A Study of Separation Difficulty^*: Its Relationship to Visual Acuity in Normal and Amblyopic Eyes

Visual stimuli presented around the time of a saccade have been shown to be perceived differently by the visual system, including a reduction in the harmful impact of flankers (crowding). However, whether the effects observed are due strictly to crowding remains controversial, and the effects have only been measured with large saccades in peripheral vision. Here we investigate how crowded stimuli placed 20 arc minutes from the center of gaze are affected by an upcoming microsaccade. The stimulus consisted of a rotated T of size 6.25 arcminutes that was either unflanked, surrounded by four flankers (Experiment 1), or surrounded by two flankers that were positioned either radially or tangentially (Experiments 2 and 3). In 80 % of trials, subjects made voluntary microsaccades to the target when cued, and in the remaining 20 % of the trials subjects continued to maintain fixation. In Experiments 1 and 2, subjects were required to saccade to the same location as the target, while in Experiment 3 subjects saccaded to a different location ∼ 20 arc min to the upper left of the target. Thus, we provide evidence for two separable pre-saccadic benefits for crowded parafoveal targets: one isotropizes the crowding zone for stimuli presented 200 to 125 ms before microsaccadic onset, and another provides a benefit exclusively for microsaccade targets surrounded by tangential flankers in the presence of imminent microsaccades. Two possible mechanisms are attentional enhancement and predictive remapping of receptive fields, respectively.

Integration of information over the CNS is an important neural process that affects our ability to perceive and react to the environment. The visual system is required to continuously integrate information arriving from two different sources (the eyes) to create a coherent percept with high spatiotemporal precision. Although this neural integration of information is assumed to be critical for visual performance, it can be impaired under some pathological or developmental conditions. Here we took advantage of a unique developmental condition, amblyopia (“lazy eye”), which is characterized by an impaired temporal synchronization between the two eyes, to meticulously study the effect of synchronization on the integration of binocular visual information. We measured the eyes’ asynchrony and compensated for it (with millisecond temporal resolution) by providing time-shifted stimuli to the eyes. We found that the re-synchronization of the ocular input elicited a significant improvement in visual functions, and binocular functions, such as binocular summation and stereopsis, were regained. This phenomenon was also evident in neurophysiological measures. Our results can shed light on other neural processing aspects and might also have translational relevance for the field of training, rehabilitation, and perceptual learning.

Crowding refers to impaired object recognition of peripheral visual targets caused by nearby flankers. It has been shown that the response to a word was faster when it was preceded by a semantically related than unrelated crowded prime, demonstrating that semantic priming survives crowding. This study examines neural correlates of semantic priming under visual crowding using magnetoencephalography with four conditions: prime (isolated, crowded) x prime-target relationship (related, unrelated). Participants judged whether the target was a word or a nonword. We found significant differences in θ activity at the left inferior frontal gyrus (IFG) for both isolated and crowded primes when comparing the unrelated and related conditions, although the activation was delayed with the crowded prime compared to the isolated prime. The locations within the IFG were also different: theta-band activation was at BA 45 in the isolated condition and at BA 47 in the crowded condition. Phase-locking-value analysis revealed that bilateral IFG was more synchronized with unrelated prime-target pairs than related pairs regardless of whether the primes were isolated or crowded, indicating the recruitment of the right hemisphere when the prime-target semantic relationship was remote. Finally, the distinct waveform patterns found in the isolated and crowded conditions from both the source localization and PLV analysis suggest different neural mechanisms for processing semantic information with isolated primes versus crowded primes.

Crowding refers to the deleterious visual interaction among nearby objects. Does maximal crowding occur when objects are closest to one another in space and time? We examined how crowding depends on the spatial and temporal proximity, retinally and perceptually, between a target and flankers. Our target was a briefly flashed T-stimulus presented at 10° right of fixation (3-o’clock position). It appeared at different target-onset-to-flanker asynchronies with respect to the instant when a pair of flanking Ts, revolving around the fixation target, reached the 3-o’clock position. Observers judged the orientation of the target-T (the crowding task), or its position relative to the revolving flankers (the flash-lag task). Performance was also measured in the absence of flanker motion: target and flankers were either presented simultaneously (closest retinal temporal proximity) with different angular spatial offsets, or were presented collinearly (closest retinal spatial proximity) with different temporal onset asynchronies. We found that neither retinal nor perceptual spatial or temporal proximity could account for when maximal crowding occurred. Simulations using a model based on feed-forward interactions between sustained and transient channels in static and motion pathways, taking into account the differential response latencies, can explain the crowding functions observed under various spatio-temporal conditions between the target and flankers.

Pure alexia and prosopagnosia traditionally have been seen as prime examples of dissociated, category-specific agnosias affecting reading and face recognition, respectively. More recent accounts have moved towards domain-independent explanations that postulate potential cross-links between different types of visual agnosia. According to one proposal, abnormal crowding, i.e. the impairment of recognition when features of adjacent objects are positioned too closely to each other, might provide a unified account for the perceptual deficits experienced by an agnosic patient. An alternative approach is based on the notion of complementary visual subsystems favouring the processing of abstract categories and specific exemplars, respectively. To test predictions of these two approaches with regard to pure alexia and prosopagnosia, we present previously unpublished data on digit recognition and visual crowding from two in the neuropsychological literature extensively studied patients, KD and MT (e.g., Campbell et al., 1986; Landis and Regard, 1988; Rentschler et al., 1994). Patient MT, diagnosed with pure alexia, showed pronounced abnormal foveal crowding, whereas KD, diagnosed with prosopagnosia, did not. These results form a distinct double dissociation with the performance of the two patients in other perceptual classification tasks involving Gabor micropatterns and textures, as well as Glass patterns, which revealed a significantly greater impairment in KD relative to MT. Based on an analysis of the specific task demands we argue that prosopagnosia and pure alexia may involve complementary deficits in instantiation and abstraction, respectively, during perceptual classification, beyond any category specificity. Such an explanation appears in line with previous distinctions between a predominantly left-hemispheric, abstract-category and a predominantly right-hemispheric, specific-exemplar subsystem underlying object recognition.

Nearby flanking objects degrade visual resolution. If the flankers are similar to the acuity target, this influence is called crowding (CW), whereas if the flanking stimuli are simple bars then the phenomenon is known as contour interaction (CI). The aim of this study was to compare the influence of the number and position of flankers on foveal CW and CI to investigate possible differences in mechanism of these two effects. Five normal observers viewed single, foveally presented Sloan letters surrounded by 1, 2 or 4 flankers (either a Sloan letter or one-stroke-width bars), presented at several edge-to-edge separations. Single flankers were presented in the right, left, top or bottom position, 2 flankers were placed equally to the right and left or top and bottom of the central target, and 4 flankers were equally spaced in all four directions. Percent correct letter identification was determined for each type, number, position and separation of flankers and confusion matrices were constructed for separations equal to 20% and 100% letter width. Increasing the number of flankers caused an increase in the magnitude of both phenomena. CW showed a greater magnitude than CI for higher numbers of flankers. Analysis of confusion matrices suggests that in addition to the edge-to-edge interaction that appears to mediate CI, letter substitution and feature pooling contribute significantly to CW when higher numbers of flankers are presented. Foveal CW is more strongly influenced by an increase in the number of flankers than CI, which can be explained by the presence of additional interaction effects.