Simultaneous processing of spatial and chromatic components of patterned stimuli by the human visual system

We report measurements on discrimination of orientation and magnification made for elements differentiated in colour and/or luminance from their background. By performing measurements at a series of background luminances and for fixed luminance of the elements, we show that with colour contrast, discrimination for both spatial parameters is unimpaired when the background is at isoluminance with the elements. Under simple luminance contrast, however, these discriminations become poorer when the background luminance is within some +/- 5% of that of the elements, and are completely absent when the two values are the same. A deuteranomalous subject is unable to make the spatial discrimination around the isoluminance point for colour contrasts which are too small for him to distinguish, but for which subjects with normal colour vision maintain spatial discriminations at isoluminance. This observation establishes that the physiological mechanisms of normal colour vision, rather than stimulus artefacts, mediate the observed spatial discriminations. We conclude that the visual processing of colour and spatial parameters such as orientation and magnification are intrinsically related to each other.     

Effects of stimulus complexity on simple spatial discriminations

We have extended previous measurements on the detection of a target, differentiated in orientation or magnification from a number of identical reference elements. In these new experiments, we have used more than one class of reference element from which the target must be discriminated. For example, in the case of orientation discrimination, the reference elements are of two classes, which differ from each other in orientation, but are otherwise identical. Representatives of the two classes are present in equal numbers, and the target to be detected is presented at an orientation angle different from that of either class of reference element. Data for discrimination of target magnification or orientation are given for simple geometric elements, namely lines, squares and triangles. In nearly all cases, the characteristic time required for 50% probability of target detection, T1/2, is greater with two classes of reference elements than with a single class. In nearly all cases, T1/2 values for the mixed reference elements increase with the number of reference elements, N, corresponding to discrimination by serial processing. This remains the case even when detection is parallel (T1/2 independent of N) for either class of reference element used separately. We discuss the properties of the spatial discrimination mechanisms which give rise to these responses.     

Spatial, colour and contrast response characteristics of mechanisms which mediate discrimination of pattern orientation and magnification

We have measured response times for the detection of a single target presented against a set of reference elements which are characterised by combinations of four different stimulus parameters; colour, contrast polarity, magnification and orientation. The aim of the experiments was to determine the response characteristics of visual mechanisms which mediate target detection through the discrimination of orientation and magnification. In the first experiments, we determined sensitivity to differences in colour and contrast polarity, and show that the mechanisms responsible for the discrimination of orientation and of magnification are both selective in their responses to colour and to contrast polarity. There are, nonetheless, residual interactions between patterns of different contrast polarities and between those of different colour, and in the latter case, weak interactions persist under equiluminance conditions. In a second set of experiments, we examined the interactions between orientation and magnification. We conclude that the responses of visual mechanisms which mediate target detection through discrimination of orientation are markedly dependent on stimulus magnification whereas those which mediate detection through discrimination of magnification are, in contrast, relatively insensitive to stimulus orientation.     

Visual discrimination of simple geometrical patterns: II. Atypical responses in a subject suffering difficulties with pattern recognition

We have applied the experimental methods described by Ike et al. (Spatial Vision, 1987, 2, 13-29) in the study of pattern discrimination by subjects who, during childhood, experienced severe difficulties with reading and spelling. Of the six subjects studied, one still suffers from such difficulties and we show that she performs atypically in certain of the discrimination tasks. In particular, her discrimination responses for elements such as delta are markedly different from those of all other subjects. We interpret these abnormal responses on the basis of the discrimination mechanisms for line orientation and rotation of the elements which were discussed by Ike et al. (1987).     

A theory for the use of visual orientation information which exploits the columnar structure of striate cortex

A neural model is constructed based on the structure of a visual orientation hypercolumn in mammalian striate cortex. It is then assumed that the perceived orientation of visual contours is determined by the pattern of neuronal activity across orientation columns. Using statistical estimation theory, limits on the precision of orientation estimation and discrimination are calculated. These limits are functions of single unit response properties such as orientation tuning width, response amplitude and response variability, as well as the degree of organization in the neural network. It is shown that a network of modest size, consisting of broadly orientation selective units, can reliably discriminate orientation with a precision equivalent to human performance. Of the various network parameters, the discrimination threshold depends most critically on the number of cells in the hypercolumn. The form of the dependence on cell number correctly predicts the results of psychophysical studies of orientation discrimination. The model system's performance is also consistent with psychophysical data in two situations in which human performance is not optimal. First, interference with orientation discrimination occurs when multiple stimuli activate cells in the same hypercolumn. Second, systematic errors in the estimation of orientation can occur when a stimulus is composed of intersecting lines. The results demonstrate that it is possible to relate neural activity to visual performance by an examination of the pattern of activity across orientation columns. This provides support for the hypothesis that perceived orientation is determined by the distributed pattern of neural activity. The results also encourage the view of neural activity. The results also are determined by the responses of many neurons rather than the sensitivity of individual cells.     

