Cortical Reorganization of Peripheral Vision Induced by Simulated Central Vision Loss.

When one's central vision is deprived, a spared part of the peripheral retina acts as a pseudofovea for fixation. The neural mechanisms underlying this compensatory adjustment remain unclear. Here we report cortical reorganization induced by simulated central vision loss. Human subjects of both sexes learned to place the target at an eccentric retinal locus outside their blocked visual field for object tracking. Before and after training, we measured visual crowding-a bottleneck of object identification in peripheral vision, using psychophysics and fMRI. We found that training led to an axis-specific reduction of crowding. The change of the crowding effect was reflected in the change of BOLD signal, as a release of cortical suppression in multiple visual areas starting as early as V1. Our findings suggest that the adult visual system is capable of reshaping its oculomotor control and sensory coding to adapt to impoverished visual input.SIGNIFICANCE STATEMENT By simulating central vision loss in normally sighted adults, we found that oculomotor training not only induces PRL, but also facilitates form processing in peripheral vision. As subjects learned to place the target at an eccentric retinal locus, "visual crowding"-the detrimental effect of clutter on peripheral object identification-was reduced. The reduction of the crowding effect was accompanied by a release of response suppression in the visual cortex. These findings indicate that the adult visual system is capable of reshaping the peripheral vision to adapt to central vision loss.

Contextual-Dependent Attention Effect on Crowded Orientation Signals in Human Visual Cortex.

A target becomes hard to identify with nearby visual stimuli. This phenomenon, known as crowding, places a fundamental limit on conscious perception and object recognition. To understand the neural representation of crowded stimuli, we used fMRI and a forward encoding model to reconstruct the target-specific feature from multivoxel activation patterns evoked by orientation patches. Orientation-selective response profiles were constructed in V1-V4 for a target embedded in different contexts. Subjects of both sexes either directed their attention over all the orientation patches or selectively to the target. In the context with a weak crowding effect, attending to the target enhanced the orientation selectivity of the response profile; such effect increased along the visual pathway. In the context with a strong crowding effect, attending to the target enhanced the orientation selectivity of the response profile in the earlier visual area, but not in V4. The increase and decrease of orientation selectivity along the visual hierarchy demonstrate a contextual-dependent attention effect on crowded orientation signals: in the context with a weak crowding effect, selective attention gradually resolves the target from nearby distractors along the hierarchy; in the context with a strong crowding effect, while selective attention maintains the target feature in the earlier visual area, its effect decreases in the downstream area. Our findings reveal how the human visual system represents the target-specific feature at multiple stages under the limit of attention selection in a cluttered scene.SIGNIFICANCE STATEMENT Using fMRI and a forward encoding model, we reconstructed orientation-selective response profiles for a target embedded in crowded contexts. In the context with a weak crowding effect, attention gradually resolves the target from nearby distractors along the visual hierarchy. In the context with a strong crowding effect, while the feature of the target is preserved in the early visual cortex, it degrades in the later visual processing stage. The increase and decrease of orientation selectivity along the visual hierarchy reveal how the human visual system strikes to present the target-specific feature under the limit of attention selection in a cluttered scene.

Effect of task difficulty on blood-oxygen-level-dependent signal: A functional magnetic resonance imaging study in a motion discrimination task.

There is much evidence that neural activity in the human brain is modulated by task difficulty, particularly in visual, frontal, and parietal cortices. However, some basic psychophysical tasks in visual perception do not give rise to this expected effect, at least not in the visual cortex. In the current study, we used functional magnetic resonance imaging (fMRI) to record brain activity while systematically manipulating task difficulty in a motion discrimination task, by varying the angular difference between the motion direction of random dots and a reference direction. We used both a blocked and an event-related design, and presented stimuli in both central and peripheral vision. The behavioral psychometric function, across angular differences of 3°, 9°, 15°, or 80°, spanned the full response range, as expected. The mean blood oxygen level dependent (BOLD) signals were also correlated within-participants between the blocked and event-related designs, across all brain areas tested. Within the visual cortex, the voxel response patterns correlated more within-conditions (e.g., 3° and 3°) than between-conditions (e.g., 3° and 9°), in both designs, further attesting to the reasonable quality of the BOLD data. Nevertheless, the BOLD-o-metric functions (i.e., BOLD activity as a function of task difficulty) were flat in the whole-brain and region-of-interest (ROI) analyses, including in the visual cortex, the parietal cortex, in both designs, and in foveal and peripheral visual fields alike. Indeed, there was little difference between BOLD activity during the 3° and 80° conditions. Some suggestive evidence of difficulty modulation was revealed only in the superior and inferior frontal gyri for the blocked design. We conclude that, in motion discrimination, there is no systematic BOLD modulation that accompanies the standard psychometric function across different hierarchies of cortical areas, except for the frontal lobe of the brain.

