Spontaneous perception of numerosity in pre-school children.

There is strong evidence that humans can make rough estimates of the numerosity of a set of items, almost from birth. However, as numerosity covaries with many non-numerical variables, the idea of a direct number sense has been challenged. Here we applied two different psychophysical paradigms to demonstrate the spontaneous perception of numerosity in a cohort of young pre-school children. The results of both tasks showed that even at that early developmental stage, humans spontaneously base the perceptual choice on numerosity, rather than on area or density. Precision in one of these tasks predicted mathematical abilities. The results reinforce strongly the idea of a primary number sense and provide further evidence linking mathematical skills to the sensory precision of the spontaneous number sense, rather than to mechanisms involved in handling explicit numerosity judgements or extensive exposure to mathematical teaching.

Adaptation to number operates on perceived rather than physical numerosity.

Humans share with many animals a number sense, the ability to estimate rapidly the approximate number of items in a scene. Recent work has shown that like many other perceptual attributes, numerosity is susceptible to adaptation. It is not clear, however, whether adaptation works directly on mechanisms selective to numerosity, or via related mechanisms, such as those tuned to texture density. To disentangle this issue we measured adaptation of numerosity of 10 pairs of connected dots, as connecting dots makes them appear to be less numerous than unconnected dots. Adaptation to a 20-dot pattern (same number of dots as the test) caused robust reduction in apparent numerosity of the connected-dot pattern, but not of the unconnected dot-pattern. This suggests that adaptation to numerosity, at least for relatively sparse dot-pattern, occurs at neural levels encoding perceived numerosity, rather than at lower levels responding to the number of elements in the scene.

A low-cost and versatile system for projecting wide-field visual stimuli within fMRI scanners.

We have constructed and tested a custom-made magnetic-imaging-compatible visual projection system designed to project on a very wide visual field (~80°). A standard projector was modified with a coupling lens, projecting images into the termination of an image fiber. The other termination of the fiber was placed in the 3-T scanner room with a projection lens, which projected the images relayed by the fiber onto a screen over the head coil, viewed by a participant wearing magnifying goggles. To validate the system, wide-field stimuli were presented in order to identify retinotopic visual areas. The results showed that this low-cost and versatile optical system may be a valuable tool to map visual areas in the brain that process peripheral receptive fields.

Temporal integration of optic flow, measured by contrast and coherence thresholds.

We measured, as a function of exposure duration, contrast sensitivity and coherence sensitivity for discerning the direction of motion of random dot patterns moving in circular, radial or translational directions. Contrast sensitivity for these patterns increased linearly with exposure duration, up to about 200-300 ms, consistent with previous estimates of temporal summation of early motion units. Coherence sensitivity, however, showed much longer summation periods, about 3 s. When the stimulus was embedded within 10 s of noise, sensitivity increased with duration up to 2-3 s, approximately linearly, as expected from an ideal integrator. When presented without the noise period, sensitivity also increased, but in a different way. For radial and circular motion the increase tended towards the theoretically predicted square root relationship for the same duration as that found with the embedded noise (about 3 s). For translation, however, the curve was steeper than the theoretical prediction (nearly linear), and the summation estimates of around 1000 ms. When the duration of the target was constant at 200 ms, but that of the flanking noise varied, sensitivity decreased with total duration over a similar interval. We interpret our results to reflect at least two stages of analysis, a threshold-limited early stage of local-motion analysis, with a time constant of 200-300 ms, and a later global-motion integration stage with a much longer time constant, around 3000 ms. There may also exist an intermediate stage, with an integration time of around 1000 ms.

Separate visual representations for perception and action revealed by saccadic eye movements.