Do alignment thresholds define a critical boundary in long-range detection facilitation with co-linear lines?

Thresholds for line contrast detection (experiment 2) were measured with a two-alternative temporal forced-choice procedure as a function of the spatial position of a vertical target line with regard to two co-linear context lines. The different spatial positions of the target line corresponded to values near the position discrimination threshold (experiment 1) reflecting the just detectable lateral offset, or non-co-linearity, between the context lines which were vertically separated by about 100 minutes of visual arc. Target and context lines were vertically separated by about 30 minutes of arc, had equal contrast polarity in one case, and opposite contrast polarity in the other. Strong line contrast detection facilitation is found at perceptually co-linear target locations. This facilitation decreases noticeably at a horizontal target offset that corresponds to the alignment threshold measured with the context lines. The effects are independent of the relative contrast polarity of target and context and, as shown in a third experiment, also independent of both the relative length or number of lines, and the magnitude of their absolute co-axial separation. This independence seems to hold, provided individual line length and co-axial distance between lines are larger than what appears to be the lower limit of the long-range spatial domain for orientation or contour integration (i.e. 20 minutes of arc), as determined by previous studies. The findings reported here suggest that alignment thresholds are likely to define a critical lateral boundary in long-range detection facilitation with co-linear lines. They support models of contour integration based on interactions between neural mechanisms that integrate local signals of contrast, orientation, and relative position or end-to-end alignment. Such mechanisms may help to explain the formation of representations of virtual contours and object contours in human perception.     

Motions Add,Orientations Don’t,in the Human Visual System

Humans can distinguish between contours of similar orientation, and between directions of visual motion. There is consensus that both of these capabilities depend on selective activation of tuned neural channels. The bandwidths of these tuned channels are estimated here by modelling previously published empirical data. Human subjects were presented with a rapid stream of randomly oriented gratings, or randomly directed motions, and asked to respond when they saw a target stimulus. For the orientation task, subjects were less likely to respond when two preceding orientations were close to the target orientation but differed from each other, presumably due to a failure of summation. For the motion data, by contrast, subjects were more likely to respond when the vector sum of two previous directions was in the target direction. Fitting a cortical signal-processing model to these data showed that the direction bandwidth of motion sensors is about three times the bandwidth of orientation sensors, and that it is the large bandwidth that allows the summation of motion stimuli. The differing bandwidths of orientation and motion sensors presumably equip them for differing tasks, such as orientation discrimination and estimation of heading, respectively.     

A model for the detection of moving targets in visual clutter inspired by insect physiology

We present a computational model for target discrimination based on intracellular recordings from neurons in the fly visual system. Determining how insects detect and track small moving features, often against cluttered moving backgrounds, is an intriguing challenge, both from a physiological and a computational perspective. Previous research has characterized higher-order neurons within the fly brain, known as 'small target motion detectors' (STMD), that respond robustly to moving features, even when the velocity of the target is matched to the background (i.e. with no relative motion cues). We recorded from intermediate-order neurons in the fly visual system that are well suited as a component along the target detection pathway. This full-wave rectifying, transient cell (RTC) reveals independent adaptation to luminance changes of opposite signs (suggesting separate ON and OFF channels) and fast adaptive temporal mechanisms, similar to other cell types previously described. From this physiological data we have created a numerical model for target discrimination. This model includes nonlinear filtering based on the fly optics, the photoreceptors, the 1(st) order interneurons (Large Monopolar Cells), and the newly derived parameters for the RTC. We show that our RTC-based target detection model is well matched to properties described for the STMDs, such as contrast sensitivity, height tuning and velocity tuning. The model output shows that the spatiotemporal profile of small targets is sufficiently rare within natural scene imagery to allow our highly nonlinear 'matched filter' to successfully detect most targets from the background. Importantly, this model can explain this type of feature discrimination without the need for relative motion cues.     

Visual search for oriented lines: the role of angular relations between targets and distractors.

Subjects can perform parallel visual search for an item of unique orientation when it is presented on a background texture composed of lines of two other orientations. A number of cues can be used to speed this search. It helps if the target has a unique categorical status (e.g., it is the only 'steep' item) (Wolfe et al., J. Exp. Psychol. Hum. Percept. Perf. 18, 34-49 1992). It helps if the two background orientations are symmetrical about a vertical axis (Wolfe and Friedman-Hill, Psychol. Sci. 3, 194-198, 1992). The experiments presented here show that it helps if the angles formed by the target with one of the distractor types are smaller than the angles formed by the two distractor orientations. These results illustrate that stimulus similarity is a complex concept even for a simple feature like orientation.     

