A BCI-Based Study on the Relationship Between the SSVEP and Retinal Eccentricity in Overt and Covert Attention

Covert attention aids us in monitoring the environment and optimizing performance in visual tasks. Past behavioral studies have shown that covert attention can enhance spatial resolution. However, electroencephalography (EEG) activity related to neural processing between central and peripheral vision has not been systematically investigated. Here, we conducted an EEG study with 25 subjects who performed covert attentional tasks at different retinal eccentricities ranging from 0.75° to 13.90°, as well as tasks involving overt attention and no attention. EEG signals were recorded with a single stimulus frequency to evoke steady-state visual evoked potentials (SSVEPs) for attention evaluation. We found that the SSVEP response in fixating at the attended location was generally negatively correlated with stimulus eccentricity as characterized by Euclidean distance or horizontal and vertical distance. Moreover, more pronounced characteristics of SSVEP analysis were also acquired in overt attention than in covert attention. Furthermore, offline classification of overt attention, covert attention, and no attention yielded an average accuracy of 91.42%. This work contributes to our understanding of the SSVEP representation of attention in humans and may also lead to brain-computer interfaces (BCIs) that allow people to communicate with choices simply by shifting their attention to them.

The Relationship Between Perifoveal L-Cone Isolating Visual Acuity and Cone Photoreceptor Spacing—Understanding the Transition Between Healthy Aging and Early AMD

Background: Age-related macular degeneration (AMD) is a multifactorial degenerative disorder that can lead to irreversible loss of visual function, with aging being the prime risk factor. However, knowledge about the transition between healthy aging and early AMD is limited. We aimed to examine the relationship between psychophysical measures of perifoveal L-cone acuity and cone photoreceptor structure in healthy aging and early AMD.Methods and Results: Thirty-nine healthy participants, 10 with early AMD and 29 healthy controls were included in the study. Multimodal high-resolution retinal images were obtained with adaptive-optics scanning-light ophthalmoscopy (AOSLO), optical-coherence tomography (OCT), and color fundus photographs. At 5 degrees retinal eccentricity, perifoveal L-cone isolating letter acuity was measured with psychophysics, cone inner segment and outer segment lengths were measured using OCT, while cone density, spacing, and mosaic regularity were measured using AOSLO. The Nyquist sampling limit of cone mosaic (Nc) was calculated for each participant. Both L-cone acuity and photoreceptor inner segment length declined with age, but there was no association between cone density nor outer segment length and age. A multiple regression showed that 56% of the variation in log L-cone acuity was accounted for by Nc when age was taken into account. Six AMD participants with low risk of progression were well within confidence limits, while two with medium-to-severe risk of progression were outliers. The observable difference in cone structure between healthy aging and early AMD was a significant shortening of cone outer segments.Conclusion: The results underscore the resilience of cone structure with age, with perifoveal functional changes preceding detectable changes in the cone photoreceptor mosaic. L-cone acuity is a sensitive measure for assessing age-related decline in this region. The transition between healthy aging of cone structures and changes in cone structures secondary to early AMD relates to outer segment shortening.

Vergence accuracy in an autostereoscopic display

When fixating an object, observers typically under or over-converge by a small amount, a phenomenon known as "fixation disparity". Fixation disparity is typically measured with physical fixation targets and dichotically presented nonius lines. Here we made fixation disparity measurements with an autostereoscopic display, varying the retinal eccentricity and disparity of the fixation targets. Measurements were made in a group of four practiced observers and in a group of thirteen experimentally naïve observers. Fixation disparities with a zero-disparity target were in the direction of fixation behind the plane of the screen and the magnitude of the fixation disparity grew with the eccentricity of the fixation targets (1-5 deg in the practiced observers and 1 – 10 deg in the naïve observers). Fixation disparity also increased with increasing disparity of the targets, especially when they were presented at crossed disparities. Fixation disparities were larger overall for naïve observers who additionally did not converge in front of the screen when vergence demand was created by crossed disparity fusion locks presented at 5 and 10 deg eccentricities.

A dynamic pink noise paradigm to assess spatial-temporal attention without biasing perceptual processing

Psychophysical paradigms measure visual attention via localized test items to which observers must react or whose features have to be discriminated. These items, however, potentially interfere with the intended measurement as they bias observers' spatial and temporal attention to their location and presentation time. Furthermore, visual sensitivity for conventional test items naturally decreases with retinal eccentricity, which prevents direct comparison of central and peripheral attention assessments. We developed a stimulus that overcomes these limitations. A brief oriented discrimination signal is seamlessly embedded into a continuously changing 1/f noise field, such that observers cannot anticipate potential test locations or times. Using our new paradigm, we demonstrate that local orientation discrimination accuracy for 1/f filtered signals is largely independent of retinal eccentricity. Moreover, we show that items present in the visual field indeed shape the distribution of visual attention, suggesting that classical studies investigating the spatiotemporal dynamics of visual attention via localized test items may have obtained a biased measure. We recommend our paradigm as an efficient method to evaluate the spatial and temporal spread of visual attention.

