The role of neural tuning in quantity perception.

Perception of quantities, such as numerosity, timing, and size, is essential for behavior and cognition. Accumulating evidence demonstrates neurons processing quantities are tuned, that is, have a preferred quantity amount, not only for numerosity, but also other quantity dimensions and sensory modalities. We argue that quantity-tuned neurons are fundamental to understanding quantity perception. We illustrate how the properties of quantity-tuned neurons can underlie a range of perceptual phenomena. Furthermore, quantity-tuned neurons are organized in distinct but overlapping topographic maps. We suggest that this overlap in tuning provides the neural basis for perceptual interactions between different quantities, without the need for a common neural representational code.

Adaptation to visual numerosity changes neural numerosity selectivity.

Perceiving numerosity, i.e. the set size of a group of items, is an evolutionarily preserved ability found in humans and animals. A useful method to infer the neural underpinnings of a given perceptual property is sensory adaptation. Like other primary perceptual attributes, numerosity is susceptible to adaptation. Recently, we have shown numerosity-selective neural populations with a topographic organization in the human brain. Here, we investigated whether numerosity adaptation can affect the numerosity selectivity of these populations using ultra-high field (7 Tesla) functional magnetic resonance imaging (fMRI). Participants viewed stimuli of changing numerosity (1 to 7 dots), which allowed the mapping of numerosity selectivity. We interleaved a low or high numerosity adapter stimulus with these mapping stimuli, repeatedly presenting 1 or 20 dots respectively to adapt the numerosity-selective neural populations. We analyzed the responses using custom-build population receptive field neural models of numerosity encoding and compared estimated numerosity preferences between adaptation conditions. We replicated our previous studies where we found several topographic maps of numerosity-selective responses. We found that overall, numerosity adaptation altered the preferred numerosities within the numerosity maps, resulting in predominantly attractive biases towards the numerosity of the adapter. The differential biases could be explained by the difference between the unadapted preferred numerosity and the numerosity of the adapter, with attractive biases being observed with higher difference. The results could link perceptual numerosity adaptation effects to changes in neural numerosity selectivity.

Adaptation reveals unbalanced interaction between numerosity and time.

Processing quantities such as the number of objects in a set, size, spatial arrangement and time is an essential means of structuring the external world and preparing for action. The theory of magnitude suggests that number and time, among other continuous magnitudes, are linked by a common cortical metric, and their specialization develops from a single magnitude system. In order to investigate potentially shared neural mechanisms underlying numerosity and time processing, we used visual adaptation, a method which can reveal the existence of a dedicated processing system. We reasoned that cross-adaptation between numerosity and duration would concur with the existence of a common processing mechanism, whereas the absence of cross-adaptation would provide evidence against it. We conducted four experiments using a rapid adaptation protocol where participants adapted to either visual numerosity or visual duration and subsequently performed a numerosity or duration discrimination task. We found that adapting to a low numerosity altered the estimation of the reference numerosity by an average of 5 dots, compared to adapting to a high numerosity. Similarly, adapting to a short duration altered the estimation of the reference duration by an average of 43 msec, compared to adapting to a long duration. In the cross-dimensional adaptation conditions, duration adaptation altered numerosity estimation by an average of 1 dot, whereas there was not sufficient evidence to either support or reject the effect of numerosity adaptation on duration judgments. These results highlight that there are partially overlapping neural mechanisms which are dedicated for processing both numerosity and time.

Visual Light Zones.

In this article, we studied perception of a particular case of light fields that is characterized by a difference in its consistent structure between parts of a scene. In architectural lighting design, such a consistent structure in a part of a light field is called a light zone. First, we explored whether human observers are sensitive to light zones, that is, zones determined primarily by light flow differences, for a natural-looking scene. We found that observers were able to distinguish the light conditions between the zones. The results suggested an effect of light zones' orientation. Therefore, in Experiment 2, we systematically examined how the orientation of light zones (left-right or front-back) with respect to a viewer influences light inferences in symmetric scenes. We found that observers are quite sensitive to the difference in the light flow of the light zones. In addition, we found that participants showed idiosyncratic behavior, especially for front-back-oriented light zones. Our findings show that observers are sensitive to differences in light field structure between two parts of a scene, which we call visual light zones.

Cortical depth dependent population receptive field attraction by spatial attention in human V1.

