Decoding working memory of stimulus contrast in early visual cortex.

Most studies of the early stages of visual analysis (V1-V3) have focused on the properties of neurons that support processing of elemental features of a visual stimulus or scene, such as local contrast, orientation, or direction of motion. Recent evidence from electrophysiology and neuroimaging studies, however, suggests that early visual cortex may also play a role in retaining stimulus representations in memory for short periods. For example, fMRI responses obtained during the delay period between two presentations of an oriented visual stimulus can be used to decode the remembered stimulus orientation with multivariate pattern analysis. Here, we investigated whether orientation is a special case or if this phenomenon generalizes to working memory traces of other visual features. We found that multivariate classification of fMRI signals from human visual cortex could be used to decode the contrast of a perceived stimulus even when the mean response changes were accounted for, suggesting some consistent spatial signal for contrast in these areas. Strikingly, we found that fMRI responses also supported decoding of contrast when the stimulus had to be remembered. Furthermore, classification generalized from perceived to remembered stimuli and vice versa, implying that the corresponding pattern of responses in early visual cortex were highly consistent. In additional analyses, we show that stimulus decoding here is driven by biases depending on stimulus eccentricity. This places important constraints on the interpretation for decoding stimulus properties for which cortical processing is known to vary with eccentricity, such as contrast, color, spatial frequency, and temporal frequency.

The role of contrast sensitivity in global motion processing deficits in the elderly.

This study compared the effects of age on the perception of translational, radial, and rotational global motion patterns. Motion coherence thresholds were measured for judging the direction of each motion type as a function of contrast (visibility) and temporal sampling rate in young and elderly participants. Coherence thresholds decreased as dot contrast increased asymptoting at high dot contrasts but were higher in elderly compared to young participants. This equated to global motion impairment in the elderly of a factor of around 2, characterized by a shift of the threshold vs. contrast function along the horizontal axes (dot contrast). The effect of contrast interacted with the temporal sampling rate. Old participants were deleteriously affected by reduced temporal sampling particularly at low contrasts. The findings suggest that age-related changes in global motion perception may be driven principally by deficits in contrast encoding, rather than by deficits in motion integration and suggest a role for increased internal noise in the older visual system.

Temporal characteristics of global motion processing revealed by transcranial magnetic stimulation.

The ability to detect the motion of objects is critical to survival, and understanding the cortical mechanisms involved in this process remains a key challenge in sensory neuroscience. A relatively new approach to this problem is to temporarily disrupt processing at specific cortical sites and measure the behavioural consequences. Several previous studies have shown that transcranial magnetic stimulation (TMS) of human visual area V5/MT disrupts global motion perception, but reports vary widely in the timescale of this effect. To resolve this issue we employed psychophysical techniques to investigate how discrimination of translational, rotational and radial global motion is affected by TMS. Prior to applying TMS we established baseline coherence thresholds for global motion perception. Adopting each observer's coherence level at threshold we examined how TMS delivered to V5/MT modulated performance. Importantly, we measured the influence of single-pulse TMS over a broad temporal range to reveal the fine temporal structure of the disruption profile for global motion perception. Results show that the disruption profile consisted of two distinct epochs during which global direction judgments were reliably impaired, separated by an interval in which performance was unaffected. The bimodal nature of the distribution profiles is consistent with feedforward and feedback processing between visual areas mediating global motion processing. We present a novel quantitative model that characterizes the contribution of each process to visual motion perception.

Choice reaction times for identifying the direction of first-order motion and different varieties of second-order motion.

This study sought to quantify the temporal properties of the human visual system by measuring forced-choice reaction times for discriminating the drift direction of first-order motion (luminance-modulated noise) and a variety of second-order motion patterns (modulations of either the contrast, polarity, orientation or spatial length of a noise carrier) over a range of stimulus modulation depths. In general, reaction times for all types of second-order motion were slower than those for first-order motion. Specifically, reaction times were similar for modulations of image contrast, polarity and orientation but were markedly slower for modulations of spatial length. There was also a tendency for reaction times to decrease as stimulus modulation depth increased. The rate of this decrease was shallowest for first-order, luminance-defined patterns. For second-order motion reaction times decreased at a similar rate for contrast, polarity and orientation but this decrease was steepest for spatial length. However, when equated in terms of visibility (multiples of direction-discrimination threshold), the rate at which reaction times decreased as modulation depth increased became comparable for patterns defined by luminance, contrast, polarity and orientation. For patterns defined by spatial length, performance could not be equated in this manner. These findings demonstrate that the time taken to encode the direction of each pattern is not an invariant response metric. The results are consistent with psychophysical and electrophysiological evidence for longer response latencies for second-order motion and may reflect the additional processing stages (e.g. filter-rectify-filter) required for its extraction.

