M. Concetta Morrone

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.

Saccadic eye movements cause compression of time as well as space

There is now considerable evidence that space is compressed when stimuli are flashed shortly before or after the onset of a saccadic eye movement. Here we report that short intervals of time between two successive perisaccadic visual (but not auditory) stimuli are also underestimated, indicating a compression of perceived time. We were even more surprised that in a critical interval before saccades, perceived temporal order is consistently reversed. The very similar time courses of spatial and temporal compression suggest that both are mediated by a common neural mechanism, probably related to the predictive shifts that occur in receptive fields of many visual areas at the time of saccades.

Neural mechanisms for timing visual events are spatially selective in real-world coordinates

It is generally assumed that perceptual events are timed by a centralized supramodal clock. This study challenges this notion in humans by providing clear evidence that visual events of subsecond duration are timed by visual neural mechanisms with spatially circumscribed receptive fields, localized in real-world, rather than retinal, coordinates.

Spatiotopic temporal integration of visual motion across saccadic eye movements

Saccadic eye movements pose many challenges for stable and continuous vision, such as how information from successive fixations is amalgamated into a single precept. Here we show in humans that motion signals are temporally integrated across separate fixations, but only when the motion stimulus falls either on the same retinal region (retinotopic integration) or on different retinal positions that correspond to the same external spatial coordinates (spatiotopic integration). We used individual motion signals that were below detection threshold, implicating spatiotopic trans-saccadic integration in relatively early stages of visual processing such as the middle temporal area (MT) or V5 of visual cortex. The trans-saccadic buildup of important congruent visual information while irrelevant non-congruent information fades could provide a simple and robust strategy to stabilize perception during eye movements.

Recognition of Positive and Negative Bandpass-Filtered Images

A study is reported in which the significance for vision of low- and high-spatial-frequency components of photographic positive and negative images was investigated by measuring recognition of bandpass-filtered photographs of faces. The results show that a 1.5 octave bandpass-filtered image contains sufficient visual information for good recognition performance, provided the filter is centred close to 20 cycles facewidth−1. At low spatial frequencies negatives are more difficult to recognize than positives, but at high spatial frequencies there is no difference in recognition, implying that it is the low-frequency components of negatives which present difficulties for the visual system.

The effects of ageing on the pattern electroretinogram and visual evoked potential in humans

We have recorded patterns electroretinograms (PERGs) and visual evoked potentials (VEPs) from 14 elderly subjects (mean age 72 yr) and 12 young subjects (mean age 21 yr) in response to stimulation by high contrast sinusoidal grating patterns of variable spatial frequency (at 9 Hz) and temporal frequency (at 1.7 c/deg). The major effect of aging on the PERG was an aspecific reduction in amplitude (of about 40%) at most spatial and temporal frequencies, together with a small but systematic phase lag. Control measurements suggest that senile miosis may be responsible for the phase lag, but not for the reduction in amplitude. The effects of aging on the VEP were more dramatic and depended on the spatial and temporal properties of the stimulus. VEP amplitudes (at 1.7 c/deg) were significantly lower for the aged at low temporal frequencies (below about 6 Hz), but were similar at high temporal frequencies. At 9 Hz, there was no effect of spatial frequency on VEP amplitude. At high temporal frequencies (above 10 Hz), the latencies of VEPs (estimated from the rate at which phase varied with temporal frequency) were similar for old and young (94 and 99 msec respectively). Below 10 Hz, however, the latencies of the old observers was much greater (153 compared with 108 msec). The second-harmonic phase of VEPs of the old but not the young decreased considerably with spatial frequency, by about 1.9 pi radians (52 msec) over the range from 0.5 to 11 c/deg. The selective reduction in amplitude at low temporal frequencies, the longer latencies at low temporal frequencies and the phase lag at high spatial frequencies are consistent with the hypothesis that mechanisms sensitive to high spatial and low temporal frequencies are selectively degraded by aging.

Evidence for edge and bar detectors in human vision

The structure of receptive fields of human visual detectors was investigated by studying their phase response. Observers were required to discriminate between pairs of periodic stimuli that differed in phase by 180 degrees (reversed in contrast). The stimuli comprised 256 harmonics, smoothly filtered in amplitude, and congruent in phase at the origin. Reversal discrimination thresholds were measured as a function of the phase of the harmonics. Thresholds were slightly higher for phases around 45 degrees, consistent with the idea that all discriminations were mediated by independent detectors with 0 or 90 degrees phase response (assuming probability summation between them). Discrimination thresholds were also measured with a pedestal stimulus, of phase complementary to that of the test gratings. For discriminations between 0 and 180 degrees (cosine phase), or 90 and 270 degrees (sine phase), the complementary pedestal had little effect, implying independence of detectors in sine and cosine phase. However, for discrimination between 45 and 225 degrees (stimuli containing both sine and cosine components) the complementary pedestal, which also contained both sine and cosine components, facilitated greatly discrimination thresholds. The results suggest that there exist two classes of detectors, one with a Fourier phase spectrum of 0, the other with a Fourier phase spectrum of 90 degrees. This implies that the receptive fields are symmetric, one class having even-symmetry (line-detectors), the other odd-symmetry (edge-detectors).

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.

Visual ageing: unspecific decline of the responses to luminance and colour.

We have investigated whether ageing affects selectively the responses to equiluminant patterns of pure colour contrast. In two groups of subjects (mean ages 29 and 72 yr) contrast thresholds were measured psychophysically for the detection and for the discrimination of the direction of motion of drifting gratings. The gratings were modulated either in pure luminance contrast (and uniform colour), or pure chromatic contrast (red-green equiluminant gratings). In subjects of the same age groups, visual evoked potentials (VEP) were recorded in response to gratings with either pure luminance contrast or pure colour contrast sinusoidally reversed in contrast at various temporal frequencies. It was shown that psychophysical contrast sensitivity for equiluminant patterns deteriorates significantly with age, and VEP latency increases. However, these effects of ageing on the responses to patterns of pure colour contrast are substantially the same as those observed in the same subjects for stimuli with pure luminance contrast. The results suggest that ageing causes a small and unspecific decline of the response of the visual system to luminance and colour contrast.

Automatic gain control contrast mechanisms are modulated by attention in humans: evidence from visual evoked potentials

This study investigated the effect of attention on the contrast response curves of steady-state visual evoked potentials (VEPs) to counter-phased sinusoidal gratings. The 1 cyc/deg gratings were modulated either in luminance or chromaticity (equiluminant red-green). The luminance grating counter-phased at 9 Hz (to favour activation of the magno-cellular system), and the chromatic grating at 2.5 Hz (to favour activation of the parvo-cellular system). Attention was directed towards the gratings (displayed in the left visual field) by requiring subjects to detect and respond to randomly occurring changes in contrast. In the control condition, attention towards the grating was minimised by requiring subjects to detect a target letter amongst distracters briefly flashed in the contra-lateral visual field. Attention increased VEP amplitudes for both luminance and chromatic stimuli, more so at high than at low contrasts, increasing the slope of the contrast amplitude curves (over the non-saturating range of contrasts). The estimates of contrast threshold from extrapolation of amplitudes were unaffected by attention. Attention also changed the VEP phases, but only for luminance gratings, where it acted to reduce the magnitude of phase advance with contrast. Attention had no effect on the average phases for chromatic gratings. The results are consistent with the notion that attention acts on cortical gain control mechanisms, which are known to be different for the magno- and parvo-cellular systems.