Motion coherence thresholds in infants--different tasks identify at least two distinct motion systems.

Optokinetic nystagmus (OKN) can be demonstrated from birth, but behavioural discrimination tasks such as habituation and preferential looking do not reveal any sensitivity to motion direction until a few weeks of age. This study compared coherence threshold for motion direction for OKN and preferential looking responses using closely comparable stimuli, in infants between 6 and 27 weeks of age. Infants were tested with two random dot motion displays, a uniform area of moving dots for OKN responses and a display in which a region was segmented on one side by differential motion direction for preferential looking responses. Coherence thresholds for each response were determined by a staircase method. For OKN responses, mean coherence thresholds were between 20% and 25%, with no significant improvement in OKN performance throughout the age range. Preferential looking thresholds were significantly higher than OKN thresholds. Preferential looking thresholds improved significantly with age, but remained higher than OKN thresholds throughout the age range tested. Experiments varying direction reversal frequency and stimulus area indicated that these differences were not simply a consequence of the spatial and temporal non-uniformity of the preferential looking stimulus. The differences in sensitivity levels and age trends for OKN and preferential looking responses we have found suggest that different directional mechanisms are involved in the two responses. We discuss the possibility that, in early infancy, OKN and preferential looking reflect the performance of subcortical and cortical directional mechanisms respectively.

Visual function and EEG reactivity in infants with perinatal brain lesions at 1 year.

The aim of this study was to examine the correlation between EEG, visual, and brain MRI findings in 19 term infants with perinatal brain lesions. All 19 had their visual acuity and visual fields assessed and had an EEG and a brain MRI performed at 1 year of age. Four of the five infants with normal optic radiations and occipital cortices on MRI had normal vision. Involvement of optic radiations and occipital cortices was only associated with visual abnormalities in eight of 14 infants. The correlation between visual abnormalities and EEG findings was stronger. All infants with a completely normal EEG from the posterior regions had normal vision and all those with an EEG non-reactive to eye closure had visual abnormalities, irrespective of MRI data. A reactive EEG with other abnormal features (such as spikes, rapid or slow activities) was accompanied by abnormal vision in five of eight participants. Results suggest that there is a better correlation between visual function and EEG activity than between visual function and involvement of the classical visual areas of the occipital cortex and optic radiations on brain MRI at 1 year of age.

Brain areas sensitive to coherent visual motion.

Detection of coherent motion versus noise is widely used as a measure of global visual-motion processing. To localise the human brain mechanisms involved in this performance, functional magnetic resonance imaging (fMRI) was used to compare brain activation during viewing of coherently moving random dots with that during viewing spatially and temporally comparable dynamic noise. Rates of reversal of coherent motion and coherent-motion velocities (5 versus 20 deg s-1) were also compared. Differences in local activation between conditions were analysed by statistical parametric mapping. Greater activation by coherent motion compared to noise was found in V5 and putative V3A, but not in V1. In addition there were foci of activation on the occipital ventral surface, the intraparietal sulcus, and superior temporal sulcus. Thus, coherent-motion information has distinctive effects in a number of extrastriate visual brain areas. The rate of motion reversal showed only weak effects in motion-sensitive areas. V1 was better activated by noise than by coherent motion, possibly reflecting activation of neurons with a wider range of motion selectivities. This activation was at a more anterior location in the comparison of noise with the faster velocity, suggesting that 20 deg s-1 is beyond the velocity range of the V1 representation of central visual field. These results support the use of motion-coherence tests for extrastriate as opposed to V1 function. However, sensitivity to motion coherence is not confined to V5, and may extend beyond the classically defined dorsal stream.

Directional motion asymmetry in infant VEPs--which direction?

Monocular viewing during early infancy reveals asymmetries in optokinetic nystagmus (OKN) and visual evoked potentials (VEPs). This study investigates the VEP asymmetry to see if it is consistent in direction with the OKN asymmetry. Steady-state VEPs were recorded from infants (5-21 weeks) viewing gratings that underwent successive displacements in the same direction, leftward or rightward. In addition, transient VEPs were recorded to the two directions of an oscillating stimulus. Both tests produced larger VEP amplitudes for nasal-to-temporal compared to temporal-to-nasal movement. Horizontal eye movements were monitored by EOG while viewing these stimuli to test whether the asymmetry was a consequence of eye movements. No difference in eye movements as a function of the stimulus was found, excluding differences in retinal slip as an explanation of the asymmetry. The stronger neural response for nasal-to-temporal displacements is opposite to the asymmetry of OKN. Oculomotor and VEP asymmetries may be related; however this relationship is not simply that the stronger neural response, indicated by the VEP, leads to a stronger optokinetic response.