Families of models for gabor paths demonstrate the importance of spatial adjacency.

This paper reports psychophysical and modelling results concerning the contour-detection paradigm of D. J. Field, A. Hayes, and R. F. Hess (1993). We measured psychophysically the maximum tolerable contour curvature (path angle) as a function of contour length. We compared these data to the predictions of an association field (D. J. Field et al., 1993) model based on the relative positions and mutual orientations of nearby elements and to models that explicitly link adjacent elements into chains and characterize each chain by its sequence of contour bends. For every stimulus, a large set of chains is produced and the target identified as the chain with the lowest maximum bend. We tested two different types of linking process: isotropic (linking one element to any other nearby) and anisotropic (linking one element to any others nearby along the orientation of its axis). All of these models can account for our data. Moreover, we show that the pattern of results due to path angle is principally a product of the distribution of spurious contours in the randomly oriented background. Given that some of the models do not embody constraints of orientation relationships between linked elements, this finding shows the importance for early vision in deciding which local elements are to be associated.

Asymmetric spatial frequency tuning of motion mechanisms in human vision revealed by masking.

PURPOSE: To investigate the spatial frequency selectivity of the human motion system by using the technique of visual masking. METHODS: Modulation-depth thresholds for identifying the direction of a sinusoidal test pattern were measured over a range of spatial frequencies (0.25-4 cyc/deg) in the absence and presence of a temporally jittering mask. RESULTS: At the lowest test frequency (0.25 cyc/deg), maximum masking occurred when the test and mask shared the same spatial frequency, decreasing as the difference in spatial frequency between the test and mask increased. However, as test spatial frequency increased, maximum masking began to shift to when the mask was presented at approximately 1 octave below the test spatial frequency. Control experiments demonstrated that the asymmetric masking functions at higher test spatial frequencies was not affected by mask amplitude nor was it an effect of speed. The results confirmed that the peak at 1 octave from the test still occurred when the potential for off-frequency looking was minimized by presenting two masks positioned equidistant in frequency from the test grating. Control experiments revealed, however, that the peak at 1 octave below the test was mediated by image size and/or the number of cycles presented on screen. CONCLUSIONS: These findings provide support for the notion that motion perception is mediated by band-pass, spatial-frequency-selective mechanisms. Moreover, asymmetric tuning of the masking functions may reflect asymmetric spatial frequency selectivity of the mechanisms in the human visual system that encode motion or inhibition between mechanisms tuned to different spatial frequencies.

Second-order optic flow processing.

Optic flow-large-field rotational and radial motion-is processed as efficiently as translational motion for first-order (luminance-defined) stimuli. However, it has been suggested recently that the same pattern does not hold for second-order (e.g. contrast-defined) stimuli. We used random dot kinematogram (RDK) stimuli to determine whether global processing of optic flow is as efficient as processing of global translational motion for both first- and second-order stimuli. For first-order stimuli, we found that coherence thresholds for radial and rotational motion were equivalent to thresholds for translational motion, supporting previous findings. For second-order stimuli we found, firstly, that given sufficient contrast, second-order optic flow can be processed as efficiently as first-order optic flow and, secondly, that rotational and translational second-order motion are processed with equal efficiency. This contradicts the suggestion that there is a loss of efficiency between integration of second-order global motion and second-order optic flow. The third interesting finding was that the processing of radial second-order motion appears to suffer from a deficit that is dependent upon both the contrast and spatial extent of the stimulus. Further experiments discounted the possibility that the observed deficit is caused by a centrifugal or centripetal bias, but demonstrated that a longer temporal integration period for radial second-order motion is responsible for the observed difference. For durations of approximately 850ms, all three types of motion are processed with equal efficiency.