Hemifield columns co-opt ocular dominance column structure in human achiasma.

In the absence of an optic chiasm, visual input to the right eye is represented in primary visual cortex (V1) in the right hemisphere, while visual input to the left eye activates V1 in the left hemisphere. Retinotopic mapping In V1 reveals that in each hemisphere left and right visual hemifield representations are overlaid (Hoffmann et al., 2012). To explain how overlapping hemifield representations in V1 do not impair vision, we tested the hypothesis that visual projections from nasal and temporal retina create interdigitated left and right visual hemifield representations in V1, similar to the ocular dominance columns observed in neurotypical subjects (Victor et al., 2000). We used high-resolution fMRI at 7T to measure the spatial distribution of responses to left- and right-hemifield stimulation in one achiasmic subject. T2-weighted 2D Spin Echo images were acquired at 0.8mm isotropic resolution. The left eye was occluded. To the right eye, a presentation of flickering checkerboards alternated between the left and right visual fields in a blocked stimulus design. The participant performed a demanding orientation-discrimination task at fixation. A general linear model was used to estimate the preference of voxels in V1 to left- and right-hemifield stimulation. The spatial distribution of voxels with significant preference for each hemifield showed interdigitated clusters which densely packed V1 in the right hemisphere. The spatial distribution of hemifield-preference voxels in the achiasmic subject was stable between two days of testing and comparable in scale to that of human ocular dominance columns. These results are the first in vivo evidence showing that visual hemifield representations interdigitate in achiasmic V1 following a similar developmental course to that of ocular dominance columns in V1 with intact optic chiasm.

Human efficiency in detecting and discriminating biological motion.

Using an "information meter" provided by ideal observer analysis, we measured the efficiency with which human observers processed different walking stimuli against luminance noise and spatial uncertainty to either detect the presence of a walker or to discriminate the walking direction. Human efficiency was examined across four renderings of a human walker: contour, point lights, silhouette, and skeleton. We replicated the previous finding of low discrimination efficiency in biological motion (Gold, Tadin, Cook, & Blake, 2008) and also found low detection efficiency for biological motion. Interestingly, in both detection and discrimination tasks, the skeleton display was among those yielding the highest level of efficiency in processing visual information. This finding suggests that structural information about the relative position of joints, highlighted in the skeleton display, provides a critical component of the internal representation for biological motion.

What Is Actually Affected by the Scrambling of Objects When Localizing the Lateral Occipital Complex?

The lateral occipital complex (LOC), the cortical region critical for shape perception, is localized with fMRI by its greater BOLD activity when viewing intact objects compared with their scrambled versions (resembling texture). Despite hundreds of studies investigating LOC, what the LOC localizer accomplishes-beyond distinguishing shape from texture-has never been resolved. By independently scattering the intact parts of objects, the axis structure defining the relations between parts was no longer defined. This led to a diminished BOLD response, despite the increase in the number of independent entities (the parts) produced by the scattering, thus indicating that LOC specifies interpart relations, in addition to specifying the shape of the parts themselves. LOC's sensitivity to relations is not confined to those between parts but is also readily apparent between objects, rendering it-and not subsequent "place" areas-as the critical region for the representation of scenes. Moreover, that these effects are witnessed with novel as well as familiar intact objects and scenes suggests that the relations are computed on the fly, rather than being retrieved from memory.

Crowding, visual awareness, and their respective neural loci.

In peripheral vision, object identification can be impeded when a target object is flanked by other objects. This phenomenon of crowding has been attributed to basic processes associated with image encoding by the visual system, but the neural origin of crowding is not known. Determining whether crowding depends on subjective awareness of the flankers can provide information on the neural origin of crowding. However, recent studies that manipulated flanker awareness have yielded conflicting results. In the current study, we suppressed flanker awareness with two methods: interocular suppression (IOS) and adaptation-induced blindness (AIB). We tested two different types of stimuli: gratings and letters. With IOS, we found that the magnitude of crowding increased as the number of physical flankers increased, even when the observers did not report seeing any of the flankers. In contrast, when flanker awareness was manipulated with AIB, the magnitude of crowding increased with the number of perceived flankers. Our results show that whether crowding is contingent on awareness of the flankers depends on the method used to suppress awareness. In addition, our results imply that the locus of crowding is upstream from the neural locus of IOS and close to or downstream from that of AIB. Neurophysiology and neuroimaging studies jointly implicate mid-to-high level visual processing stages for IOS, while direct evidence regarding the neural locus of AIB is limited. The most consistent interpretation of our empirical findings is to place the neural locus of crowding at an early cortical site, such as V1 or V2.

