Evolutionary and developmental specialization of foveal cell types in the marmoset.

In primates, high-acuity vision is mediated by the fovea, a small specialized central region of the retina. The fovea, unique to the anthropoid lineage among mammals, undergoes notable neuronal morphological changes during postnatal maturation. However, the extent of cellular similarity across anthropoid foveas and the molecular underpinnings of foveal maturation remain unclear. Here, we used high-throughput single-cell RNA sequencing to profile retinal cells of the common marmoset (Callithrix jacchus), an early divergent in anthropoid evolution from humans, apes, and macaques. We generated atlases of the marmoset fovea and peripheral retina for both neonates and adults. Our comparative analysis revealed that marmosets share almost all their foveal types with both humans and macaques, highlighting a conserved cellular structure among primate foveas. Furthermore, by tracing the developmental trajectory of cell types in the foveal and peripheral retina, we found distinct maturation paths for each. In-depth analysis of gene expression differences demonstrated that cone photoreceptors and Müller glia (MG), among others, show the greatest molecular divergence between these two regions. Utilizing single-cell ATAC-seq and gene-regulatory network inference, we uncovered distinct transcriptional regulations differentiating foveal cones from their peripheral counterparts. Further analysis of predicted ligand-receptor interactions suggested a potential role for MG in supporting the maturation of foveal cones. Together, these results provide valuable insights into foveal development, structure, and evolution.

Toward a theory of perspective perception in pictures.

This paper reviews projection models and their perception in realistic pictures, and proposes hypotheses for three-dimensional (3D) shape and space perception in pictures. In these hypotheses, eye fixations, and foveal vision play a central role. Many past theories and experimental studies focus solely on linear perspective. Yet, these theories fail to explain many important perceptual phenomena, including the effectiveness of nonlinear projections. Indeed, few classical paintings strictly obey linear perspective, nor do the best distortion-avoidance techniques for wide-angle computational photography. The hypotheses here employ a two-stage model for 3D human vision. When viewing a picture, the first stage perceives 3D shape for the current gaze. Each fixation has its own perspective projection, but, owing to the nature of foveal and peripheral vision, shape information is obtained primarily for a small region of the picture around the fixation. As a viewer moves their eyes, the second stage continually integrates some of the per-gaze information into an overall interpretation of a picture. The interpretation need not be geometrically stable or consistent over time. It is argued that this framework could explain many disparate pictorial phenomena, including different projection styles throughout art history and computational photography, while being consistent with the constraints of human 3D vision. The paper reviews open questions and suggests new studies to explore these hypotheses.

Featural Representation and Internal Noise Underlie the Eccentricity Effect in Contrast Sensitivity.

Human visual performance for basic visual dimensions (e.g., contrast sensitivity and acuity) peaks at the fovea and decreases with eccentricity. The eccentricity effect is related to the larger visual cortical surface area corresponding to the fovea, but it is unknown if differential feature tuning contributes to this eccentricity effect. Here, we investigated two system-level computations underlying the eccentricity effect: featural representation (tuning) and internal noise. Observers (both sexes) detected a Gabor embedded in filtered white noise which appeared at the fovea or one of four perifoveal locations. We used psychophysical reverse correlation to estimate the weights assigned by the visual system to a range of orientations and spatial frequencies (SFs) in noisy stimuli, which are conventionally interpreted as perceptual sensitivity to the corresponding features. We found higher sensitivity to task-relevant orientations and SFs at the fovea than that at the perifovea, and no difference in selectivity for either orientation or SF. Concurrently, we measured response consistency using a double-pass method, which allowed us to infer the level of internal noise by implementing a noisy observer model. We found lower internal noise at the fovea than that at the perifovea. Finally, individual variability in contrast sensitivity correlated with sensitivity to and selectivity for task-relevant features as well as with internal noise. Moreover, the behavioral eccentricity effect mainly reflects the foveal advantage in orientation sensitivity compared with other computations. These findings suggest that the eccentricity effect stems from a better representation of task-relevant features and lower internal noise at the fovea than that at the perifovea.

Dynamic changes in ocular shape during human development and its implications for retina fovea formation.

The human fovea is known for its distinctive pit-like appearance, which results from the displacement of retinal layers superficial to the photoreceptors cells. The photoreceptors are found at high density within the foveal region but not the surrounding retina. Efforts to elucidate the mechanisms responsible for these unique features have ruled out cell death as an explanation for pit formation and changes in cell proliferation as the cause of increased photoreceptor density. These findings have led to speculation that mechanical forces acting within and on the retina during development underly the formation of foveal architecture. Here we review eye morphogenesis and retinal remodeling in human embryonic development. Our meta-analysis of the literature suggests that fovea formation is a protracted process involving dynamic changes in ocular shape that start early and continue throughout most of human embryonic development. From these observations, we propose a new model for fovea development.

