned for only a single feature. The tuning properties of most jointly tuned neurons did not dynamically evolve over time, nor did jointly tuned neurons respond earlier than neurons tuned for only a single feature. The response selectivity of V1 neurons is low and decreases the information that a neuron represents about a stimulus. Together these results suggest that distinct stimulus features are integrated very early in visual processing. Furthermore, V1 generates a distributed representation through low response selectivity that avoids the curse of dimensionality by using separable joint tuning functions. [ABSTRACT FROM AUTHOR]

Studies of the primary visual cortex (V1) have produced models that account for neuronal responses to synthetic stimuli such as sinusoidal gratings. Little is known about how these models generalize to activity during natural vision. We recorded neural responses in area V1 of awake macaques to a stimulus with natural spatiotemporal statistics and to a dynamic grating sequence stimulus. We fit nonlinear receptive field models using each of these data sets and compared how well they predicted time-varying responses to a novel natural visual stimulus. On average, the model fit using the natural stimulus predicted natural visual responses more than twice as accurately as the model fit to the synthetic stimulus. The natural vision model produced better predictions in > 75% of the neurons studied. This large difference in predictive power suggests that natural spatiotemporal stimulus statistics activate nonlinear response properties in a different manner than the grating stimulus. To characterize this modulation, we compared the temporal and spatial response properties of the model fits. During natural stimulation, temporal responses often showed a stronger late inhibitory component, indicating an effect of nonlinear temporal summation during natural vision. In addition, spatial tuning underwent complex shifts, primarily in the inhibitory, rather than excitatory, elements of the response profile. These differences in late and spatially tuned inhibition accounted fully for the difference in predictive power between the two models. Both the spatial and temporal statistics of the natural stimulus contributed to the modulatory effects. [ABSTRACT FROM AUTHOR]

In this study, we compared the organization of orientation preference in visual areas V1, V2, and V3. Within these visual areas, we also quantified the relationship between orientation preference and cytochrome oxidase (CO) staining patterns. V1 maps of orientation preference contained both pinwheels and linear zones. The location of CO blobs did not relate in a systematic way to maps of orientation; although, as in other primates, there were approximately twice as many pinwheels as CO blobs. V2 contained bands of high and low orientation selectivity. The bands of high orientation selectivity were organized into pinwheels and linear zones, but iso-orientation domains were twice as large as those in V1. Quantitative comparisons between bands containing high or low orientation selectivity and CO dark and light bands suggested that at least four functional compartments exist in V2, CO dense bands with either high or low orientation selectivity, and CO light bands with either high or low selectivity. We also demonstrated that two functional compartments exist in V3, with zones of high orientation selectivity corresponding to CO dense areas and zones of low orientation selectivity corresponding to CO pale areas. Together with previous findings, these results suggest that the modular organization of V1 is similar across primates and indeed across most mammals. V2 organization in owl monkeys also appears similar to that of other simians but different from that of prosimians and other mammals. Finally, V3 of owl monkeys shows a compartmental organization for orientation selectivity that remains to be demonstrated in other primates. [ABSTRACT FROM AUTHOR]

The integration of visual information is a critical task that is performed by neurons in the extrastriate cortex of the primate brain. For motion signals, integration is complicated by the geometry of the visual world, which renders some velocity measurements ambiguous and others incorrect. The ambiguity arises because neurons in the early stages of visual processing have small receptive fields, which can only recover the component of motion perpendicular to the orientation of a contour (the aperture problem). Unambiguous motion signals are located at end points and corners, which are referred to as terminators. However, when an object moves behind an occluding surface, motion measurements made at the terminators formed by the intersection of the object and the occluder are generally not consistent with the direction of object motion. To study how cortical neurons integrate these different motion cues, we used variations on the classic "barber pole" stimulus and measured the responses of neurons in the middle temporal area (MT or V5) of extrastriate cortex of alert macaque monkeys. Our results show that MT neurons are more strongly influenced by the unambiguous motion signals generated by terminators than to the ambiguous signals generated by contours. Furthermore, these neurons respond better to terminators that are intrinsic to a moving object than to those that are accidents of occlusion. V1 neurons show similar response patterns to local cues (contours and terminators), but for large stimuli, they do not reflect the global motion direction computed by MT neurons. These observations are consistent with psychophysical findings that show that our perception of moving objects often depends on the motion of terminators. [ABSTRACT FROM AUTHOR]

