Transient attention enhances perceptual performance and FMRI response in human visual cortex

When a visual stimulus suddenly appears, it captures attention, producing a transient improvement of performance on basic visual tasks. We investigate the effect of transient attention on stimulus representations in early visual areas using rapid event-related fMRI. Participants discriminated the orientation of one of two gratings preceded or followed by a nonpredictive peripheral cue. Compared to control conditions, precueing the target location improved performance and produced a larger fMRI response in corresponding retinotopic areas. This enhancement progressively increased from striate to extrastriate areas. Control conditions indicated that the enhanced fMRI response was not due to sensory summation of cue and target signals. Thus, an uninformative precue increases both perceptual performance and the concomitant stimulus-evoked activity in early visual areas. These results provide evidence regarding the retinotopically specific neural correlate for the effects of transient attention on early vision.     

Three-dimensional shape representation in monkey cortex

Using fMRI in anesthetized monkeys, this study investigates how the primate visual system constructs representations of three-dimensional (3D) shape from a variety of cues. Computer-generated 3D objects defined by shading, random dots, texture elements, or silhouettes were presented either statically or dynamically (rotating). Results suggest that 3D shape representations are highly localized, although widely distributed, in occipital, temporal, parietal, and frontal cortices and may involve common brain regions regardless of shape cue. This distributed network of areas cuts across both "what" and "where" processing streams, reflecting multiple uses for 3D shape representation in perception, recognition, and action.     

Effects of Spatial Frequency Similarity and Dissimilarity on Contour Integration

We examined the effects of spatial frequency similarity and dissimilarity on human contour integration under various conditions of uncertainty. Participants performed a temporal 2AFC contour detection task. Spatial frequency jitter up to 3.0 octaves was applied either to background elements, or to contour and background elements, or to none of both. Results converge on four major findings. (1) Contours defined by spatial frequency similarity alone are only scarcely visible, suggesting the absence of specialized cortical routines for shape detection based on spatial frequency similarity. (2) When orientation collinearity and spatial frequency similarity are combined along a contour, performance amplifies far beyond probability summation when compared to the fully heterogenous condition but only to a margin compatible with probability summation when compared to the fully homogenous case. (3) Psychometric functions are steeper but not shifted for homogenous contours in heterogenous backgrounds indicating an advantageous signal-to-noise ratio. The additional similarity cue therefore not so much improves contour detection performance but primarily reduces observer uncertainty about whether a potential candidate is a contour or just a false positive. (4) Contour integration is a broadband mechanism which is only moderately impaired by spatial frequency dissimilarity.     

Performance data indicate summation for pictorial depth-cues in slanted surfaces

Over recent years much has been learned about the way in which depth cues are combined (e.g. Landy et al.. 1995). The majority of this work has used subjective measures, a rating scale or a point of subjective equality, to deduce the relative contributions of different cues to perception. We have adopted a very different approach by using two interval forced-choice (21FC) performance measures and a signal processing framework. We performed summation experiments for depth cue increment thresholds between pairs of pictorial depth cues in displays depicting slanted planar surfaces made from arrays of circular 'contrast' elements. Summation was found to be ideal when size-gradient was paired with contrast-gradient for a wide range of depth-gradient magnitudes in the null stimulus. For a pairing of size-gradient and linear perspective, substantial summation (> 1.5 dB) was found only when the null stimulus had intermediate depth gradients; when flat or steeply inclined surfaces were depicted, summation was diminished or abolished. Summation was also abolished when one of the target cues was (i) not a depth cue, or (ii) added in conflict. We conclude that vision has a depth mechanism for the constructive combination of pictorial depth cues and suggest two generic models of summation to describe the results. Using similar psychophysical methods. Bradshaw and Rogers (1996) revealed a mechanism for the depth cues of motion parallax and binocular disparity. Whether this is the same or a different mechanism from the one reported here awaits elaboration.     

Strength of feature contrast mediates interaction among feature domains

Traditional theories of texture segregation suggest that elementary visual features are processed in parallel by independent modules at early visual stages. Here we show that, for small feature contrasts and large values evoking perceptual popout, different forms of module interaction exist. While discrimination of highly salient features rests on independent feature specific pathways, information is summed across domains when barely noticeable ones are to be detected in homogeneous textures.     