Some 30 years ago, Trevarthen [1] introduced the idea of two separate visual systems, a focal system for fine motor acts and an ambient system for gross body movements such as ambulation. More recent developments indicating anatomically and physiologically separate pathways in primate vision [2] have led to a different idea of separate visual systems, one for conscious perception and one for action [3]. It has received empirical support from several studies showing that pointing, reaching, and grasping can remain accurate while the perceived position or size of objects is subject to illusory distortion [4-6]. However, much of this evidence has been challenged on the grounds of methodological flaws, particularly failure to match perfectly the conditions for verbal and motor tasks and failure to replicate results [7-10]. Here we take advantage of the strong compression of perceived position that occurs around the time of saccadic eye movements [11, 12]. Under normal lighting conditions, stimuli flashed briefly over a wide range of spatial positions just before saccadic onset are neither seen nor reached for in their veridical positions, but are compressed toward the saccadic target. We validate the idea of separate systems by showing that, in the dark, subjects are able to point accurately to the correct target position, even though their verbal reports are still subject to compression.

Dependency of reaction times to motion onset on luminance and chromatic contrast.

We measured reaction times for detecting the onset of motion of sinusoidal gratings of 1 c/deg, modulated in either luminance or chromatic contrast, caused to move abruptly at speeds ranging from 0.25 to 10 deg/s (0.25-10 Hz). At any given luminance or chromatic contrast, RTs varied linearly with temporal periodicity (r2 congruent with 0.97), yielding a Weber fraction of period. The value of the Weber fraction varied inversely with contrast, differently for luminance and chromatic contrast. The results were well simulated with a simple model that accumulated change in contrast over time until a critical threshold had been reached. Two crucial aspects of the model are a second-stage temporal integration mechanism, capable of accumulating information for periods of up to 2 s, and contrast gain control, different for luminance than for chromatic stimuli. The contrast response for luminance shows very low semi-saturating contrasts and high gain, similar to LGN M-cells and cells in MT; that for colour shows high semi-saturating contrasts and low gain, similar to LGN P-cells. The results suggest that motion onset for luminance and chromatic gratings are detected by different mechanisms, probably by the magno- and parvo-cellular systems.

Cardinal axes for radial and circular motion, revealed by summation and by masking.

Both electro-physiological and psychophysical studies point to the existence of detectors specialised for the analysis of optic flow. However, it is unclear whether these detectors are tuned to specific 'cardinal directions' (such as radial and circular motion), or whether they respond equally to all directions of optic-flow motion, including intermediate spiral motions. Here summation and masking studies of motion coherence sensitivity are reported that suggest that optic flow may be tuned to radial and circular cardinal directions. Strong summation was found between two orthogonal directions of spiral motion, but much weaker summation between radial and circular motion. As orthogonal spiral motions always contain a common radial or circular component, the stronger summation for these motions implies that detectors are tuned to radial and circular directions. Similarly, the most effective masking stimuli (placed adjacent to but not superimposed on the test stimuli) tended to be those in the radial or circular directions, even for spiral targets, further suggesting that flow-field motion is detected and discriminated by mechanisms tuned to these 'cardinal' directions.

Changes in visual perception at the time of saccades.

We frequently reposition our gaze by making rapid ballistic eye movements that are called saccades. Saccades pose problems for the visual system, because they generate rapid, large-field motion on the retina and change the relationship between the object position in external space and the image position on the retina. The brain must ignore the one and compensate for the other. Much progress has been made in recent years in understanding the effects of saccades on visual function and elucidating the mechanisms responsible for them. Evidence suggests that saccades trigger two distinct neural processes: (1) a suppression of visual sensitivity, specific to the magnocellular pathway, that dampens the sensation of motion and (2) a gross perceptual distortion of visual space in anticipation of the repositioning of gaze. Neurophysiological findings from several laboratories are beginning to identify the neural substrates involved in these effects.

A cortical area that responds specifically to optic flow, revealed by fMRI.

The continuously changing optic flow on the retina provides information about direction of heading and about the three-dimensional structure of the environment. Here we use functional magnetic resonance imaging (fMRI) to demonstrate that an area in human cortex responds selectively to components of optic flow, such as circular and radial motion. This area is within the region commonly referrred to as V5/MT complex, but is distinct from the part of this region that responds to translation. The functional properties of these two areas of the V5/MT complex are also different; the response to optic flow was obtained only with changing flow stimuli, whereas response to translation occurred during exposure to continuous motion.