Speed and direction of locally-paired dot patterns.

Phenomenal transparency in random-dot kinematograms is abolished when two motion directions are 'locally-balanced' by pairing limited-lifetime dots at each location [Qian, Andersen and Adelson (1994). Journal of Neuroscience, 14, 7357-7366]. Qian et al. also report that locally-paired stimuli appear as directionless flicker when the paired dots differ in their directions by 90 degrees or more. They attribute this to local inhibition between motion detectors more than 45 degrees apart. We investigated perceived motion in such displays, by requiring subjects to make direction and speed judgements with locally-paired stimuli containing two directions 60, 90 or 120 degrees apart. Subjects perceived coherent motion in these displays and made reliable direction judgements, indicating that the two motions are combined rather than interfering destructively. Our results show that the judged motion of locally-paired stimuli is in the vector-average direction of the two components. This vector-averaging rule also applies when the two sets of component dots differ in their velocity. Similarly, speed judgements comply with a vector-averaging rule for a range of speeds as well as for mixed-speed stimuli. These results suggest that the abolition of transparency does not necessarily imply abolition of a global motion percept. The local interaction abolishing transparency is not exclusively inhibitory, at least for directions up to 120 degrees apart, but generates a vector combination of the superimposed motions.

Form and motion coherence activate independent, but not dorsal/ventral segregated, networks in the human brain.

There is much evidence in primates' visual processing for distinct mechanisms involved in object recognition and encoding object position and motion, which have been identified with 'ventral' and 'dorsal' streams, respectively, of the extra-striate visual areas [1] [2] [3]. This distinction may yield insights into normal human perception, its development and pathology. Motion coherence sensitivity has been taken as a test of global processing in the dorsal stream [4] [5]. We have proposed an analogous 'form coherence' measure of global processing in the ventral stream [6]. In a functional magnetic resonance imaging (fMRI) experiment, we found that the cortical regions activated by form coherence did not overlap with those activated by motion coherence in the same individuals. Areas differentially activated by form coherence included regions in the middle occipital gyrus, the ventral occipital surface, the intraparietal sulcus, and the temporal lobe. Motion coherence activated areas consistent with those previously identified as V5 and V3a, the ventral occipital surface, the intraparietal sulcus, and temporal structures. Neither form nor motion coherence activated area V1 differentially. Form and motion foci in occipital, parietal, and temporal areas were nearby but showed almost no overlap. These results support the idea that form and motion coherence test distinct functional brain systems, but that these do not necessarily correspond to a gross anatomical separation of dorsal and ventral processing streams.

Developmental changes in optokinetic mechanisms in the absence of unilateral cortical control.

Animal models suggest that the asymmetry of monocular optokinetic nystagmus (OKN) in young infants can be explained by a direct pathway from retina to the midbrain nucleus of the optic tract. However, earlier studies with hemispherectomized infants showed no evidence for OKN responses towards the damaged cortex that could be ascribed to this subcortical pathway. In longitudinal testing of two infants with very extensive unilateral cortical damage, we have now shown that OKN responses in both directions do occur before 10 months of age. OKN towards the damaged cortex, indicating functioning of the direct pathways in the absence of cortical control, drops out in the later development. The neural circuitry responsible for OKN in humans appears to undergo a plastic reorganization.

How does noise influence the estimation of speed?

Local motion signals have to be combined in space and time, to yield a coherent motion percept as it is involved in a variety of visual tasks. This combination necessarily means to trade-off between loosing spatio-temporal resolution by pooling local signals and maintaining perceptually significant segmentation between them. When signals are pooled to detect the presence of coherent motion in large amounts of random noise, the question raised is how the noise affects the perceived quality, in particular speed, of the coherent motion. Is there an analogy to the well-known reduction in the perceived speed of moving gratings at low contrast? Using a two-interval forced-choice procedure, we have investigated the assessment of speed in random-dot kinematograms containing different proportions of noise. Under the conditions investigated, there is no strong reduction of perceived speed with increasing noise, as long as coherence levels remain well above the thresholds for directional judgements. This basic result, which could suggest considerable but not perfect segregation of signal and noise motion components in the pooling process leading to speed estimation, is discussed in relation to a model that is designed to decode speed from a population of elementary motion detectors (EMDs) of the correlation type. A strategy to estimate speed from a set of EMDs with a variety of spatio-temporal tuning does not only provide a velocity predictor unambiguous with the spatial structure of the stimulus, but also is largely independent of noise.