Retinal factors of visual sensitivity in the human fovea

Humans direct their gaze towards visual objects of interest such that the retinal images of fixated objects fall onto the fovea, a small anatomically and physiologically specialized region of the retina displaying highest visual fidelity. One striking anatomical feature of the fovea is its non-uniform cellular topography, with a steep decline of cone photoreceptor density and outer segment length with increasing distance from its center. We here assessed in how far the specific cellular organization of the foveola is reflected in visual function. Increment sensitivity to small spot visual stimuli (1 x 1 arcmin, 543 nm light) was recorded psychophysically in 4 human participants at 17 locations placed concentric within a 0.2-degree diameter around the preferred retinal locus of fixation with adaptive optics scanning laser ophthalmoscopy based microstimulation. While cone density as well as maximum outer segment length differed significantly among the four tested participants, the range of observed threshold was similar, yielding an average increment threshold of 3.3 ± 0.2 log10 photons at the cornea. Thresholds were correlated with retinal eccentricity, as well as cone density and outer segment length. Biophysical simulation allowed to develop a model of foveal sensitivity based on these parameters, explaining at least 37% of the observed threshold variability. Based on high reproducibility in replicate testing, the residual variability is assumed to be caused by individual cone and bipolar cell weighting at the specific target locations.

Investigating Visual Crowding of Objects in Complex Real-World Scenes

Visual crowding, the impairment of object recognition in peripheral vision due to flanking objects, has generally been studied using simple stimuli on blank backgrounds. While crowding is widely assumed to occur in natural scenes, it has not been shown rigorously yet. Given that scene contexts can facilitate object recognition, crowding effects may be dampened in real-world scenes. Therefore, this study investigated crowding using objects in computer-generated real-world scenes. In two experiments, target objects were presented with four flanker objects placed uniformly around the target. Previous research indicates that crowding occurs when the distance between the target and flanker is approximately less than half the retinal eccentricity of the target. In each image, the spacing between the target and flanker objects was varied considerably above or below the standard (0.5) threshold to either suppress or facilitate the crowding effect. Experiment 1 cued the target location and then briefly flashed the scene image before participants could move their eyes. Participants then selected the target object’s category from a 15-alternative forced choice response set (including all objects shown in the scene). Experiment 2 used eye tracking to ensure participants were centrally fixating at the beginning of each trial and showed the image for the duration of the participant’s fixation. Both experiments found object recognition accuracy decreased with smaller spacing between targets and flanker objects. Thus, this study rigorously shows crowding of objects in semantically consistent real-world scenes.

The spatial distribution of ERGs reflecting luminance and L-/M-cone-opponent signals

Purpose To study the spatial retinal distribution of electroretinographic (ERG) responses that reflect signals in the L-/M-cone-opponent and luminance post-receptoral pathways. Methods ERG recordings to heterochromatic stimuli (sinusoidal counter-phase modulation of red and green LED light sources) were performed, while varying fractions of red and green modulation. Two temporal frequencies of the stimuli were employed: 12 Hz to record ERGs that reflect L-/M-cone-opponent signal and 36 Hz for recording ERG signals sensitive to stimulus luminance. Stimuli were about 20° in diameter and projected on various retinal locations: the fovea and four eccentricities (10°, 19°, 28° and 35°), each presented nasally, temporally, inferiorly and superiorly from the fovea. Results The 36 Hz stimuli elicited responses that strongly varied with red fraction and were minimal at iso-luminance. Moreover, response phases changed abruptly at the minimum by 180°. In contrast, the responses to the 12 Hz stimuli had amplitudes and phases that changed more gradually with red fraction. The 36 Hz response amplitudes were maximal close to the fovea and sharply decreased with increasing distance from the fovea. The responses to 12 Hz stimuli were more broadly distributed across the retina. Conclusions In the present study, it was found that retinal eccentricity and direction from the fovea have distinct effects on ERGs reflecting different post-receptoral mechanisms. The results are in accord with previous findings that ERGs to 12 Hz stimuli are predominantly determined by the red–green chromatic content of the stimuli, thus reflecting activation in the L-/M-cone-opponent pathway, while responses to 36 Hz stimuli manifest post-receptoral luminance-dependent activation. We found that the response in the cone-opponent pathway is broadly comparable across the retina; in comparison, response amplitude of the luminance pathway strongly depends on retinal stimulus position.