Parafoveal processing in bilingual readers: Semantic access within but not across languages.

Prior research has investigated the quality of information a reader can extract from upcoming parafoveal words. However, very few studies have considered parafoveal processing in bilingual readers, who may differ from monolinguals due to slower lexical access and susceptibility to cross-language activation. This eye-tracking experiment, therefore, investigated how bilingual readers process parafoveal semantic information within and across languages. We used the boundary technique to replace a preview word in a sentence with a different target word during the first rightward saccade from the pretarget region. We manipulated both preview language (nonswitch vs. code-switch) and semantic relatedness (synonym/translation vs. unrelated) between previews and targets. Upon fixation, target words always appeared in the same language as the rest of the sentence to create an essentially monolingual language context. Semantic preview benefits emerged for nonswitched synonym previews but not for code-switched translation previews. Furthermore, participants skipped code-switched previews less often than nonswitched previews and no more often than previews that were unfamiliar to them. These data suggest that bilinguals can extract within-language semantic information from the parafovea in both native and nonnative languages, but that cross-language words are not accessible while reading in a monolingual language mode, as per the partial selectivity hypothesis of bilingual language control. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Foveal feedback in perceptual processing: Contamination of neural representations and task difficulty effects.

Visual object recognition was traditionally believed to rely on a hierarchical feedforward process. However, recent evidence challenges this notion by demonstrating the crucial role of foveal retinotopic cortex and feedback signals from higher-level visual areas in processing peripheral visual information. The nature of the information conveyed through foveal feedback remains a topic of debate. To address this, we conducted a study employing a foveal mask paradigm with varying stimulus-mask onset asynchronies in a peripheral same/different task, where peripheral objects exhibited different degrees of similarity. Our hypothesis posited that simultaneous arrival of feedback and mask information in the foveal cortex would lead to neural contamination, biasing perception. Notably, when the two peripheral objects were identical, we observed a significant increase in the number of "different" responses, peaking at approximately 100 ms. Similar effect was found when the objects were dissimilar, but with an overall later timing (around 150 ms). No significant difference was found when comparing easy (dissimilar objects) and difficult trials (similar objects). The findings challenge the hypothesis that foveation planning alone accounts for the observed effects. Instead, these and previous observations support the notion that the foveal cortex serves as a visual sketchpad for maintaining and manipulating task-relevant information.

Maturation and refinement of the maculae and foveae in the Anolis sagrei lizard.

The fovea is a pit in the center of the macula, which is a region of the retina with a high concentration of photoreceptor cells, which accounts for a large degree of visual acuity in primates. The maturation of this primate visual acuity area is characterized by the shallowing and widening of the foveal pit, a decrease in the diameter of the rod-free zone, and an increase in photoreceptor cells packing after birth. Maturation occurs concurrently with progressing age, increasing eye size, and retinal length/area. These observations have led to the hypothesis that the maturation of the fovea might be a function of mechanical variables that remodel the retina. However, this has never been explored outside of primates. Here, we take advantage of the Anolis sagrei lizard, which has a bifoveated retina, to study maturation of the fovea and macula. Eyes were collected from male and female lizards-hatchling, 2-month, 4-month, 6-month, and adult. We found that Anolis maculae undergo a maturation process somewhat different than what has been observed in primates. Anole macular diameters actually increase in size and undergo minimal photoreceptor cell packing, possessing a near complete complement of these cells at the time of hatching. As the anole eye expands, foveal centers experience little change in overall retina cell density with most cell redistribution occurring at macular borders and peripheral retina areas. Gene editing technology has recently been developed in lizards; this study provides a baseline of normal retina maturation for future genetic manipulation studies in anoles.

Out of sight, out of mind: Foveal processing is necessary for semantic integration of words into sentence context.

Word recognition begins before a reader looks directly at a word, as demonstrated by the parafoveal preview benefit and word skipping. Both low-level form and high-level semantic features can be accessed in parafoveal vision and used to promote reading efficiency. However, words are not recognized in isolation during reading; once a semantic representation is retrieved, it must be integrated with the broader sentence context. One open question about parafoveal processing is whether it is limited to shallow stages of lexico-semantic activation or extends to semantic integration. In the present two-experiment study, we recorded event-related brain potentials in response to a sentence-final word that was presented in foveal or parafoveal vision and was either expected, unexpected, or anomalous in the sentence context. We found that word recognition, indexed by the N400, ensued regardless of perception location whereas identification of the semantic fit of a word in its sentence context, indexed by the late positive component, was only observed for foveally perceived but not parafoveally perceived words. This pattern was not sensitive to task differences that promote different levels of orthographic scrutiny, as manipulated between the two experiments. These findings demonstrate separate roles for parafoveal and foveal processing in reading. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Cross-language semantic parafoveal preview benefits in bilinguals.