The influences of visibility and anomalous integration processes on the perception of global spatial form versus motion in human amblyopia.

Do amblyopes demonstrate general irregularities in processes of global image integration? Or are these anomalies stimulus specific? To address these questions we employed directly analogous global-orientation and global-motion stimuli using a method that allows us to factor out any influence of the low-level visibility loss [Simmers, A. J., Ledgeway, T., Hess, R. F., & McGraw, P. V. (2003). Deficits to global motion processing in human amblyopia. Vision Research 43, pp. 729-738]. The combination of orientation and motion coherence thresholds reported here provides comparable psychophysical measures of global processing by spatial-sensitive and motion-sensitive mechanisms in the amblyopic visual system. The results show deficits in both global-orientation and global-motion processing in amblyopia, which appear independent of any low-level visibility loss, but with the most severe deficit affecting the extraction of global motion. This provides evidence for the existence of a dominant temporal processing deficit in amblyopia.

Attentional modulation of threshold sensitivity to first-order motion and second-order motion patterns.

Previous studies [e.g. Vision Research 40 (2000) 173] have shown that when observers are required to selectively attend to one of two, spatially-adjacent patches containing either first-order (luminance-defined) or second-order (contrast-defined) motion, threshold sensitivity for identifying the direction of second-order motion, but not first-order motion, is enhanced for the attended stimuli. The processing of second-order motion, unlike first-order motion, may, therefore, require attention. However, other studies have found little evidence for differential effects of attention on the processing of first-order and second-order motion [Investigative Ophthalmology and Visual Science 42(4) (2001) 5061]. We investigated the effects of attention instructions on the ability of observers to identify the directions and spatial orientations of luminance-defined and contrast-defined motion stimuli. Pairs of motion stimuli were presented simultaneously and threshold performance was measured over a wide range of drift temporal frequencies and stimulus durations. We found: (1) direction discrimination thresholds for attended motion stimuli were lower than those for unattended stimuli for both types of motion. The magnitude of this effect was reduced when the observers were not given prior knowledge of which patch of motion (attended or unattended) they had to judge first. (2) Direction discrimination for first-order motion was similarly affected at all temporal frequencies and durations examined, but for second-order motion the effects of attention depended critically on the drift temporal frequency and stimulus duration used. (3) Orientation discrimination showed little or no influence of attention instructions. Thus, whether or not attention influences the processing of second-order motion depends crucially on the precise stimulus parameters tested. Furthermore under appropriate conditions the processing of first-order motion is also influenced by attention, albeit to a lesser extent than second-order motion.

Rules for combining the outputs of local motion detectors to define simple contours.

We know something about the fidelity with which motion can be detected in local regions of the visual field but nothing about how these local motion signals are combined across space to define contours. To investigate such linking rules, we measured the detectability of motion-defined contours using an adaptation of the paradigm of Field, Hayes, and Hess (Vision Research, 33 (1993) 173) in which subjects are asked to detect the presence of simple contours defined solely by local motion direction that are embedded in a field of otherwise random local motions. We show that contours defined by motion whose direction is along the contour are more detectable than contours defined by motions of any common direction. Furthermore, the contour configuration is important in that straight and moderately curved contours, though not highly curved ones, can support this specialized form of motion integration.

Poor encoding of position by contrast-defined motion.

Second-order (contrast-defined) motion stimuli lead to poor performance on a number of tasks, including discriminating form from motion and visual search. To investigate this deficiency, we tested the ability of human observers to monitor multiple regions for motion, to code the relative positions of shapes defined by motion, and to simultaneously encode motion direction and location. Performance with shapes from contrast-defined motion was compared with that obtained from luminance-defined (first-order) stimuli. When the position of coherent motion was uncertain, direction-discrimination thresholds were elevated similarly for both luminance-defined and contrast-defined motion, compared to when the stimulus location was known. The motion of both luminance- and contrast-defined structure can be monitored in multiple visual field locations. Only under conditions that greatly advantaged contrast-defined motion, were observers able to discriminate the positional offset of shapes defined by either type of motion. When shapes from contrast-defined and luminance-defined motion were presented under comparable conditions, the positional accuracy of contrast-defined motion was found to be poorer than its luminance-defined counterpart. These results may explain some, but possibly not all, of the deficits found previously with second-order motion.