What motion distributions yield global transparency and spatial segmentation?

We have examined the ability of observers to parse bimodal local-motion distributions into two global motion surfaces, either overlapping (yielding transparent motion) or spatially segregated (yielding a motion boundary). The stimuli were random dot kinematograms in which the direction of motion of each dot was drawn from one of two rectangular probability distributions. A wide range of direction distribution widths and separations was tested. The ability to discriminate the direction of motion of one of the two motion surfaces from the direction of a comparison stimulus was used as an objective test of the perception of two discrete surfaces. Performance for both transparent and spatially segregated motion was remarkably good, being only slightly inferior to that achieved with a single global motion surface. Performance was consistently better for segregated motion than for transparency. Whereas transparent motion was only perceived with direction distributions which were separated by a significant gap, segregated motion could be seen with abutting or even partially overlapping direction distributions. For transparency, the critical gap increased with the range of directions in the distribution. This result does not support models in which transparency depends on detection of a minimum size of gap defining a bimodal direction distribution. We suggest, instead, that the operations which detect bimodality are scaled (in the direction domain) with the overall range of distribution. This yields a flexible, adaptive system that determines whether a gap in the direction distribution serves as a segmentation cue or is smoothed as part of a unitary computation of global motion.

Development of illusory-contour perception in infants.

We investigated whether infants from 8-22 weeks of age were sensitive to the illusory contour created by aligned line terminators. Previous reports of illusory-contour detection in infants under 4 months old could be due to infants' preference for the presence of terminators rather than their configuration. We generated preferential-looking stimuli containing sinusoidal lines whose oscillating, abutting terminators give a strong illusory contour in adult perception. Our experiments demonstrated a preference in infants 8 weeks old and above for an oscillating illusory contour compared with a stimulus containing equal terminator density and movement. Control experiments excluded local line density, or attention to alignment in general, as the basis for this result. In the youngest age group (8-10 weeks) stimulus velocity appears to be critical in determining the visibility of illusory contours, which is consistent with other data on motion processing at this age. We conclude that, by 2 months of age, the infant's visual system contains the nonlinear mechanisms necessary to extract an illusory contour from aligned terminators.

Infant emmetropization: longitudinal changes in refraction components from nine to twenty months of age.

Rapid emmetropization is described in pediatrically normal infants from 9 months of age during the following year. The infants, obtained from various categories of the Cambridge population screening program, provided a broad range of refractive errors. The large group of 254 nonanisometropic infants studied allowed the mean rate of change and dependence on the initial refraction value to be determined. Refraction was measured by cycloplegic retinoscopy. Rapid emmetropization changes occurred in the following refractive components: mean spherical equivalent (MSE), astigmatism magnitude, the horizontal astigmatism component, the infant's most positive meridian, and the infant's most negative meridian. The MSE and astigmatism rates of change (diopters/year), were highly dependent on their respective initial powers (r = -0.61 and r = -0.76). The percentage weighted mean proportional rate of change for MSE was -30% (SE 4%) and for astigmatism magnitude it was -59% (SE 14%). There was much individual variation, with some exhibiting fast emmetropization and others not. The MSE and astigmatism changes, however, were almost independent of each other. The refractive errors of the most positive and most negative meridians emmetropize because they are both derived from the MSE and half the astigmatism. With-the-rule astigmatism was more prevalent than against-the-rule astigmatism at 9 months of age, and with-the-rule astigmatism exhibited a significantly greater proportional rate of change. The relationship of emmetropization and refractive screening is considered. A new component "MOMS" is introduced, the maximum ocular meridional separation, when both eyes are considered. Thus incorporating astigmatism and anisometropia may be a good single indicator of conditions associated with later amblyopia. The almost independent emmetropization of the MSE and astigmatism components is an important result to consider in theories of emmetropization, refractive screening, clinical prescribing, and the evaluation of infants in treatment trials.

Integration across directions in dynamic random dot displays: vector summation or winner take all?