The aim of this project was to identify factors contributing to cross-language semantic preview benefits. In Experiment 1, Russian-English bilinguals read English sentences with Russian words presented as parafoveal previews. The gaze-contingent boundary paradigm was used to present sentences. Critical previews were cognate translations of the target word (CTAPT-START), noncognate translations (CPOK-TERM), or interlingual homograph translations (MOPE-SEA). A semantic preview benefit (i.e., shorter fixation durations for related than unrelated previews) was observed for cognate and interlingual homograph translations, but not for noncognate translations. In Experiment 2, English-French bilinguals read English sentences with French words used as parafoveal previews. Critical previews were interlingual homograph translations of the target word (PAIN-BREAD) or interlingual homograph translations with a diacritic added (PÁIN-BREAD). A robust semantic preview benefit was found only for interlingual homographs without diacritics, although both preview types produced a semantic preview benefit in the total fixation duration. Our findings suggest that semantically related previews need to have substantial orthographic overlap with words in the target language to produce cross-language semantic preview benefits in early eye fixation measures. In terms of the Bilingual Interactive Activation+ model, the preview word may need to activate the language node for the target language before its meaning is integrated with that of the target word. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Investigating the role of the foveal cortex in peripheral object discrimination.

Peripheral object discrimination is hindered by a central dynamic mask presented between 150 and 300 ms after stimulus onset. The mask is thought to interfere with task-relevant feedback coming from higher visual areas to the foveal cortex in V1. Fan et al. (2016) supported this hypothesis by showing that the effect of mask can be further delayed if the task requires mental manipulation of the peripheral target. The main purpose of this study was to better characterize the temporal dynamics of foveal feedback. Specifically, in two experiments we have shown that (1) the effect of foveal noise mask is sufficiently robust to be replicated in an online data collection (2) in addition to a change in sensitivity the mask affects also the criterion, which becomes more conservative; (3) the expected dipper function for sensitivity approximates a quartic with a global minimum at 94 ms, while the best fit for criterion is a quintic with a global maximum at 174 ms; (4) the power spectrum analysis of perceptual oscillations in sensitivity data shows a cyclic effect of mask at 3 and 12 Hz. Overall, our results show that foveal noise affects sensitivity in a cyclic manner, with a global dip emerging earlier than previously found. The noise also affects the response bias, even though with a different temporal profile. We, therefore, suggest that foveal noise acts on two distinct feedback mechanisms, a faster perceptual feedback followed by a slower cognitive feedback.

Fixation-related fMRI analysis reveals the neural basis of parafoveal processing in self-paced reading of Chinese words.

While parafoveal word processing plays an important role in natural reading, the underlying neural mechanism remains unclear. The present study investigated the neural basis of parafoveal processing during Chinese word reading with the co-registration of eye-tracking and functional magnetic resonance imaging (fMRI) using fixation-related fMRI analysis. In the gaze-contingent boundary paradigm, preview conditions (words that are identical, orthographically similar, and unrelated to target words), pre-target word frequency and target word frequency were manipulated. When fixating the pre-target word, the identical preview condition elicited lower brain activation in the left fusiform gyrus relative to unrelated and orthographically similar preview conditions and there were significant interactions of preview condition and pre-target word frequency on brain activation of the left middle frontal gyrus, left fusiform gyrus and supplementary motor area. When fixating the target word, there was a significant main effect of preview condition on brain activation of the right fusiform gyrus and a significant interaction of preview condition and pre-target word frequency on brain activation of the left middle frontal gyrus. These results suggest that fixation-related brain activation provides immediate measures and new perspectives to understand the mechanism of parafoveal processing in self-paced reading.

Severe distortion in the representation of foveal visual image locations in short-term memory.

The foveal visual image region provides the human visual system with the highest acuity. However, it is unclear whether such a high fidelity representational advantage is maintained when foveal image locations are committed to short-term memory. Here, we describe a paradoxically large distortion in foveal target location recall by humans. We briefly presented small, but high contrast, points of light at eccentricities ranging from 0.1 to 12°, while subjects maintained their line of sight on a stable target. After a brief memory period, the subjects indicated the remembered target locations via computer controlled cursors. The biggest localization errors, in terms of both directional deviations and amplitude percentage overshoots or undershoots, occurred for the most foveal targets, and such distortions were still present, albeit with qualitatively different patterns, when subjects shifted their gaze to indicate the remembered target locations. Foveal visual images are severely distorted in short-term memory.