Mislocalization of visual stimuli: independent effects of static and dynamic attention

Shifts of visual attention cause systematic distortions of the perceived locations of visual objects around the focus of attention. In the attention repulsion effect, the perceived location of a visual target is shifted away from an attention-attracting cue when the cue is presented before the target. Recently it has been found that, if the visual cue is presented after the target, the perceived location of the target shifts toward the location of the following cue. One unanswered question is whether a single mechanism underlies both attentional repulsion and attraction effects. We presented participants with two disks at diagonal locations as visual cues and two vertical lines as targets. Participants were asked to perform a forced-choice task to judge targets' positions. The present study examined whether the magnitude of the repulsion effect and the attraction effect would differ (Experiment 1), whether the two effects would interact (Experiment 2), and whether the location or the dynamic shift of attentional focus would determine the distortions effects (Experiment 3). The results showed that the effect size of the attraction effect was slightly larger than the repulsion effect and the preceding and following cues have independent influences on the perceived positions. The repulsion effect was caused by the location of attnetion and the attraction effect was due to the dynamic shift of attentional focus, suggesting that the underlying mechanisms for the retrospective attraction effect might be different from those for the repulsion effect.     

A general rule for sensory cue summation: evidence from photographic, musical, phonetic and cross-modal stimuli

The Euclidean and MAX metrics have been widely used to model cue summation psychophysically and computationally. Both rules happen to be special cases of a more general Minkowski summation rule , where m = 2 and ∞, respectively. In vision research, Minkowski summation with power m = 3-4 has been shown to be a superior model of how subthreshold components sum to give an overall detection threshold. Recently, we have previously reported that Minkowski summation with power m = 2.84 accurately models summation of suprathreshold visual cues in photographs. In four suprathreshold discrimination experiments, we confirm the previous findings with new visual stimuli and extend the applicability of this rule to cue combination in auditory stimuli (musical sequences and phonetic utterances, where m = 2.95 and 2.54, respectively) and cross-modal stimuli (m = 2.56). In all cases, Minkowski summation with power m = 2.5-3 outperforms the Euclidean and MAX operator models. We propose that this reflects the summation of neuronal responses that are not entirely independent but which show some correlation in their magnitudes. Our findings are consistent with electrophysiological research that demonstrates signal correlations (r = 0.1-0.2) between sensory neurons when these are presented with natural stimuli.     

Second-Order Conditioning and Conditioned Inhibition: Influences of Speed versus Accuracy on Human Causal Learning

In human causal learning, excitatory and inhibitory learning effects can sometimes be found in the same paradigm by altering the learning conditions. This study aims to explore whether learning in the feature negative paradigm can be dissociated by emphasising speed over accuracy. In two causal learning experiments, participants were given a feature negative discrimination in which the outcome caused by one cue was prevented by the addition of another. Participants completed training trials either in a self-paced fashion with instructions emphasising accuracy, or under strict time constraints with instructions emphasising speed. Using summation tests in which the preventative cue was paired with another causal cue, participants in the accuracy groups correctly rated the preventative cue as if it reduced the probability of the outcome. However, participants in the speed groups rated the preventative cue as if it increased the probability of the outcome. In Experiment 1, both speed and accuracy groups later judged the same cue to be preventative in a reasoned inference task. Experiment 2 failed to find evidence of similar dissociations in retrospective revaluation (release from overshadowing vs. mediated extinction) or learning about a redundant cue (blocking vs. augmentation). However in the same experiment, the tendency for the accuracy group to show conditioned inhibition and the speed group to show second-order conditioning was consistent even across sub-sets of the speed and accuracy groups with equivalent accuracy in training, suggesting that second-order conditioning is not merely a consequence of poorer acquisition. This dissociation mirrors the trade-off between second-order conditioning and conditioned inhibition observed in animal conditioning when training is extended.     

Conjunctions of colour, luminance and orientation: the role of colour and luminance contrast on saliency and proximity grouping in texture segregation

To examine whether perceptual grouping on the basis of orientation can be performed simultaneously with or only subsequently to grouping according to colour or luminance, we tested whether subjects are able to segregate arrays of texture elements that differ from surrounding elements by conjunctions of either (i) colour and orientation, or (ii) luminance contrast and orientation, or (iii) luminance contrast polarity and orientation. Subjects were able to use conjunctions between luminance and orientation for segregation but not conjunctions between colour or contrast polarity and orientation. Our results suggest that (i) in agreement with earlier findings, there seem to exist no specific conjunction detectors for colour and orientation or contrast polarity and orientation, and (ii) when orientation defined textures are to be distinguished by virtue of differences in luminance, colour, or contrast polarity, luminance provides a much stronger cue than colour or contrast polarity for saliency-based orientation grouping.     

US specificity of occasion setting: Hierarchical or configural learning?

