Visual neuroscience: Resonating to natural images.

Visual neurons may be optimized to produce sparse, distributed responses to natural scenes. This proposal, along with recent results from monkey fMRI and electrophysiology, may force us to re-interpret many neuroimaging results.

The effects of aging on visual memory: evidence for functional reorganization of cortical networks.

Recent evidence suggests that the mature human brain is capable of substantial functional reorganization following injury. The fact that the brain retains a great deal of plasticity raises the possibility that cortical reorganization may occur during normal aging. We examined this issue by using positron emission tomography (PET) to measure the brain activity associated with short-term memory for simple visual attributes in young and old observers. A two-interval forced choice procedure was used to measure spatial frequency discrimination thresholds for sine wave gratings presented at different inter-stimulus intervals (ISI). Memory load was manipulated by varying the duration of the ISI and by presenting an irrelevant masking stimulus in the middle of the ISI. Old and young observers performed the experiment equally well. However, the neural systems correlated with good performance differed for the two age groups. The results support the hypothesis that the functional networks that underlie visual memory undergo reorganization during aging.

Time course of amodal completion revealed by a shape discrimination task.

We measured the extent of amodal completion as a function of stimulus duration over the range of 15-210 msec, for both moving and stationary stimuli. Completion was assessed using a performance-based measure; a shape discrimination task that is easy if the stimulus is amodally completed and difficult if it is not. Specifically, participants judged whether an upright rectangle was longer horizontally or vertically, when the rectangle was unoccluded, occluded at its corners by four negative-contrast squares, or occluded at its corners by four zero-contrast squares. In the zero-contrast condition, amodal completion did not occur because there were no occlusion cues; in the unoccluded condition, the entire figure was present. Thus, comparing performance in the negative-contrast condition to these two extremes provided a quantitative measure of amodal completion. This measure revealed a rapid but measurable time course for amodal completion. Moving and stationary stimuli took the same amount of time to be completed (approximately 75 msec), but moving stimuli had slightly stronger completion at long durations.

The effects of occlusion and past experience on the allocation of object-based attention.

There is considerable evidence indicating that cuing a specific portion of an object results in the entire object's being attended to. In the present study, we examined whether previous experience with an object could halt perceptual (i.e., amodal) completion. In Experiment 1, two parallel rectangles were initially displayed, and then the middle portions of these objects were occluded. Attentional cuing effects were found for both discrete portions of the completed rectangles. In the final two experiments, four discrete objects were initially displayed, followed by the same occluder as that used in the first experiment. The appearance of the occluder (500 msec before the cue in Experiment 2,100 msec before the cue in Experiment 3) allowed the four discrete objects to be completed into two rectangles. Attentional cuing effects were found for the completed rectangles in both experiments, indicating that previous experience was not sufficient to halt the amodal completion of objects.

Generalized common fate: grouping by common luminance changes.

A critical step toward object recognition is the segmentation of a scene into relevant regions. One of the most important cues for segmentation is that of common fate: Elements that move together are grouped together Here we describe a new instantiation of common fate, in which elements move together not through physical space, but through luminance space. Experiment I shows that when elements of a scene become brighter together or darker together, observers group those elements together Experiments 2 and 3 show that this effect is not due to the availability of fixed luminance differences between target and background regions, but requires common changes within each region in the direction of luminance over time. The effect is differentiated from the recently discovered grouping cue of temporal synchrony, and is considered instead to be an extension of Wertheimer's original grouping factor of common fate. Common fate for luminance, or generalized common fate, is an extremely strong cue for the segmentation of a scene, yielding a tremendous advantage over grouping by fixed luminance cues.

Corticolimbic interactions associated with performance on a short-term memory task are modified by age.

Aging has been associated with a decline in memory abilities dependent on hippocampal processing. We investigated whether the functional interactions between the hippocampus and related cortical areas were modified by age. Young and old subjects' brain activity was measured using positron emission tomography (PET) while they performed a short-term memory task (delayed visual discrimination) in which they determined which of two successively presented sine-wave gratings had the highest spatial frequency. Behavioral performance was equal for the two groups. Partial least squares (PLS) analysis of PET images identified a hippocampal voxel whose activity was similarly correlated with performance across groups. Using this voxel as a seed, a second PLS analysis identified cortical regions functionally connected to the hippocampus. Quantification of the neural interactions with structural equation modeling suggested that a different hippocampal network supported performance in the elderly. Unlike the neural network engaged by the young, which included prefrontal cortex Brodmann's area (BA) 10, fusiform gyrus, and posterior cingulate gyrus, the network recruited by the old included more anterior areas, i.e., dorsolateral prefrontal cortex (BA 9/46), middle cingulate gyrus, and caudate nucleus. Recruitment of a distinct corticolimbic network for visual memory in the elderly suggests that age-related neurobiological deterioration not only results in focal changes but also in the modification of large-scale network operations.

Deriving behavioural receptive fields for visually completed contours.