Recent studies have clearly demonstrated that the activity of directionally selective neuronal populations in the middle temporal (MT) and medial superior temporal (MST) cortical areas plays a direct role in the judgment of the direction of visual motion. However, the way in which the information is derived from a population of neurons remains unknown. Two principal models have been suggested in the past: the vector summation model suggests that the responses of neurons encoding all directions of motion are weighted and pooled to obtained an accurate estimate of the mean direction of motion; the winner-take-all model is based on a competition between different direction-specific channels, so that decisions are cast in favor of the channel generating the strongest directional signal. To discriminate between these two models we generated random dot stimuli that contained an asymmetric distribution of directions of motion. Human subjects were asked to adjust the global direction of motion to the upward vertical direction. When the directional signals were of similar strength, subjects tended to perceive global motion in the mean direction of motion (corresponding to vector summation), but as one directional signal became more prominent, most subjects' settings diverged from the mean towards the modal direction of motion. Some subjects could either match the mean or the modal direction of motion in the display, depending on the task instructions. These results suggest that the perceptual judgment of direction of motion is not based on any rigid algorithm generating a single valued output. Rather, human observers are able to judge different aspects of the distribution of activity in a cortical area depending on the task requirements.

What is noise for the motion system?

Motion coherence thresholds in random-dot patterns have been widely adopted as a measure of performance in visual motion processing. However, there has been diversity in the type of "noise" in which a coherent motion signal has to be detected. Here we compare coherence thresholds for three ways of creating motion noise: dots replotted in random positions in each new frame; dots with a set displacement but following a random walk from frame to frame; or dots moving in random directions which remain constant for a given dot over a sequence of displacements. In each case, the signal dots may either remain the same throughout the display sequence, or the signal dots may be re-selected afresh on each frame ("different"). With our display (3 deg square, 120 msec exposure, velocity = 5 or 10 deg sec-1), all these different noise conditions yielded similar thresholds around 5-8%. There were some small but systematic differences between conditions. Thresholds in random-direction displays were consistently higher than those in random-walk or random-position displays, especially at the lower velocity. However, this effect is much smaller than would be expected from the increased standard error of the noise mean in random direction, perhaps because the motion system integrates information most effectively over a local region of space and/or time. Subjects" performance could not be explained by a strategy of identifying individual signal dots with extended trajectories. The similarity between random-walk and random-position thresholds implies that subjects do not exploit the marked differences in speed distribution between signal and noise dots in the latter case. The practical message for the design and interpretation of experiments using coherence thresholds is that the results are not much affected by the choice of noise, at least within the range of stimuli tested here.

The temporal integration and resolution of velocity signals.

The temporal properties of human visual motion detection were explored. Experiment 1 measured thresholds for speed discrimination as a function of stimulus duration. Thresholds fell asymptotically to a Weber fraction around 0.06 over a period of approx. 100 msec, with faster speeds asymptoting at slightly shorter stimulus durations. A second experiment required subjects to discriminate a pattern that was modulated between two speeds from one which remained at a constant speed. The minimum depth of the modulation required to make this judgement was found to be equivalent to a Weber fraction of 0.3 at low modulation rates, around five times greater than when the velocities were presented in isolation (expt 1). At some higher modulation rate performance dramatically declined. The modulation rate at which this occurred decreased with stimulus speed, and increased with stimulus size. The results of expt 1 seem consistent with the known properties of primary motion sensors, while the results of the latter experiments may arise from a later stage integrating the output of these primary motion sensors.

Discrimination of spatial phase shows a qualitative difference between foveal and peripheral processing.

Detection and discrimination of compound grating stimuli were examined in foveal and peripheral vision. At the fovea, stimuli containing two components (spatial frequencies F and 3F) can be discriminated on the basis of their relative spatial phase when the 3F component is at a contrast below its independent detection threshold. This is no longer the case at increasing retinal eccentricity, where phase discrimination thresholds fall off much more steeply than simple detection thresholds. This relative fall-off in discrimination performance is still present for stimuli scaled for the cortical magnification factor, and is not attributable to fading of peripheral images due to the Troxler effect. The results therefore must imply a qualitative change in the processing of phase information between foveal and peripheral vision.

Pre-attentive detection of a target defined by stereoscopic slant.