Asymmetries in oriented-line detection indicate two orthogonal filters in early vision.

Visual detection of a line target differing in orientation from a background of lines may be achieved speedily and effortlessly. Such performance is assumed to occur early in vision and to involve filter mechanisms acting in parallel over the visual field. This study establishes orientational limits on this performance and analytically derives some generic properties of the underlying filters. It was found that, in brief displays, target orientation detection thresholds increased approximately linearly with background orientation, from minima at 0 degree (vertical) and 90 degrees, whereas background orientation detection thresholds decreased approximately linearly with target orientation, from maxima at 0 degree and 90 degrees. Target and background threshold functions were exactly antisymmetric. These data are shown to indicate a model of early line processing dominated by two classes of orientation-sensitive filter with axes close to the vertical and horizontal and orientation-tuning half-widths each of approximately 30 degrees at half-height.     

Horizontal-vertical filters in early vision predict anomalous line-orientation identification frequencies.

A characteristic of early visual processing is a reduction in the effective number of filter mechanisms acting in parallel over the visual field. In the detection of a line target differing in orientation from a background of lines, performance with brief displays appears to be determined by just two classes of orientation-sensitive filter, with preferred orientations close to the vertical and horizontal. An orientation signal represented as a linear combination of responses from such filters is shown to provide a quantitative prediction of the probability density function for identifying the perceived orientation of a target line. This prediction was confirmed in an orientation-matching experiment, which showed that the precision of orientation estimates was worst near the vertical and horizontal and best at about 30 degrees each side of the vertical, a result that contrasts with the classical oblique effect in vision, when scrutiny of the image is allowed. A comparison of predicted and observed frequency distributions showed that the hypothesized orientation signal was formed as an opponent combination and horizontal and vertical filter responses.     

Spatial heterogeneity in the perception of face and form attributes

The identity of an object is a fixed property, independent of where it appears, and an effective visual system should capture this invariance [1-3]. However, we now report that the perceived gender of a face is strongly biased toward male or female at different locations in the visual field. The spatial pattern of these biases was distinctive and stable for each individual. Identical neutral faces looked different when they were presented simultaneously at locations maximally biased to opposite genders. A similar effect was observed for perceived age of faces. We measured the magnitude of this perceptual heterogeneity for four other visual judgments: perceived aspect ratio, orientation discrimination, spatial-frequency discrimination, and color discrimination. The effect was sizeable for the aspect ratio task but substantially smaller for the other three tasks. We also evaluated perceptual heterogeneity for facial gender and orientation tasks at different spatial scales. Strong heterogeneity was observed even for the orientation task when tested at small scales. We suggest that perceptual heterogeneity is a general property of visual perception and results from undersampling of the visual signal at spatial scales that are small relative to the size of the receptive fields associated with each visual attribute.     

Two-dimensional pattern discrimination by the honeybee

For a reward of sugar, bees will learn to prefer a pattern rather than an alternative similar one. This visual discrimination allows us to measure resolution, and to search for the cues that the bees remember and later use to recognize the rewarded pattern. Two systems in parallel, analogous to low pass and high pass filters, are distinguished. The first system discriminates the location and size of at least one area of contrast on each side of the target, with inputs from blue and green receptors, but the ability to discriminate the location of colour depends upon fixation. The bees remember less than a low resolution copy of the image, even when they fixate on a vertical pattern. The second system amplifies the contrast at edges in the pattern, ignoring the direction of contrast, and controls fixation upon the target. Edges are discriminated according to their orientation and radial or tangential arrangement. An axis of bilateral symmetry is detected. However, the relative locations of cues within the image are lost, apparently because the relevant neurones have very large fields. Only the cues, not the whole patterns, are preserved in memory. This system is colour blind because its input is restricted to the receptors with peak sensitivity in the green. The two systems together discriminate many simple patterns, but not all, because the filters are limited in variety.     

Sensory adaptation for timing perception

Recent sensory experience modifies subjective timing perception. For example, when visual events repeatedly lead auditory events, such as when the sound and video tracks of a movie are out of sync, subsequent vision-leads-audio presentations are reported as more simultaneous. This phenomenon could provide insights into the fundamental problem of how timing is represented in the brain, but the underlying mechanisms are poorly understood. Here, we show that the effect of recent experience on timing perception is not just subjective; recent sensory experience also modifies relative timing discrimination. This result indicates that recent sensory history alters the encoding of relative timing in sensory areas, excluding explanations of the subjective phenomenon based only on decision-level changes. The pattern of changes in timing discrimination suggests the existence of two sensory components, similar to those previously reported for visual spatial attributes: a lateral shift in the nonlinear transducer that maps relative timing into perceptual relative timing and an increase in transducer slope around the exposed timing. The existence of these components would suggest that previous explanations of how recent experience may change the sensory encoding of timing, such as changes in sensory latencies or simple implementations of neural population codes, cannot account for the effect of sensory adaptation on timing perception.     