The visual system is constantly faced with the problem of identifying partially occluded objects from incomplete images cast on the retinae. Phenomenologically, the visual system seems to fill in missing information by interpolating illusory and occluded contours at points of occlusion, so that we perceive complete objects. Previous behavioural [1] [2] [3] [4] [5] [6] [7] and physiological [8] [9] [10] [11] [12] studies suggest that the visual system treats illusory and occluded contours like luminance-defined contours in many respects. None of these studies has, however, directly shown that illusory and occluded contours are actually used to perform perceptual tasks. Here, we use a response-classification technique [13] [14] [15] [16] [17] [18] [19] [20] to answer this question directly. This technique provides pictorial representations - 'classification images' - that show which parts of a stimulus observers use to make perceptual decisions, effectively deriving behavioural receptive fields. Here we show that illusory and occluded contours appear in observers' classification images, providing the first direct evidence that observers use perceptually interpolated contours to recognize objects. These results offer a compelling demonstration of how visual processing acts on completed representations, and illustrate a powerful new technique for constraining models of visual completion.

Motion and shape in common fate.

We determined how much motion coherence was needed to detect a target group of four moving dots in a dynamic visual noise (DVN) background. The lifetimes of the trajectories of the target and that of the noise dots were the same. In addition to parallel trajectories and collinear dot arrangements, divergent, convergent, or crossing trajectories and non-collinear dot arrangements were also tested. Performance saturated at a lifetime of approximately 600 ms. It was best for parallel trajectories and collinear dots, and worse for crossed trajectories with non-collinear dots, where it approached performance in a no-motion, form-only control experiment. Results illustrate the importance of common fate in motion perception in DVN, when other factors are equated.

Effects of aging on the useful field of view.

Previous research has shown that the useful field of view (UFOV) is a useful tool in predicting driving ability, and the UFOV also seems to decline with age. The goals of the current study were first, to examine UFOV changes systematically as a function of age (15-84), and, second, to determine the effect of dividing attention on the UFOV. Our results show that the deterioration in the UFOV begins early in life (by 20 years, or younger). This deterioration is best conceptualized as a decrease in the efficiency with which observers can extract information from a cluttered scene, rather than by shrinking of the field of view per se. The diminished efficiency among elderly observers is exacerbated when conditions require the division of attention between central and peripheral tasks.

Interactions between symmetry and elongation in determining reference frames for object perception.

Many theories of object recognition posit that objects are encoded with respect to a perceptual frame of reference. Such theories assume that factors such as symmetry and elongation are critical for the assignment of an object's primary axis, and consequently for the extraction of an object's reference frame. The present experiments directly examined the relative roles played by symmetry and elongation in the determination of an object's primary axis, and the extent to which symmetry and elongation interact with one another. We found that observers use both symmetry and elongation in extracting an object's primary axis, that the extent to which each cue dominates depends on its relative salience, and that symmetry and elongation are processed interactively, rather than in encapsulated modules.

Recruitment of unique neural systems to support visual memory in normal aging.

The performance of many cognitive tasks changes in normal aging [1] [2] [3]. Recent behavioral work has identified some tasks that seem to be performed in an age-invariant manner [4]. To understand the brain mechanisms responsible for this, we combined psychophysical measurements of visual short-term memory with positron emission tomography (PET) in young and old individuals. Participants judged the differences between two visual stimuli, and the memory load was manipulated by interposing a delay between the two stimuli. Both age groups performed the task equally well, but the neural systems supporting performance differed between young and old individuals. Although there was some overlap in the brain regions supporting performance (for example, occipital, temporal and inferior prefrontal cortices, and caudate), the functional interconnections between these common regions were much weaker in old participants. This suggests that the regions were not operating effectively as a network in old individuals. Old participants recruited unique areas, however, including medial temporal and dorsolateral prefrontal cortices. These unique areas were strongly interactive and their activity was related to performance only in old participants. Therefore, these areas may have acted to compensate for reduced interactions between the other brain areas.

Effects of aging on calculation efficiency and equivalent noise.

Contrast sensitivity under photopic conditions declines with age; however, the cause of this decline remains unknown. To address this issue, we measured detection thresholds for sine wave gratings in noise, under various conditions of spatial-frequency uncertainty, and estimated observers' internal noise and calculation efficiency. Statistical analyses revealed that efficiencies were lower for old (median age at 68 years) than for young (median age at 22 years) observers; no significant differences in internal noise were found. A control experiment ruled out the possibility that reduced retinal illuminance causes the decline in efficiency with age. Our results demonstrate that age-related neural changes play a major role in the decline in contrast sensitivity with age. Possible contributing mechanisms are considered.

Collisions between moving visual targets: what controls alternative ways of seeing an ambiguous display?