Does the visual system represent stereoscopic depth purely as a map of local disparities, or does it explicitly represent local relationships of disparity, such as disparity gradients? Experiments are reported in which visual search for a target containing the same disparity range as other elements in the display, but differing in the relationship of the disparities (stereo slant), was used to determine whether the target showed 'pop-out' like a unitary feature, or the serial search characteristic of feature conjunctions. Each stereo pair of elements was selected randomly from a range of outline parallelograms leaning to the right or to the left, so that the target could not be identified using any monocular shape cue. Response times for detection of the target (present on 50% of the trials) were independent of the number of elements in the display. This result was confirmed by varying element size and spacing, and by using oblique crosses rather than parallelograms as stimuli. It is concluded that stereoscopically defined slant, or disparity gradient, can be processed and compared in parallel across the display, and acts in this respect as an explicit unitary visual property. This contrasts with findings in analogous experiments on movement, which show that targets defined by divergence or deformation of optic flow can only be identified by serial search.

Serial search for targets defined by divergence or deformation of optic flow.

The optic flow field can be described in terms of the local differential measures, divergence, deformation, and rotation, which are informative about observer motion and the 3-D structure of the environment. Does an explicit representation of these measures exist in human visual processing in the form of a feature map? Triesman's criteria were used to investigate this; ie is there 'pop-out' for a target defined as different in local divergence or deformation from surrounding elements, or is a serial search necessary? The stimulus arrays contained 3, 5, or 9 square or rectangular elements, which each underwent repeated cycles of expansion, contraction, or deformation. The time required to detect a target undergoing the opposite transformation increased steeply with the number of elements, implying very slow serial search. (The mean time was 210 ms per element for divergence targets and 542 ms per element for deformation). The process was clearly still serial when the density and number of elements was increased up to 48 in an array 2.16 deg x 2.16 deg. In contrast, a single line element undergoing the opposite direction of translation motion to the rest of the display did show pop-out. It is concluded that no parallel processes seem to exist which are sensitive to the spatial uniformity of divergence and of deformation of optic flow. These differential properties may be derived as conjunctions of signals from a primary process which extracts local velocity. This result contrasts with our findings for targets defined by stereo disparity gradient, which show parallel processing in analogous experiments.

Direction discrimination for band-pass filtered random dot kinematograms.

When an array of random dots is displaced, the ability to report the direction of apparent motion is subject to an upper spatial limit (dmax). As the size of the displacement is increased, direction discrimination errors show a monotonic increase that becomes asymptotic at a chance level. We have measured direction discrimination using spatially band-pass filtered random dots. These stimuli do not yield a monotonic increase in errors. Rather, for displacements greater than around 1 cycle of the stimulus centre frequency (Fc), performance oscillates about chance, with displacements of 1 1/4 cycles of Fc yielding systematic errors in perceived direction. We analyse this pattern of performance in terms of the stimulus autocorrelation function and conclude that dmax can be taken as lying on the initial rising portion of the displacement versus error function. Using this definition we find, in line with the results of Chang and Julesz (1985), that dmax scales inversely with Fc. Contrary to the results of Chang and Julesz, we find that this scaling holds beyond 4 c/deg.

Masking of low frequency information in short-range apparent motion.

When an array of random dots is displaced, the ability to report the direction of apparent motion is subject to an upper spatial limit (dmax). Using spatially low-pass filtered random dot kinematograms we show that dmax is dependent on the upper cut-off frequency of the stimulus (Fh). The extent of this dependence is critically dependent on the size of the stimulus. Our results suggest a process whereby low spatial frequency motion information is masked by the presence of high spatial frequencies in the same region of the field, analogous to phenomena occurring in the perception of static form (e.g. the Abraham Lincoln effect). The effects of stimulus size on dmax, found for broad-band stimuli by ourselves and others, result from a loss of high frequency sensitivity at increased retinal eccentricities; this loss reduces the masking effect of high frequencies, as stimulus size increases.

Differences in the processing of short-range apparent motion at small and large displacements.

Using random dot patterns we have compared performance on direction discrimination tasks for single and multi-step sequences of apparent motion at a range of displacement sizes. Performance was measured by varying the correlation between the frames. For "small" displacements we found that no improvement in performance occurs with stimulus duration (number of frames) if the movement of individual elements within the pattern was restricted to one step, whereas if elements undergo multiple steps, performance improves with duration. For "large" displacements, on the contrary, performance improves with increasing stimulus duration irrespective of whether individual elements are restricted to single steps. These results suggest that small and large displacements are processed in different ways. We review possible psychophysical and physiological correlates of this suggestion.