Four experiments in rats examined whether occasion setters and target CSs play qualitatively different roles in occasion-setting discriminations. Two visual occasion setters, A and B, signalled reinforcement of two auditory target CSs, x and y, with sucrose and oil (A…x→suc, B…y→oil, A-, B-, x-, y-); in addition two transfer CSs w and z were paired with sucrose and oil (w→suc, z→oil). When w and z were substituted for x and y (A…w, B…w, A…z, B…z) more responding was observed when both stimuli had been paired with the same outcome (Experiments 1 and 3a). No effect was observed when two visual "pseudo-occasion setters", C and D (paired with sucrose and oil in a trace relation to the US:C…→suc, D…→oil), were substituted for the occasion setters A and B (C…x, D…x, C…y, D…y; Experiments 2, 3b and 4). These results could not be explained in terms of Pavlovian summation: responding to combinations of Pavlovian CSs paired with same or different outcomes was either the same, or lower when both stimuli had been paired with the same outcome (Experiment 4). Implications of these results for theories of occasion setting and configural learning are discussed.     

Area summation in human vision at and above detection threshold

The initial image-processing stages of visual cortex are well suited to a local (patchwise) analysis of the viewed scene. But the world's structures extend over space as textures and surfaces, suggesting the need for spatial integration. Most models of contrast vision fall shy of this process because (i) the weak area summation at detection threshold is attributed to probability summation (PS) and (ii) there is little or no advantage of area well above threshold. Both of these views are challenged here. First, it is shown that results at threshold are consistent with linear summation of contrast following retinal inhomogeneity, spatial filtering, nonlinear contrast transduction and multiple sources of additive Gaussian noise. We suggest that the suprathreshold loss of the area advantage in previous studies is due to a concomitant increase in suppression from the pedestal. To overcome this confound, a novel stimulus class is designed where: (i) the observer operates on a constant retinal area, (ii) the target area is controlled within this summation field, and (iii) the pedestal is fixed in size. Using this arrangement, substantial summation is found along the entire masking function, including the region of facilitation. Our analysis shows that PS and uncertainty cannot account for the results, and that suprathreshold summation of contrast extends over at least seven target cycles of grating.     

Lack of evidence for inhibitory processes in over-selectivity

Stimulus over-selectivity can be defined as control over behavior being exerted by one aspect of the environment at the expense of other equally salient aspects of the environment, and is a common problem for discrimination learning under conditions of cognitive strain, and in intellectual disorders. Non-clinical participants exposed to a concurrent task load were trained and tested on a two-component trial-and-error discrimination task to investigate whether inhibition plays a role in producing under-selectivity by using both summation and retardation tests. Experiment 1 found evidence for the over-selectivity effect, and replicated the finding that revaluation of a previously over-selected stimulus allows emergence of control by a previous under-selected stimulus, despite the latter stimulus receiving no direct conditioning. The under-selected cue was not found to gain any conditioned inhibitory status, as reflected by summation (Experiment 2), and retardation (Experiment 3), tests. The results extend the literature explaining over-selectivity as a post-acquisition failure.     

On the organization of memory in the optomotor system of the crab pachygrapsus crassipes

Memory responses of the optomotor system to rotations of various stripe patterns were studied. The separate elements of the visual background are individually remembered in terms of the parts of the eye on which their images fell. A visual illusion resulting from this property is described. All parts of the retina have an equal capacity to contribute to memory. The memory response results from the summation of contributions from individual elements rather than the maintenance of a fixation upon any particular feature of the situation. Both the separation between background elements for angles from 6° up to 60° and the number of elements present affect the size of the memory evoked response.     

On the organization of memeory in the optomotor system of the crab Pachygrapsus crassipes.

Memory responses of the optomotor system to rotations of various stripe patterns were studied. The separate elements of the visual background are individually remembered in terms of the parts of the eye on which their images fell. A visual illusion resulting from this property is described. All parts of the retina have an equal capacity to contribute to memory. The memory response results from the summation of contributions from individual elements rather than the maintenance of a fixation upon any particular feature of the situation. Both the separation between background elements for angles from 6 degrees up to 60 degrees and the number of elements present affect the size of the memory evoked response.     

Long-range, pattern-dependent contextual effects in early human visual cortex

The standard view of neurons in early visual cortex is that they behave like localized feature detectors. Here we demonstrate that processing in early visual areas goes beyond feature detection by showing that neural responses are greater when a feature deviates from its context compared to when it does not deviate from its context. Using psychophysics, fMRI, and electroencephalography methodologies, we measured neural responses to an oriented Gabor ("target") embedded in various visual patterns as defined by the relative orientation of flanking stimuli. We first show using psychophysical contrast adaptation and fMRI that a target that differs from its context results in more neural activity compared to a target that is contained within an alternating sequence, suggesting that neurons in early visual cortex are sensitive to large-scale orientation patterns. Next, we use event-related potentials to show that orientation deviations affect the earliest sensory components of the target response. Finally, we use forced-choice classification of "noise" stimuli to show that we are more likely to "see" orientations that deviate from the context. Our results suggest that early visual cortex is sensitive to global patterns in images in a way that is markedly different from the predictions of standard models of cortical visual processing.     