Parafoveal words can modulate sentence meaning: Electrophysiological evidence from an RSVP-with-flanker task.

During natural reading, readers can take up some visual information from not-yet-fixated words to the right of the current fixation and it is well-established that this parafoveal preview facilitates the subsequent foveal processing of the word. However, the extraction and integration of word meaning from parafoveal words and their possible influence on the semantic content of the sentence are controversial. In the current study, we recorded event-related potentials (ERPs) in the RSVP-with-flankers paradigm to test whether and how updates of sentential meaning, based only on parafoveal information, may influence the subsequent foveal processing. In Chinese sentences, the congruency of parafoveal and foveal target words with the sentence was orthogonally manipulated. In contrast to previous research, we also controlled for potentially confounding effects of parafoveal-to-foveal repetition priming (identity preview effects) on the N400. Crucially, we found that the classic effect of foveal congruency on the N400 component only appeared when the word in preview had been congruent with sentence meaning; in contrast, there was no N400 as a function of foveal incongruency when the preview word had also been incongruent. These results indicate that sentence meaning rapidly adapts to parafoveal preview, altering the semantic context for the subsequently fixated word. We also show that correct parafoveal preview generally attenuates the N400 once a word is fixated, regardless of congruency. Taken together, our findings underline the highly generative and adaptive framework of language comprehension.

Achromatic doublet electrowetting prism array for beam steering device in foveated display.

A foveated display is a technology that can solve the problem of insufficient angular resolution (relative to the human eye) for near-eye display. In a high-resolution foveated display, a beam steering element is required to track the human gaze. An electrowetting prism array is a transmissive non-mechanical beam steering device, that allows a light and compact optical system to be configured and a large aperture possible. However, the view is obstructed by the sidewall of the prism array. When the size of the cell prism is 7mm, the prism array has an 87% fill-factor. To push the fill-factor to 100%, the cell prisms were magnified using a lens array. Image processing was performed such that the image produced by the lens array was identical to the original. Beam steering by refraction is accompanied by chromatic dispersion, which causes chromatic aberration, making colors appear blurry. The refractive index condition to reduce chromatic dispersion was obtained using the doublet structure of the electrowetting prism. The chromatic dispersion was reduced by 70% on average.

Preview frequency effects in reading: evidence from Chinese.

Studies about sentence reading have shown that visual and lexical information beyond the currently fixated word can be integrated across fixations. The gaze-contingent boundary paradigm has been used widely to explore the extent to which parafoveal information can be processed before a word is fixated on. However, a critical review of the current literature suggests that unrelated mask previews are an unlikely baseline control with zero lexical activation, blurring the nature of experimental effects observed in the paradigm. The present study, therefore, aimed at shedding light on the effect of parafoveal mask properties through a manipulation of preview word frequency. Low-frequency preview words that are unrelated to target words elicited a larger interference than high-frequency preview words. We discuss implications of the preview frequency effect for computational models of eye-movement control in reading.

The perceptual span is dynamically adjusted in response to foveal load by beginning readers.

The perceptual span describes the size of the visual field from which information is obtained during a fixation in reading. Its size depends on characteristics of writing system and reader, but-according to the foveal load hypothesis-it is also adjusted dynamically as a function of lexical processing difficulty. Using the moving window paradigm to manipulate the amount of preview, here we directly test whether the perceptual span shrinks as foveal word difficulty increases. We computed the momentary size of the span from word-based eye-movement measures as a function of foveal word frequency, allowing us to separately describe the perceptual span for information affecting spatial saccade targeting and temporal saccade execution. First fixation duration and gaze duration on the upcoming (parafoveal) word N + 1 were significantly shorter when the current (foveal) word N was more frequent. We show that the word frequency effect is modulated by window size. Fixation durations on word N + 1 decreased with high-frequency words N, but only for large windows, that is, when sufficient parafoveal preview was available. This provides strong support for the foveal load hypothesis. To investigate the development of the foveal load effect, we analyzed data from three waves of a longitudinal study on the perceptual span with German children in Grades 1 to 6. Perceptual span adjustment emerged early in development at around second grade and remained stable in later grades. We conclude that the local modulation of the perceptual span indicates a general cognitive process, perhaps an attentional gradient with rapid readjustment. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Foveal processing of emotion-informative facial features.