Deficits to global motion processing in human amblyopia.

We investigated global motion processing in a group of adult amblyopes using a method that allows us to factor out any influence of the known contrast sensitivity deficit. We show that there are independent global motion processing deficits in human amblyopia that are unrelated to the contrast sensitivity deficit, and that are more extensive for contrast-defined than for luminance-defined stimuli. We speculate that the site of these deficits must include the extra-striate cortex and in particular the dorsal pathway.

The ups and downs of global motion perception: a paradoxical advantage for smaller stimuli in the aging visual system.

Recent evidence suggests that normal aging is typically accompanied by impairment in the ability to perceive the global (overall) motion of visual objects in the world. The purpose of this study was to examine the interplay between age-related changes in the ability to perceive translational global motion (up vs. down) and important factors such as the spatial extent (size) over which movement occurs and how cluttered the moving elements are (density). We used random dot kinematograms (RDKs) and measured motion coherence thresholds (% signal elements required to reliably discriminate global direction) for young and older adults. We did so as a function of the number and density of local signal elements, and the aperture area in which they were displayed. We found that older adults' performance was relatively unaffected by changes in aperture size, the number and density of local elements in the display. In young adults, performance was also insensitive to element number and density but was modulated markedly by display size, such that motion coherence thresholds decreased as aperture area increased (participants required fewer local elements to move coherently to determine the overall image direction). With the smallest apertures tested, young participants' motion coherence thresholds were considerably higher (~1.5 times worse) than those of their older counterparts. Therefore, when RDK size is relatively small, older participants were actually better than young participants at processing global motion. These findings suggest that the normal (disease-free) aging process does not lead to a general decline in perceptual ability and in some cases may be visually advantageous. The results have important implications for the understanding of the consequences of aging on visual function and a number of potential explanations are explored. These include age-related changes in spatial summation, reduced cortical inhibition, neural blur and attentional resource allocation.

The influence of spatial pattern on visual short-term memory for contrast.

Several psychophysical studies of visual short-term memory (VSTM) have shown high-fidelity storage capacity for many properties of visual stimuli. On judgments of the spatial frequency of gratings, for example, discrimination performance does not decrease significantly, even for memory intervals of up to 30 s. For other properties, such as stimulus orientation and contrast, however, such "perfect storage" behavior is not found, although the reasons for this difference remain unresolved. Here, we report two experiments in which we investigated the nature of the representation of stimulus contrast in VSTM using spatially complex, two-dimensional random-noise stimuli. We addressed whether information about contrast per se is retained during the memory interval by using a test stimulus with the same spatial structure but either the same or the opposite local contrast polarity, with respect to the comparison (i.e., remembered) stimulus. We found that discrimination thresholds got steadily worse with increasing duration of the memory interval. Furthermore, performance was better when the test and comparison stimuli had the same local contrast polarity than when they were contrast-reversed. Finally, when a noise mask was introduced during the memory interval, its disruptive effect was maximal when the spatial configuration of its constituent elements was uncorrelated with those of the comparison and test stimuli. These results suggest that VSTM for contrast is closely tied to the spatial configuration of stimuli and is not transformed into a more abstract representation.

Visual motion integration is mediated by directional ambiguities in local motion signals.

The output of primary visual cortex (V1) is a piecemeal representation of the visual scene and the response of any one cell cannot unambiguously guide sensorimotor behavior. It remains unsolved how subsequent stages of cortical processing combine ("pool") these early visual signals into a coherent representation. We (Webb et al., 2007, 2011) have shown that responses of human observers on a pooling task employing broadband, random dot motion can be accurately predicted by decoding the maximum likelihood direction from a population of motion-sensitive neurons. Whereas Amano et al. (2009) found that the vector average velocity of arrays of narrowband, two-dimensional (2-d) plaids predicts perceived global motion. To reconcile these different results, we designed two experiments in which we used 2-d noise textures moving behind spatially distributed apertures and measured the point of subjective equality between pairs of global noise textures. Textures in the standard stimulus moved rigidly in the same direction, whereas their directions in the comparison stimulus were sampled from a set of probability distributions. Human observers judged which noise texture had a more clockwise (CW) global direction. In agreement with Amano and colleagues, observers' perceived global motion coincided with the vector average stimulus direction. To test if directional ambiguities in local motion signals governed perceived global direction, we manipulated the fidelity of the texture motion within each aperture. A proportion of the apertures contained texture that underwent rigid translation and the remainder contained dynamic (temporally uncorrelated) noise to create locally ambiguous motion. Perceived global motion matched the vector average when the majority of apertures contained rigid motion, but with increasing levels of dynamic noise shifted toward the maximum likelihood direction. A class of population decoders utilizing power-law non-linearities can accommodate this flexible pooling.