Stimulus saliency modulates pre-attentive processing speed in human visual cortex

The notion of a saliency-based processing architecture [1] underlying human vision is central to a number of current theories of visual selective attention [e.g., 2]. On this view, focal-attention is guided by an overall-saliency map of the scene, which integrates (sums) signals from pre-attentive sensory feature-contrast computations (e.g., for color, motion, etc.). By linking the Posterior Contralateral Negativity (PCN) component to reaction time (RT) performance, we tested one specific prediction of such salience summation models: expedited shifts of focal-attention to targets with low, as compared to high, target-distracter similarity. For two feature-dimensions (color and orientation), we observed decreasing RTs with increasing target saliency. Importantly, this pattern was systematically mirrored by the timing, as well as amplitude, of the PCN. This pattern demonstrates that visual saliency is a key determinant of the time it takes for focal-attention to be engaged onto the target item, even when it is just a feature singleton.     

Summation of perceptual cues in natural visual scenes

Natural visual scenes are rich in information, and any neural system analysing them must piece together the many messages from large arrays of diverse feature detectors. It is known how threshold detection of compound visual stimuli (sinusoidal gratings) is determined by their components' thresholds. We investigate whether similar combination rules apply to the perception of the complex and suprathreshold visual elements in naturalistic visual images. Observers gave magnitude estimations (ratings) of the perceived differences between pairs of images made from photographs of natural scenes. Images in some pairs differed along one stimulus dimension such as object colour, location, size or blur. But, for other image pairs, there were composite differences along two dimensions (e.g. both colour and object-location might change). We examined whether the ratings for such composite pairs could be predicted from the two ratings for the respective pairs in which only one stimulus dimension had changed. We found a pooling relationship similar to that proposed for simple stimuli: Minkowski summation with exponent 2.84 yielded the best predictive power (r=0.96), an exponent similar to that generally reported for compound grating detection. This suggests that theories based on detecting simple stimuli can encompass visual processing of complex, suprathreshold stimuli.     

Mapping the arrestin-receptor interface. Structural elements responsible for receptor specificity of arrestin proteins

Arrestins selectively bind to phosphorylated activated forms of their cognate G protein-coupled receptors. Arrestin binding prevents further G protein activation and often redirects signaling to other pathways. The comparison of the high-resolution crystal structures of arrestin2, visual arrestin, and rhodopsin as well as earlier mutagenesis and peptide inhibition data collectively suggest that the elements on the concave sides of both arrestin domains most likely participate in receptor binding directly, thereby dictating its receptor preference. Using comparative binding of visual arrestin/arrestin2 chimeras to the preferred target of visual arrestin, light-activated phosphorylated rhodopsin (PRh*), and to the arrestin2 target, phosphorylated activated m2 muscarinic receptor (P-m2 mAChR*), we identified the elements that determine the receptor specificity of arrestins. We found that residues 49-90 (beta-strands V and VI and adjacent loops in the N-domain) and 237-268 (beta-strands XV and XVI in the C-domain) in visual arrestin and homologous regions in arrestin2 are largely responsible for their receptor preference. Only 35 amino acids (22 of which are nonconservative substitutions) in the two elements are different. Simultaneous exchange of both elements between visual arrestin and arrestin2 fully reverses their receptor specificity, demonstrating that these two elements in the two domains of arrestin are necessary and sufficient to determine their preferred receptor targets.     

Occasion setting in Pavlovian ambiguous target discriminations

Rats were trained in a Pavlovian serial ambiguous target discrimination, in which a target cue was reinforced if it was preceded by one stimulus (P-->T+) but was not reinforced if it was preceded by another stimulus (N-->T-). Test performance indicated that stimulus control by these features was weaker than that acquired by features trained within separate serial feature positive (P-->T+, T-) and serial feature negative (N-->W-, W+) discriminations. The form of conditioned responding and the patterns of transfer observed suggested that the serial ambiguous target discrimination was solved by occasion setting. The data are discussed in terms of the use of retrospective coding strategies when solving Pavlovian serial conditional discriminations, and the acquisition of special properties by both feature and target stimuli.     

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.