Certain facial features provide useful information for recognition of facial expressions. In two experiments, we investigated whether foveating informative features of briefly presented expressions improves recognition accuracy and whether these features are targeted reflexively when not foveated. Angry, fearful, surprised, and sad or disgusted expressions were presented briefly at locations which would ensure foveation of specific features. Foveating the mouth of fearful, surprised and disgusted expressions improved emotion recognition compared to foveating an eye or cheek or the central brow. Foveating the brow led to equivocal results in anger recognition across the two experiments, which might be due to the different combination of emotions used. There was no consistent evidence suggesting that reflexive first saccades targeted emotion-relevant features; instead, they targeted the closest feature to initial fixation. In a third experiment, angry, fearful, surprised and disgusted expressions were presented for 5 seconds. Duration of task-related fixations in the eyes, brow, nose and mouth regions was modulated by the presented expression. Moreover, longer fixation at the mouth positively correlated with anger and disgust accuracy both when these expressions were freely viewed (Experiment 2b) and when briefly presented at the mouth (Experiment 2a). Finally, an overall preference to fixate the mouth across all expressions correlated positively with anger and disgust accuracy. These findings suggest that foveal processing of informative features is functional/contributory to emotion recognition, but they are not automatically sought out when not foveated, and that facial emotion recognition performance is related to idiosyncratic gaze behaviour.

Perceptual-driven approach to statically foveated head-mounted displays.

A foveated display is a promising technique to realize displays offering both a large field of view (FOV) and high spatial resolution. Although several prior works have attempted to apply a foveation method to the design of a head-mounted display (HMD) system, the common method is based on a dual-resolution dynamic foveation scheme which is inevitably complex and has a high cost due to the requirements for multiple display sources, a 2D steering mechanism, and eye tracker. In this paper, a new perceptual-driven approach to the design of a statically foveated HMD is proposed with the goal of offering a wide FOV across which the degradation of the perceived image resolution is nearly imperceptible or minimal within regions of frequent eye movements. Compared to a dual-resolution discrete and dynamic foveation approach in the prior art, the static foveation approach will not only maintain resolution continuity but also eliminate the need for a scanning mechanism, multiple display sources, and an eyetracker, and therefore minimize hardware complexity. We present the general approach for creating a static foveation scheme, performance metrics for evaluating the perceived image quality, and the process of optimizing a foveation scheme to meet different requirements. Finally, we experimentally demonstrate and validate the proposed foveation scheme using a testbed system. Overall, we demonstrate a statically foveated scheme is capable of offering a display with a total 160° FOV, a constant resolution of 0.5 or 1 arcminutes per pixel within the ±10° region where frequent eye movements occur, an adequate resolution no less than 45% of peak resolution within the parafovea region of ±30°, and a data sampling efficiency as high as 90%.

Neural evidence for lexical parafoveal processing.

In spite of the reduced visual acuity, parafoveal information plays an important role in natural reading. However, competing models on reading disagree on whether words are previewed parafoveally at the lexical level. We find neural evidence for lexical parafoveal processing by combining a rapid invisible frequency tagging (RIFT) approach with magnetoencephalography (MEG) and eye-tracking. In a silent reading task, target words are tagged (flickered) subliminally at 60 Hz. The tagging responses measured when fixating on the pre-target word reflect parafoveal processing of the target word. We observe stronger tagging responses during pre-target fixations when followed by low compared with high lexical frequency targets. Moreover, this lexical parafoveal processing is associated with individual reading speed. Our findings suggest that reading unfolds in the fovea and parafovea simultaneously to support fluent reading.

Fast and nonuniform dynamics of perisaccadic vision in the central fovea.

Humans use rapid eye movements (saccades) to inspect stimuli with the foveola, the region of the retina where receptors are most densely packed. It is well established that visual sensitivity is generally attenuated during these movements, a phenomenon known as saccadic suppression. This effect is commonly studied with large, often peripheral, stimuli presented during instructed saccades. However, little is known about how saccades modulate the foveola and how the resulting dynamics unfold during natural visual exploration. Here we measured the foveal dynamics of saccadic suppression in a naturalistic high-acuity task, a task designed after primates' social grooming, which-like most explorations of fine patterns-primarily elicits minute saccades (microsaccades). Leveraging on recent advances in gaze-contingent display control, we were able to systematically map the perisaccadic time course of sensitivity across the foveola. We show that contrast sensitivity is not uniform across this region and that both the extent and dynamics of saccadic suppression vary within the foveola. Suppression is stronger and faster in the most central portion, where sensitivity is generally higher and selectively rebounds at the onset of a new fixation. These results shed light on the modulations experienced by foveal vision during the saccade-fixation cycle and explain some of the benefits of microsaccades.