Binocular summation of second-order global motion signals in human vision.

Although many studies have examined the principles governing first-order global motion perception, the mechanisms that mediate second-order global motion perception remain unresolved. This study investigated the existence, nature and extent of the binocular advantage for encoding second-order (contrast-defined) global motion. Motion coherence thresholds (79.4% correct) were assessed for determining the direction of radial, rotational and translational second-order motion trajectories as a function of local element modulation depth (contrast) under monocular and binocular viewing conditions. We found a binocular advantage for second-order global motion processing for all motion types. This advantage was mainly one of enhanced modulation sensitivity, rather than of motion-integration. However, compared to findings for first-order motion where the binocular advantage was in the region of a factor of around 1.7 (Hess et al., 2007), the binocular advantage for second-order global motion was marginal, being in the region of around 1.2. This weak enhancement in sensitivity with binocular viewing is considerably less than would be predicted by conventional models of either probability summation or neural summation.

Psychophysical correlates of global motion processing in the aging visual system: a critical review.

The consequences of visual decline in aging have a fundamental and wide-reaching impact on age-related quality of life. It is of concern therefore that evidence suggests that normal aging is accompanied by impairments in the ability to effectively encode global motion. Global motion perception is a fundamentally important process. It enables us to determine the overall velocity of spatially extensive objects in the world and provides us with information about our own body movements. Here, we review what is currently known about the effects of age on performance for encoding the global motion information available in random dot kinematograms (RDKs), a class of stimuli widely used to probe the mechanisms underlying motion perception. We conclude that age-related deficits in global motion perception are not all encompassing. Rather, they appear to be specific to certain stimulus conditions. We also examine evidence for an interaction between age and gender and consider the efficacy of techniques such as visual perceptual learning that may attenuate some of the visual deficits in the older adult population.

Temporal frequency modulates reaction time responses to first-order and second-order motion.

This study investigated the effect of temporal frequency and modulation depth on reaction times for discriminating the direction of first-order (luminance-defined) and second-order (contrast-defined) motion, equated for visibility using equal multiples of direction-discrimination threshold. Results showed that reaction times were heavily influenced by temporal frequency, especially in the case of second-order motion. At 1 Hz, reaction times were faster for first-order compared with second-order motion. As temporal frequency increased, reaction times for first-order motion decreased slightly, but those for second-order motion decreased more rapidly. At 8 Hz, reaction times for second-order motion were, in many cases, faster than those for first-order motion. Reaction times decreased as stimulus modulation depth increased at approximately the same rate for both motion types. The findings demonstrate that behavioral response latencies to first-order and second-order motion are dependent on specific stimulus parameters and may, in some cases, be shorter in response to second-order compared with first-order motion.

Interaction between luminance gratings and disparity gratings.

It was shown from geometry and photographic measurement that the shading pattern for a sinusoidal corrugated surface of frequency f approximates to a luminance-defined grating of frequency f, 2f or f + 2f in specific relative phase. It was confirmed that a luminance grating modifies the appearance of a suprathreshold stereoscopic corrugated surface, suggesting an interaction between shading and binocular disparity. Disparity thresholds for detecting random-dot, disparity-defined gratings of spatial frequency 0.2 or 0.4 c/deg were measured in the presence of luminance gratings of spatial frequency 0.4 c/deg with the same orientation. Phase-specific facilitation of disparity thresholds was greatest for a phase relationship inconsistent with shading of a corrugated surface, and was disrupted by positional uncertainty. The presence of texture-defined lines (which served to mark explicitly the successive spatial locations of salient depth features in the image) produced a similar pattern of facilitation, in the absence of shape-from-shading cues. The pattern of results indicates direct local interactions, including spatial cueing, rather than interaction of depth cues.