Feature-based integration of orientation signals in visual search.

We have measured orientation discrimination in the presence of a variable number of neutral distracters for two distinct tasks: identification of the orientation of a tilted target and location of its position. Both tasks were performed in the presence of visual noise of variable contrasts. Under a range of conditions, subjects could identify the direction of target tilt at thresholds well below those necessary to locate its position. The location thresholds showed only weak dependency on set-size, consistent with a stimulus uncertainty of parallel search of the output of independent orientation analysers, while the identification thresholds showed a much stronger dependency, varying with the square root of set-size over a wide range noise contrasts. The square root relationship suggests perceptual summation of target and distracters. Manipulating the spread of visual noise suggests that the summation is feature-based, possibly operating on the outputs of first-stage orientation analysers. Pre-cueing the target eliminates the effects of set-size, showing that the summation is under rapid attentional control; the visual system can choose between high performance over a limited area and poorer performance over a much larger area.

Saccadic suppression precedes visual motion analysis.

There is now good evidence that perception of motion is strongly suppressed during saccades (rapid shifts of gaze), presumably to blunt the disturbing sense of motion that saccades would otherwise elicit. Other aspects of vision, such as contrast detection of high-frequency or equiluminant gratings, are virtually unaffected by saccades [1] [2] [3] [4] [5]. This has led to the suggestion that saccades may suppress selectively the magnocellular pathway (which is strongly implicated in motion perception), leaving the parvocellular pathway unaffected [5] [6]. Here, we investigate the neural level at which perception of motion is suppressed. We used a simple technique in which an impression of motion is generated from only two frames, allowing precise control over the stimulus [7] [8]. One frame has a certain fixed contrast, whereas the contrast of the other (the test frame) is varied to determine the threshold for motion discrimination (that is, the lowest test-frame contrast level at which the direction of motion can be correctly guessed). Contrast thresholds of the test depended strongly and non-monotonically on the contrast of the fixed-contrast frame, with a minimum at medium contrast. To study the effect of saccadic suppression, we triggered the two-frame sequence by a voluntary saccade. Thresholds during saccades increased in a way that suggested that saccadic suppression precedes motion analysis: when the test frame was first in the motion sequence there was a general depression of sensitivity, whereas when it was second, the contrast response curve was shifted to a higher contrast range, sometimes even resulting in higher sensitivity than without a saccade. The dependence on presentation order suggests that saccadic suppression occurs at an early stage of visual processing, on the single frames themselves rather than on the combined motion signal. As motion detection itself is thought to occur at an early stage, saccadic suppression must take place at a very early phenomenon.

Cardinal directions for visual optic flow.

As we move through our environment, the flow of deforming images on the retinae provides a rich source of information about the three-dimensional structure of the external world and how to navigate through it. Recent evidence from psychophysical [1] [2] [3] [4], electrophysiological [5] [6] [7] [8] [9] and imaging [10] [11] studies suggests that there are neurons in the primate visual system - in the medial superior temporal cortex - that are specialised to respond to this type of complex 'optic flow' motion. In principle, optic flow could be encoded by a small number of neural mechanisms tuned to 'cardinal directions', including radial and circular motion [12] [13]. There is little support for this idea at present, however, from either physiological [6] [7] or psychophysical [14] research. We have measured the sensitivity of human subjects for detection of motion and for discrimination of motion direction over a wide and densely sampled range of complex motions. Average sensitivity was higher for inward and outward radial movement and for both directions of rotation, consistent with the existence of detectors tuned to these four types of motion. Principle component analysis revealed two clear components, one for radial stimuli (outward and inward) and the other for circular stimuli (clockwise and counter-clock-wise). The results imply that the mechanisms that analyse optic flow in humans tend to be tuned to the cardinal axes of radial and rotational motion.

The effects of ageing on reaction times to motion onset.