Modulation of backward pattern masking by focal visual attention

The effect of focal visual attention on backward pattern masking was investigated using an orientation discrimination task. The results show that attention reduces primarily the effect of interruption masking, the later component of pattern masking, which occurs when the delay between the target and mask onset is about 50-150 ms. The strongest spatial cueing effect, i.e. the strongest reduction of the orientation discrimination threshold due to focal attention, was observed at intermediate (approximately 100 ms) target-to-mask stimulus onset asynchrony (SOA). There was a weak effect of cueing at shorter SOAs, and no or a very weak attentional effect was present at longer target-to-mask SOAs, where the pattern masking effect is absent. The dynamics of attentional modulation of backward pattern masking correlates closely with the dynamics of the attentional modulation of neuronal responses in the early visual cortex.     

A spatial perturbation technique for the investigation of discrete internal representations of visual patterns

A technique is proposed for the investigation of discrete internal representations assumed to be formed by the visual system in response to pattern stimuli. The proposed technique involves applying a local 1-parameter group of spatial transformations to a pattern to generate a continuum of patterns. The visual discriminability of pairs of perturbed patterns corresponding to small fixed increments in the transformation parameter is determined at various points in the parameter range. By means of a simple rule, the characteristics of this discrimination performance are then related to the probability density functions assumed to underlie the hypothesized discrete internal representations. Two experimental applications of the proposed technique are described. The first is concerned with a discrete internal representation involving the specification of the collinearity or non-collinearity of the points in a pattern; the second is concerned with a discrete internal representation involving the specification of the acuteness or obtuseness of the angle between two lines in a pattern.     

Visual behaviors of the migratory grasshopper,Melanoplus sanguinipes F. (Orthoptera: Acrididae)

The visual behavior of adult Melanoplus sanguinipesF. (Orthoptera: Acrididae) was investigated by placing individuals in the center of an arena and recording their orientation responses to visual targets at the perimeter of the arena. Targets that reflected more 540- to 570-nm light were approached more frequently; however, when reflectance in the 540 -to 570-nm region was combined with reflectance in the 400- to 520-nm region, orientation responses were reduced significantly. This suggests that spectral discrimination in M. sanguinipesinvolves at least two classes of photoreceptors, which respond to different regions of the wavelength spectrum. In addition, grasshoppers oriented to vertical, but not horizontal, contrasting stripes. However, when vertical stripes were added to targets reflecting 520- to 650-nm light, responses to verticals on these targets were not enhanced relative to verticals presented against a target background of 400- to 520-nm + 520- to 650-nm light. Thus, spectral discrimination and vertical stripe fixation appear to be two distinct visual behaviors, controlled by outputs from two classes of photoreceptors and a single class of photoreceptors, respectively, and may be used in different physiological or ecological contexts.     

A human extrastriate area functionally homologous to macaque V4

Extrastriate area V4 is crucial for intermediate form vision and visual attention in nonhuman primates. Human neuroimaging suggests that an area in the lingual sulcus/fusiform gyrus may correspond to ventral V4 (V4v). We studied a human neurological patient, AR, with a putative V4v lesion. The lesion does not affect early visual processing (luminance, orientation, and motion perception). However, it does impair hue perception, intermediate form vision, and visual attention in the upper contralateral visual field. Form deficits occur during discrimination of illusory borders, Glass patterns, curvature, and non-Cartesian patterns. Attention deficits occur during discrimination of the relative positions of object parts, detection of low-salience targets, and orientation discrimination in the presence of distractors. This pattern of deficits is consistent with the known properties of area V4 in nonhuman primates, indicating that AR's lesion affects a cortical region functionally homologous to macaque V4.     

A Computational Model of Afterimage Rotation in the Peripheral Drift Illusion Based on Retinal ON/OFF Responses

Human observers perceive illusory rotations after the disappearance of circularly repeating patches containing dark-to-light luminance. This afterimage rotation is a very powerful phenomenon, but little is known about the mechanisms underlying it. Here, we use a computational model to show that the afterimage rotation can be explained by a combination of fast light adaptation and the physiological architecture of the early visual system, consisting of ON- and OFF-type visual pathways. In this retinal ON/OFF model, the afterimage rotation appeared as a rotation of focus lines of retinal ON/OFF responses. Focus lines rotated clockwise on a light background, but counterclockwise on a dark background. These findings were consistent with the results of psychophysical experiments, which were also performed by us. Additionally, the velocity of the afterimage rotation was comparable with that observed in our psychophysical experiments. These results suggest that the early visual system (including the retina) is responsible for the generation of the afterimage rotation, and that this illusory rotation may be systematically misinterpreted by our high-level visual system.