Visual spatial attention concentrates neural resources at the attended location. Recently, we demonstrated that voluntary spatial attention attracts population receptive fields (pRFs) toward its location throughout the visual hierarchy. Theoretically, both a feed forward or feedback mechanism could underlie pRF attraction in a given cortical area. Here, we use sub-millimeter ultra-high field functional MRI to measure pRF attraction across cortical depth and assess the contribution of feed forward and feedback signals to pRF attraction. In line with previous findings, we find consistent attraction of pRFs with voluntary spatial attention in V1. When assessed as a function of cortical depth, we find pRF attraction in every cortical portion (deep, center and superficial), although the attraction is strongest in deep cortical portions (near the gray-white matter boundary). Following the organization of feed forward and feedback processing across V1, we speculate that a mixture of feed forward and feedback processing underlies pRF attraction in V1. Specifically, we propose that feedback processing contributes to the pRF attraction in deep cortical portions.

Acute alerting effects of light: A systematic literature review.

Periodic, well timed exposure to light is important for our health and wellbeing. Light, in particular in the blue part of the spectrum, is thought to affect alertness both indirectly, by modifying circadian rhythms, and directly, giving rise to acute effects. We performed a systematic review of empirical studies on direct, acute effects of light on alertness to evaluate the reliability of these effects. In total, we identified 68 studies in which either light intensity, spectral distribution, or both were manipulated, and evaluated the effects on behavioral measures of alertness, either subjectively or measured in reaction time performance tasks. The results show that increasing the intensity of polychromatic white light has been found to increase subjective ratings of alertness in a majority of studies, though a substantial proportion of studies failed to find significant effects, possibly due to small sample sizes or high baseline light intensities. The effect of the color temperature of white light on subjective alertness is less clear. Some studies found increased alertness with higher color temperatures, but other studies reported no detrimental effects of filtering out the short wavelengths from the spectrum. Similarly, studies that used monochromatic light exposure showed no systematic pattern for the effects of blue light compared to longer wavelengths. Far fewer studies investigated the effects of light intensity or spectrum on alertness as measured with reaction time tasks and of those, very few reported significant effects. In general, the small sample sizes used in studies on acute alerting effects of light make it difficult to draw definitive conclusions and better powered studies are needed, especially studies that allow for the construction of dose-response curves.

Depth in box spaces.

Human observers adjust the frontal view of a wireframe box on a computer screen so as to look equally deep and wide, so that in the intended setting the box looks like a cube. Perspective cues are limited to the size-distance effect, since all angles are fixed. Both the size on the screen, and the viewing distance from the observer to the screen were varied. All observers prefer a template view of a cube over a veridical rendering, independent of picture size and viewing distance. If the rendering shows greater or lesser foreshortening than the template, the box appears like a long corridor or a shallow slab, that is, like a 'deformed' cube. Thus observers ignore 'veridicality'. This does not fit an 'inverse optics' model. We discuss a model of 'vision as optical user interface'.

Perception of 3D Slant Out of the Box.

Evidence for contextual effects is widespread in visual perception. Although this suggests that contextual effects are the result of a generic property of the visual system, current explanations are limited to the domain in which they occur. In this paper we propose a more general mechanism of global influences on the perception of slant. We review empirical data and evaluate proposed explanations of contextual biases. By assessing not only a model about three-dimensional slant perception but also evaluating more generic mechanisms of contextual modulation, we show that surround suppression of neural responses explains the major phenomena in the empirical data on contextual biases. Moreover, contextual biases may be part of a mechanism of grouping and segmentation.

Detecting gradual visual changes in colour and brightness agnosia: a double dissociation.

Two patients, one with colour agnosia and one with brightness agnosia, performed a task that required the detection of gradual temporal changes in colour and brightness. The results for these patients, who showed anaverage or an above-average performance on several tasks designed to test low-level colour and luminance (contrast) perception in the spatial domain, yielded a double dissociation; the brightness agnosic patient was within the normal range for the coloured stimuli, but much slower to detect brightness differences, whereas the colour agnosic patient was within the normal range for the achromatic stimuli, but much slower for the coloured stimuli. These results suggest that a modality-specific impairment in the detection of gradual temporal changes might be related to, if not underlie, the phenomenon of visual agnosia.

Interactions between colour and synaesthetic colour: an effect of simultaneous colour contrast on synaesthetic colours.