We have measured reaction time (RT) to motion onset in two groups of subjects (average ages: 70 and 29 years), for horizontal gratings of 1 c deg-1, modulated in either luminance or colour (equiluminant red-green), for various contrasts and speeds. For both old and young subjects, RTs depended on both speed and contrast, being faster at high speeds and high contrasts, and showed a stronger contrast dependency for chromatic gratings. The older subjects were systematically slower than the younger subjects. The difference between old and young RTs varied with condition, being 30-40 ms more at the slow than at the fast speed. The relative difference in RTs in different stimulus conditions shows that at least some of the increase in response time with age has a sensory origin. The results relate well to previous work on visual evoked potentials.

Large receptive fields for optic flow detection in humans.

We used a psychophysical summation technique to study the properties of detectors tuned to radial, circular and translational motion, and to determine the spatial extent of their receptive fields. Signal-to-noise motion thresholds were measured for patterns curtailed spatially in various ways. Sensitivity for radial, circular and translational motion increased with stimulus area at a rate predicted by an ideal integrator. When sectors of noise were added to the stimulus, sensitivity decreased at a rate consistent with an ideal integrator. Summation was tested for large annular stimuli, and shown to hold up to 70 degrees in some cases, suggesting very large receptive fields for this type of motion (consistent with the physiology of neurones in the dorsal region of the medial superior temporal area (MSTd)). This is a far greater area than observed for summation of contrast sensitivity to gratings (Anderson SJ and Burr DC, Vis Res 1987;29:621-635, and to this type of stimuli (Morrone MC, Burr DC and Vaina LM, Nature 1995;376:507-509, consistent with the suggestion that the two techniques examine different levels of motion analysis.

Seeing biological motion.

One of the more stunning examples of the resourcefulness of human vision is the ability to see 'biological motion', which was first shown with an adaptation of earlier cinematic work: illumination of only the joints of a walking person is enough to convey a vivid, compelling impression of human animation, although the percept collapses to a jumble of meaningless lights when the walker stands still. The information is sufficient to discriminate the sex and other details of the walker, and can be interpreted by young infants. Here we measure the ability of the visual system to integrate this type of motion information over space and time, and compare this capacity with that for viewing simple translational motion. Sensitivity to biological motion increases rapidly with the number of illuminated joints, far more rapidly than for simple motion. Furthermore, this information is summed over extended temporal intervals of up to 3 seconds (eight times longer than for simple motion). The steepness of the summation curves indicates that the mechanisms that analyse biological motion do not integrate linearly over space and time with constant efficiency, as may occur for other forms of complex motion, but instead adapt to the nature of the stimulus.

Reaction time to motion onset of luminance and chromatic gratings is determined by perceived speed.

We measured reaction times for detecting motion onset for sinusoidal gratings whose contrast was modulated in either luminance or chromaticity, for various drift rates and contrasts. In general, reaction times to chromatic gratings were slower than to luminance gratings of matched cone contrast, but the difference in response depended critically on both contrast and speed. At high image speeds there was virtually no difference, whereas at low speeds, the difference was pronounced, especially at low contrasts. At high image speeds there was little dependence of reaction times on contrast (for either luminance or colour), whereas at low speeds the dependence was greater, particularly for chromatic stimuli. This pattern of results is reminiscent of those found for apparent speed of drifting luminance and chromatic gratings. We verified the effects of contrast on perceived speed, and went on to show that the effects of contrast on reaction times are totally predictable by the perceived speed of the stimuli, as if it were perceived rather than physical speed that determined reaction times. Our results support that idea of separate systems for fast and slow motion (with separate channels for luminance and colour at slower speeds), and further suggest that apparent speed and reaction times may be determined at a similar stage of motion analysis.

Capture and transparency in coarse quantized images.