Disparity selectivity in the striate cortex has generally been studied with uniform disparity fields covering the receptive field (RF). In four awake behaving monkeys, we quantitatively characterized the spatial three-dimensional structure of 55 V1 RFs using random dot stereograms in which disparity varied as a sinusoidal function of vertical position ("corrugations"). At low spatial frequencies, this produced a modulation in neuronal firing at the temporal frequency of the stimulus. As the spatial frequency increased, the modulation reduced. The mean response rate changed little and was close to that produced by a uniform stimulus at the mean disparity of the corrugation. In 48 of 55 (91%) neurons, the modulation strength was a lowpass function of spatial frequency. These results are compatible with a response determined only by the weighted mean of the disparities of the dots (the weights being set by the RF envelope) and suggest that there is no disparity-based surround inhibition or selectivity for disparity gradients. This simple weighting scheme predicts a relationship between RF size and the high-frequency cutoff. Comparison with independent measurements of RF size was compatible with this. All of this behavior closely matches the binocular energy model. The mean cutoff frequency, 0.5 cycles per degree, is similar to equivalent measures of decline in human psychophysical sensitivity for such depth corrugations as a function of frequency (Tyler, 1974; Prince and Rogers, 1998; Banks et al., 2004). This suggests that human cyclopean acuity for disparity modulations is limited by the selectivity of V1 neurons. This in turn is primarily limited by the RF size, because we find no sensitivity for disparity gradients or other disparity differences within the RFs. [ABSTRACT FROM AUTHOR]

In the primary visual cortex (V1), the single-neuron response to a grating stimulus placed in the classical receptive field (CRF) is suppressed by a similar stimulus presented in the CRF surround. To assess the input mechanism underlying the surround suppression, we tested the effects of iontophoretically administered GABAA-receptor antagonist, bicuculline methiodide (BMI), for the 46 V1 neurons in anesthetized cats. First, the stimulus-size tuning curves were studied, with or without BMI administration, for each neuron by changing the size of the grating patch. During the BMI administration, the shape of the normalized size tuning curve did not change considerably. Second, the dependency of surround suppression on the orientation of the surround grating was examined. In the control, the surround suppression showed the clear orientation tuning that peaked at an orientation the same as the optimal orientation of the CRF response. The BMI administration did not change the orientation dependency of surround suppression. We also estimated the relative contribution of excitation and inhibition to the size and orientation tuning of surround suppression. It was concluded that cortical excitation and inhibition were well balanced, having similar tuning profiles for both stimulus size and orientation of the surround grating. Furthermore, surround stimuli used for V1 neurons suppressed the CRF response of neurons in the lateral geniculate nucleus. These results suggest that surround suppression is not primarily attributable to the intracortical inhibition, but because of a reduction of thalamocortical inputs, which drive the cortical excitation and inhibition, and a subsequent decrease in the cortical excitatory interactions. [ABSTRACT FROM AUTHOR]

Stimulation of the suppressive surround of a cortical neuron affects the responsivity and tuning of the classical receptive field (CRF) on several stimulus dimensions. In V1 and V2 of macaques prepared for acute electrophysiological experiments, we explored the chromatic sensitivity of the surround and its influence on the chromatic tuning of the CRF. We studied receptive fields of single neurons with patches of drifting grating of optimal spatial frequency and orientation and variable size, modulated along achromatic or isoluminant color directions. The responses of most neurons declined as the patch was enlarged beyond the optimal size (surround suppression). In V1 the suppression evoked by isoluminant gratings was less than one-half that evoked by achromatic gratings. Consequently, many cells were most sensitive to achromatic modulation when patches just covered the CRF but were most sensitive to isoluminant modulation when patches were enlarged to cover the suppressive surround. Non-oriented neurons that were strongly chromatically opponent generally lacked suppressive surrounds. In V2 most neurons showed equal surround suppression from isoluminant gratings and achromatic gratings. This makes the relative sensitivity of V2 neurons to achromatic and isoluminant gratings mainly independent of the size of the grating. We also measured the chromatic properties of the CRF in the presence of differently colored surrounds. In neither V1 nor V2 did the surround alter the chromatic tuning of the CRF. Cortical mechanisms sensitive to chromatic contrast seem to provide little input to the suppressive surrounds of V1 neurons but substantial input to those of V2 neurons. [ABSTRACT FROM AUTHOR]

Two incongruent images viewed by the two eyes cause binocular rivalry, during which observers perceive continuous alternations between these two visual images. Previous studies in both humans and monkeys have shown that the primary visual cortex (V1) plays a critical role in the rivalry perception. However, it is unclear whether the rivalry activity observed in V1 relies on conscious influences. Here, we examine the responses of V1 in monkeys under general anesthesia. With intrinsic signal optical imaging and single-trial analysis, alternating activation of ocular dominance columns in V1 was observed during binocularly incongruent stimulation. Left- and right-eye columns exhibited counterphase activation, which were modulated by stimulus features in ways similar to those found in conscious human observers. These observations indicated that binocular rivalry occurs in V1 without consciousness, suggesting that the low-level automatic mechanisms play a more important role than previously believed in handling visual ambiguities., Significance Statement: When visual input is ambiguous, for example, in viewing bistable images, human subjects normally perceive one of the interpretations at a particular moment. Previous studies have shown that both low-level visual processing and high-level attention contribute to the establishment of the final visual perception. However, it is not clear whether attention is indispensable in such a process. Here we show that rivalry-like neural activity persisted in monkey V1 when the monkeys were anesthetized and viewed binocularly incongruent stimuli. Such activity has many key features similar to those observed in conscious human subjects. These findings indicate that low-level visual processes play a critical role in solving visual ambiguity such as binocular rivalry., (Copyright © 2016 the authors 0270-6474/16/363231-12$15.00/0.)