We investigated whether simultaneous colour contrast affects the synaesthetic colour experience and normal colour percept in a similar manner. We simultaneously presented a target stimulus (i.e. grapheme) and a reference stimulus (i.e. hash). Either the grapheme or the hash was presented on a saturated background of the same or opposite colour category as the synaesthetic colour and the other stimulus on a grey background. In both conditions, grapheme-colour synaesthetes were asked to colour the hash in a colour similar to the synaesthetic colour of the grapheme. Controls that were pair-matched to the synaesthetes performed the same experiment, but for them, the grapheme was presented in the colour induced by the grapheme in synaesthetes. When graphemes were presented on a grey and the hash on a coloured background, a traditional simultaneous colour-contrast effect was found for controls as well as synaesthetes. When graphemes were presented on colour and the hash on grey, the controls again showed a traditional simultaneous colour-contrast effect, whereas the synaesthetes showed the opposite effect. Our results show that synaesthetic colour experiences differ from normal colour perception; both are susceptible to different surrounding colours, but not in a comparable manner.

Shape contrast: a global mechanism?

We investigated whether a shape contrast bias is caused by local contrast enhancement or by a global mechanism. In a baseline condition, observers performed a shape discrimination task on an isolated hinged plane. But in the experimental conditions, five dihedral surfaces, of which we varied the dihedral angle distribution, were added on each side. Shape perception was influenced not only by the adjacent surface but also by the mean of the shape distribution in the extended surround. Thus, shape contrast is not locally determined and has to be understood from a global mechanism. We propose divisive normalization of shape signals as such a mechanism.

Uncertainty reveals surround modulation of shape.

Noisy estimations of shape can be partially resolved by incorporating relevant information from the context. The effect of surround stimuli on shape perception becomes clear in illusions of shape contrast and assimilation. In this study, we answer the question how a surround-induced bias depends on the reliability of shape signals. This way, we assess the processes by which an observer incorporates relevant data from the context into the shape estimate. We selectively added visual noise to the center and surround and compared a bias in shape perception with a control condition where no noise was added. In the conditions where shape and surround stimuli were well defined, we found a shape-contrast bias. When the surround stimuli were degraded, this contrast bias decreased. Most interestingly, when the central shape was degraded, an assimilation bias was observed. This bias was larger when the entire stimulus was degraded compared to when only the central shape was degraded. This suggests that shape contrast is the result of inference processes relying on local representations in early visual areas whereas assimilation is related to inference processes by global representations in higher visual areas.

Perception of 3D shape in context: contrast and assimilation.

Whereas integration of shape and surround is held to occur through cue-dependent representations, we show that both cue-invariant and cue-dependent representations are involved. A central hinged plane and larger flanking plane were defined by either binocular disparity or motion. In a 'within-cue' condition, shape and surround were defined by the same cue and in a 'cross-cue' condition they were defined by a different cue. Observers compared the dihedral angle of the central shape with a constant reference. When the central shape was defined by disparity, the surround stimuli invoked a contrast bias in the within-cue condition, but shape assimilation occurred in the cross-cue condition. When the central shape was defined by motion there were overall no significant results, but if a contrast bias was observed, it was in the within-cue condition where integration could occur through cue-dependent representations.

Adaptive center-surround interactions in human vision revealed during binocular rivalry.

We used binocular rivalry as a psychophysical probe to explore center-surround interactions in orientation, motion and color processing. Addition of the surround matching one of the rival targets dramatically altered rivalry dynamics. For all visual sub-modalities tested, predominance of the high-contrast rival target matched to the surround was greatly reduced-a result that disappeared at low contrast. At low contrast, addition of the surround boosted dominance of orientation and motion targets matched to the surround. This contrast-dependent modulation of center-surround interactions seems to be a general property of the visual system and may reflect an adaptive balance between surround suppression and spatial summation.

Material-illumination ambiguities and the perception of solid objects.

The appearance of objects depends on their material, shape, and on the illumination conditions. Conversely, object appearance provides us with cues about the illumination and the material. This so-called inverse problem is basically underdetermined and therefore we expect that material and illumination perception are confounded. To gain insight into the relevant mechanisms, we rendered a set of artificial spheres for vastly different canonical light fields and reflectance functions. We used four physics-based bidirectional reflectance distribution functions (BRDFs) representing glossy, pitted, velvety, and matte material. The six illumination conditions were collimated illumination from four directions, hemispherical diffuse illumination, and fully diffuse (Ganzfeld) illumination. In three sub-experiments we presented pairs of stimuli and asked human observers to judge whether the material was the same, whether the illumination was the same, and for a subset in which either the illumination or the material was the same to judge which of the two was constant. We found that observers made many errors in all sub-experiments. In experiment 2 the illumination direction was chosen at random. Using an interactive interface, we asked human observers to match the illumination direction of a sphere of one of the four materials with that of a Lambertian sphere. We found systematical material-dependent deviations from veridical performance. Theoretical analysis of the radiance patterns suggests that judgments were based mainly on the position of the shadow edge. In conclusion, we found no evidence for 'material constancy' for perception of smooth rendered spheres despite vast quantitative and qualitative differences in illumination and in BRDF between the stimuli. Although human observers demonstrated some 'illumination constancy', they made systematic errors depending on the material reflectance, suggesting that they used mainly the location of the shadow edge. Our results suggest that material perception and light-field perception are basically confounded.