The role of sensory uncertainty in simple contour integration

Perceptual organization is the process of grouping scene elements into whole entities. A classic example is contour integration, in which separate line segments are perceived as continuous contours. Uncertainty in such grouping arises from scene ambiguity and sensory noise. Some classic Gestalt principles of contour integration, and more broadly, of perceptual organization, have been re-framed in terms of Bayesian inference, whereby the observer computes the probability that the whole entity is present. Previous studies that proposed a Bayesian interpretation of perceptual organization, however, have ignored sensory uncertainty, despite the fact that accounting for the current level of perceptual uncertainty is one of the main signatures of Bayesian decision making. Crucially, trial-by-trial manipulation of sensory uncertainty is a key test to whether humans perform near-optimal Bayesian inference in contour integration, as opposed to using some manifestly non-Bayesian heuristic. We distinguish between these hypotheses in a simplified form of contour integration, namely judging whether two line segments separated by an occluder are collinear. We manipulate sensory uncertainty by varying retinal eccentricity. A Bayes-optimal observer would take the level of sensory uncertainty into account—in a very specific way—in deciding whether a measured offset between the line segments is due to non-collinearity or to sensory noise. We find that people deviate slightly but systematically from Bayesian optimality, while still performing “probabilistic computation” in the sense that they take into account sensory uncertainty via a heuristic rule. Our work contributes to an understanding of the role of sensory uncertainty in higher-order perception.

EEG signatures of contextual influences on visual search with real scenes

The use of scene context is a powerful way by which biological organisms guide and facilitate visual search. Although many studies have shown enhancements of target-related electroencephalographic activity (EEG) with synthetic cues, there have been fewer studies demonstrating such enhancements during search with scene context and objects in real world scenes. Here, observers covertly searched for a target in images of real scenes while we used EEG to measure the steady state visual evoked response to objects flickering at different frequencies. The target appeared in its typical contextual location or out of context while we controlled for low-level properties of the image including target saliency against the background and retinal eccentricity. A pattern classifier using EEG activity at the relevant modulated frequencies showed target detection accuracy increased when the target was in a contextually appropriate location. A control condition for which observers searched the same images for a different target orthogonal to the contextual manipulation, resulted in no effects of scene context on classifier performance, confirming that image properties cannot explain the contextual modulations of neural activity. Pattern classifier decisions for individual images was also related to the aggregated observer behavioral decisions for individual images. Together, these findings demonstrate target-related neural responses are modulated by scene context during visual search with real world scenes and can be related to behavioral search decisions.Significance StatementContextual relationships among objects are fundamental for humans to find objects in real world scenes. Although there is a larger literature understanding the brain mechanisms when a target appears at a location indicated by a synthetic cue such as an arrow or box, less is known about how the scene context modulates target-related neural activity. Here we show how neural activity predictive of the presence of a searched object in cluttered real scenes increases when the target object appears at a contextual location and diminishes when it appears at a place that is out of context. The results increase our understanding of how the brain processes real scenes and how context modulates object processing.

Neural sources of letter and Vernier acuity

Visual acuity can be measured in many different ways, including with letters and Vernier offsets. Prior psychophysical work has suggested that the two acuities are strongly linked given that they both depend strongly on retinal eccentricity and both are similarly affected in amblyopia. Here we used high-density EEG recordings to ask whether the underlying neural sources are common as suggested by the psychophysics or distinct. To measure visual acuity for letters, we recorded evoked potentials to 3 Hz alternations between intact and scrambled text comprised of letters of varying size. To measure visual acuity for Vernier offsets, we recorded evoked potentials to 3 Hz alternations between bar gratings with and without a set of Vernier offsets. Both alternation types elicited robust activity at the 3 Hz stimulus frequency that scaled in amplitude with both letter and offset size, starting near threshold. Letter and Vernier offset responses differed in both their scalp topography and temporal dynamics. The earliest evoked responses to letters occurred on lateral occipital visual areas, predominantly over the left hemisphere. Later responses were measured at electrodes over early visual cortex, suggesting that letter structure is first extracted in second-tier extra-striate areas and that responses over early visual areas are due to feedback. Responses to Vernier offsets, by contrast, occurred first at medial occipital electrodes, with responses at later time-points being more broadly distributed—consistent with feedforward pathway mediation. The previously observed commonalities between letter and Vernier acuity may be due to common bottlenecks in early visual cortex but not because the two tasks are subserved by a common network of visual areas.