This study examines the effect of coarse quantization (blocking) on image recognition, and explores possible mechanisms. Thresholds for noise corruption showed that coarse quantization reduces drastically the recognizability of both faces and letters, well beyond the levels expected by equivalent blurring. Phase-shifting the spurious high frequencies introduced by the blocking (with an operation designed to leave both overall and local contrast unaffected, and feature localization) greatly improved recognizability of both faces and letters. For large phase shifts, the low spatial frequencies appear in transparency behind a grid structure of checks or lines. We also studied a more simple example of blocking, the checkerboard, that can be considered as a coarse quantized diagonal sinusoidal plaid. When one component of the plaid was contrast-inverted, it was seen in transparency against the checkerboard, while the other remained "captured" within the block structure. If the higher harmonics are then phase-shifted by pi, the contrast-reversed fundamental becomes captured and the other seen in transparency. Intermediate phase shifts of the higher harmonics cause intermediate effects, which we measured by adjusting the relative contrast of the fundamentals until neither orientation dominated. The contrast match varied considerably with the phase of the higher harmonics, over a range of about 1.5 log units. Simulations with the local energy model predicted qualitatively the results of the recognizability of both faces and letters, and quantitatively the apparent orientation of the modified checkerboard pattern. More generally, the model predicts the conditions under which an image will be "captured" by coarse quantization, or seen in transparency.

Apparent position of visual targets during real and simulated saccadic eye movements.

It is now well established that briefly flashed single targets are mislocalized in space, not only during saccades but also before them. We show here by several techniques (including a vernier judgment that did not require absolute location in space) that errors appear up to 100 msec before saccades are made and are maximal just before they start. The size and even the sign of errors depend strongly on position in the visual field, the complete pattern of errors suggesting a compression of visual space around the initial fixation point and the target of the impending saccade. The compression was confirmed by displaying multiple rather than single targets and was found to be powerful enough to reduce or even to remove vernier offset for pairs of bars shown simultaneously and to create offsets for colinear bars separated in time by 75 msec. It also reduced the apparent number of parallel bars. When saccades were simulated by moving the display at saccadic speed, there were sometimes errors of location, but only for tasks requiring absolute judgment of position. The pattern of errors differed greatly from that during saccades and, in particular, showed no signs of compression. We can model our saccade results by assuming a shift in the point in space associated with eye position compression of eccentricity along the axis of saccades.

Compression of visual space before saccades.

Saccadic eye movements, in which the eye moves rapidly between two resting positions, shift the position of our retinal images. If our perception of the world is to remain stable, the visual directions associated with retinal sites, and others they report to, must be updated to compensate for changes in the point of gaze. It has long been suspected that this compensation is achieved by a uniform shift of coordinates driven by an extra-retinal position signal, although some consider this to be unnecessary. Considerable effort has been devoted to a search for such a signal and to measuring its time course and accuracy. Here, by using multiple as well as single targets under normal viewing conditions, we show that changes in apparent visual direction anticipate saccades and are not of the same size, or even in the same direction, for all parts of the visual field. We also show that there is a compression of visual space sufficient to reduce the spacing and even the apparent number of pattern elements. The results are in part consistent with electrophysiological findings of anticipatory shifts in the receptive fields of neurons in parietal cortex and superior colliculi.

Motion deblurring in human vision.

Under normal viewing conditions we are little conscious of blur in moving objects, despite the persistence of vision. Moving objects look more blurred in brief than in long exposures, suggesting an active mechanism for suppressing motion blur. To see whether blur suppression would improve visual discrimination of objects, we measured blur discrimination thresholds for moving Gaussian-blurred edges and bars. The observer's task was to decide which of two moving stimuli, presented successively, was the more blurred. It is known that for stationary objects the just-noticeable difference in blur increases with baseline blur; therefore, if motion increases blur, it would be expected to increase the just-noticeable difference in blur. An active deblurring mechanism, on the other hand, would be expected to counteract the detrimental effects of motion blur on discrimination performance. We found, however, that motion increased thresholds for blur discrimination, both for brief (40 ms) and for longer (150 ms) exposures. We conclude that motion deblurring is a subjective effect, which does not enhance visual discrimination performance. Moving objects appear sharp, not because of some special mechanism that removes blur, but because the visual system is unable to perform the discrimination necessary to decide whether the moving object is really sharp or not.