Center-surround inhibition and facilitation as a function of size and contrast at multiple levels of visual motion processing.

Visual context often plays a crucial role in visual processing. In the domain of visual motion processing, the response to a stimulus presented to a neuron's classical receptive field can be modulated by presenting stimuli to its surround. The nature of these center-surround interactions is often inhibitory; the neural response decreases when the same direction of motion is presented to center and surround. Here we use binocular rivalry as a tool to study center-surround interactions. We show that magnitude of surround suppression varies as a function of luminance contrast and surround width. Increasing the size of surround motion increased surround suppression at high contrast. Furthermore, large, high-contrast surrounds facilitated opposite-direction motion in the center. For stimuli presented at low contrast, surround suppression peaked at a smaller surround width. In addition, we provide evidence that surround inhibition occurs at multiple levels of visual processing: Surround inhibition in motion processing is likely to originate from both monocular and binocular processing stages.

Center-surround interactions in visual motion processing during binocular rivalry.

When each eye is confronted with a dissimilar stimulus, the percept will generally alternate between the two. This phenomenon is known as binocular rivalry. Although binocular rivalry occurs at locations where targets overlap spatially, the area surrounding rivalrous targets can modulate their dominance. Here we show that during binocular rivalry of oppositely moving gratings, a surrounding grating moving in the same direction as one of the two leads to increased dominance of the opposite direction of motion in the center. This increased dominance of the opposite direction in the center was observed irrespective of the eye to which the surround was presented. Inspection of the results for different conditions reveals that the preference for the opposite direction of motion cannot be explained by a single mechanism operating beyond binocular fusion. We therefore suggest that this phenomenon is the outcome of center-surround interactions at multiple levels along the pathway of visual motion processing.

Visual discrimination of spectral distributions.

Human observers seem to be able to use different features that classify materials with a large degree of accuracy. In this paper, we look at human perception of statistical properties of the spectral distribution in a scene. We investigated whether human observers can discriminate just as accurately between coloured textures that have a spectral distribution due either to shading only or to both shading and specular reflectance as between uniform colours. Thresholds for the discrimination of coloured textures are about 15 times as high as thresholds for the discrimination of uniform colours, provided there is a sharp transition between the two colours. However, the coloured texture thresholds are only 1.5 times higher when we introduce a gradual transition between the two colours. There are also distinct qualitative differences in discrimination thresholds for different base colours. These differences cannot be predicted from discrimination thresholds for uniform colours. Human observers are surprisingly good at discriminating between a material edge and a shadow edge in complex scenes. Statistical differences in the orientation of the colour distributions in colour space might be used to accomplish this. In a second experiment we investigated how well observers can discriminate between two linear distributions in colour space that have the same base colour but different orientations. When we vary the line-length in R, G, B space, thresholds cannot be predicted completely by the conservation of the average distance between the two distributions. This means that observers use not only the maximum colour difference in the stimulus to do the task, but other cues are also involved.

Illumination direction from texture shading.

We investigate the ability of human observers to judge the direction of illumination from image texture. Photographs of 61 real surfaces were used, taken from the Columbia-Utrecht Reflectance and Texture (Curet) database (http:/www.cs.columbia.edu/CAVE/curet). All samples were normally viewed but obliquely illuminated, the elevation of the source being 22.5 degrees, 45.0 degrees, or 67.5 degrees. The illumination was with a collimated, parallel beam. Stimuli were presented in random orientation, and observers had to judge both the elevation and the azimuth of the source. Observers judged the azimuth within approximately 15 degrees, except for the fact that they committed random (with approximately 50% probability) sign flips (180 degrees flips). Connected with this finding is the fact that observers judged the illumination to be from above rather than below in the overwhelming majority of cases, despite the fact that each case occurred with equal probability. The elevation of the illumination can be judged to some extent but is not far above chance level. The data are in good agreement with a simple model that bases the estimate of illumination direction on the second-order statistics of local luminance gradients. This locates the locus of the probable mechanism very early in the visual stream.