When identical visual targets move directly toward and then past one another, they appear either to stream past one another or to bounce off each other. Bertenthal et al (1993 Perception 22 193-207) accounted for the relative strengths of these two percepts by invoking a directional bias, arising from cooperative interactions within a network of motion detectors. We tested this explanation by devising conditions that would enhance or diminish the strength of such a directional bias. In separate experiments we varied (i) the presence or absence of temporal transients (pausing, disappearance, occlusion); (ii) the distances travelled by the targets; and (iii) their acceleration or deceleration before and after collision. The tendency to see the objects stream past one another was not related to the strength of an hypothesized directional bias, suggesting that the perception of this ambiguous motion display was not mediated by directional recruitment. Instead, the results suggest that perceived direction reflects the operation of neural constraints that mirror the constraints operating upon moving objects in the three-dimensional natural world.

Identification of band-pass filtered letters and faces by human and ideal observers.

To better understand how the visual system makes use of information across spatial scales when identifying different kinds of complex patterns, we measured human and ideal contrast identification thresholds to estimate identification efficiency for 1- and 2-octave wide band-pass filtered letters and faces embedded in 2-D dynamic Gaussian noise. Varying stimulus center frequency from 1 to 70 c/object had different effects on letter and face identification efficiency. In the 2-octave conditions, identification efficiencies decreased by 0.25-0.5 log units for letters and 0.5-1.2 log units for faces as center frequency increased from 6.2 to 49.5 c/object, but only letters were identifiable at center frequencies below 6.2 c/object. In the 1-octave conditions, letter identification efficiencies increased by about 0.5 log units as center frequency increased from 1.1 to 2.2 c/object, and were nearly constant from 2.2 to 35 c/object. Letters were unidentifiable by human observers at 70 c/object. Surprisingly, face identification was impossible for human observers at all center frequencies except 8.8 c/object for one observer, and 8.8 and 17.5 c/object for a second observer. Ideal observer thresholds were obtained for both letters and faces in all conditions, so information was always available to perform the task. Thus, the failure to identify faces reflects constraints on visual processing rather than a lack of stimulus information. Selective spatial sampling may account for some of the differences between letter and face identification efficiencies.

Signal but not noise changes with perceptual learning.

Perceptual discrimination improves with practice. This 'perceptual learning' is often specific to the stimuli presented during training, indicating that practice may alter the response characteristics of cortical sensory neurons. Although much is known about how learning modifies cortical circuits, it remains unclear how these changes relate to behaviour. Different theories assume that practice improves discrimination by enhancing the signal, diminishing internal noise or both. Here, to distinguish among these alternatives, we fashioned sets of faces and textures whose signal strength could be varied, and we trained observers to identify these patterns embedded in noise. Performance increased by up to 400% across several sessions over several days. Comparisons of human performance to that of an ideal discriminator showed that learning increased the efficiency with which observers encoded task-relevant information. Observer response consistency, measured by a double-pass technique in which identical stimuli are shown twice in each experimental session, did not change during training, showing that learning had no effect on internal noise. These results indicate that perceptual learning may enhance signal strength, and provide important constraints for theories of learning.

Axis of elongation can determine reference frames for object perception.

Object perception remains constant in the face of much retinal image variability. One way to achieve such constancy is to represent objects with respect to a reference frame, and many theories of object recognition assume that, among other factors, observers use objects' primary axes of elongation to derive reference frames. However, the limited research directly addressing this assumption suggests that the role of axis of elongation may not be as central as previously though (Palmer, 1990; Quinlan & Humphreys, 1993). The present study re-examines elongation's role in determining reference frames, adapting Palmer's paradigm to determine the extent to which surrounds bias the perception of ambiguously oriented objects: equilateral triangles. When surrounds were oriented inconsistently with experimentally constrained correct responses, response times increased with surrounds' elongation. These results hold for both symmetric and asymmetric surrounds, suggesting that elongation alone is sufficient to construct a reference frame for object perception.

Spatial phase differences can drive apparent motion.

Can shape differences drive apparent motion? Results from previous research are equivocal. Much of the confusion may be due to the use of relatively complex stimuli: letters or geometric shapes, comprising many spatial frequencies, phases, orientations, and contrasts. We focus on relatively simple stimuli: Gaussian damped f+nf compound sinewave gratings. We examine whether relative phase differences, which are critical for shape perception, can drive apparent motion. We find that some, but not all, phase differences can drive apparent motion. Specifically, stimuli that are easily discriminable and perceptually dissimilar can affect the solution of the correspondence problem. In this case, observers consistently perceive stimuli in one frame moving to the position of perceptually similar stimuli in the next frame. This general result holds over a wide range of spatial frequencies, orientations, and contrasts. Implications for theories of motion processing are discussed.

Pigeons do not complete partly occluded figures.

One of the most common obstacles to object perception is the fact that objects often occlude parts of themselves and parts of other objects. Perceptual completion has been studied extensively in humans, and researchers have shown that humans do complete partly occluded objects. In an effort to understand more about the mechanisms underlying completion, recent research has extended the study of perceptual completion to other mammalian species. Monkeys and mice also seem to complete two-dimensional representations of partly occluded objects. The present study addresses the question of whether this capacity generalizes to a nonmammalian species, the pigeon (Columba livia). The results point to a limit of the generalizability of perceptual completion: pigeons do not complete partly occluded figures.