Object crowding in age-related macular degeneration.

Crowding, the phenomenon of impeded object identification due to clutter, is believed to be a key limiting factor of form vision in the peripheral visual field. The present study provides a characterization of object crowding in age-related macular degeneration (AMD) measured at the participants' respective preferred retinal loci with binocular viewing. Crowding was also measured in young and age-matched controls at the same retinal locations, using a fixation-contingent display paradigm to allow unlimited stimulus duration. With objects, the critical spacing of crowding for AMD participants was not substantially different from controls. However, baseline contrast energy thresholds in the noncrowded condition were four times that of the controls. Crowding further exacerbated deficits in contrast sensitivity to three times the normal crowding-induced contrast energy threshold elevation. These findings indicate that contrast-sensitivity deficit is a major limiting factor of object recognition for individuals with AMD, in addition to crowding. Focusing on this more tractable deficit of AMD may lead to more effective remediation and technological assistance.

The Lateral Occipital Complex shows no net response to object familiarity.

In 1995, Malach et al. discovered an area whose fMRI BOLD response was greater when viewing intact, familiar objects than when viewing their scrambled versions (resembling texture). Since then hundreds of studies have explored this late visual region termed the Lateral Occipital Complex (LOC), which is now known to be critical for shape perception (James, Culham, Humphrey, Milner, & Goodale, 2003). Malach et al. (1995) discounted a role of familiarity by showing that "abstract" Henry Moore sculptures, unfamiliar to the subjects, also activated this region. This characterization of LOC as a region that responds to shape independently of familiarity has been accepted but never tested with control of the same low-level features. We assessed LOC's response to objects that had identical parts in two different arrangements, one familiar and the other novel. Malach was correct: There is no net effect of familiarity in LOC. However, a multivoxel correlation analysis showed that LOC does distinguish familiar from novel objects.

Automated retinofugal visual pathway reconstruction with multi-shell HARDI and FOD-based analysis.

Diffusion MRI tractography provides a non-invasive modality to examine the human retinofugal projection, which consists of the optic nerves, optic chiasm, optic tracts, the lateral geniculate nuclei (LGN) and the optic radiations. However, the pathway has several anatomic features that make it particularly challenging to study with tractography, including its location near blood vessels and bone-air interface at the base of the cerebrum, crossing fibers at the chiasm, somewhat-tortuous course around the temporal horn via Meyer's Loop, and multiple closely neighboring fiber bundles. To date, these unique complexities of the visual pathway have impeded the development of a robust and automated reconstruction method using tractography. To overcome these challenges, we develop a novel, fully automated system to reconstruct the retinofugal visual pathway from high-resolution diffusion imaging data. Using multi-shell, high angular resolution diffusion imaging (HARDI) data, we reconstruct precise fiber orientation distributions (FODs) with high order spherical harmonics (SPHARM) to resolve fiber crossings, which allows the tractography algorithm to successfully navigate the complicated anatomy surrounding the retinofugal pathway. We also develop automated algorithms for the identification of ROIs used for fiber bundle reconstruction. In particular, we develop a novel approach to extract the LGN region of interest (ROI) based on intrinsic shape analysis of a fiber bundle computed from a seed region at the optic chiasm to a target at the primary visual cortex. By combining automatically identified ROIs and FOD-based tractography, we obtain a fully automated system to compute the main components of the retinofugal pathway, including the optic tract and the optic radiation. We apply our method to the multi-shell HARDI data of 215 subjects from the Human Connectome Project (HCP). Through comparisons with post-mortem dissection measurements, we demonstrate the retinotopic organization of the optic radiation including a successful reconstruction of Meyer's loop. Then, using the reconstructed optic radiation bundle from the HCP cohort, we construct a probabilistic atlas and demonstrate its consistency with a post-mortem atlas. Finally, we generate a shape-based representation of the optic radiation for morphometry analysis.

Feasibility of Structural and Functional MRI Acquisition with Unpowered Implants in Argus II Retinal Prosthesis Patients: A Case Study.

PURPOSE: Magnetic resonance imaging (MRI) can measure the effects of vision loss and recovery on brain function and structure. In this case study, we sought to determine the feasibility of acquiring anatomical and functional MRI data in recipients of the Argus II epiretinal prosthesis system. METHODS: Following successful implantation with the Argus II device, two retinitis pigmentosa (RP) patients completed MRI scans with their implant unpowered to measure primary visual cortex (V1) functional responses to a tactile task, whole-brain morphometry, V1 cortical thickness, and diffusion properties of the optic tract and optic radiation. Measurements in the subjects with the Argus II implant were compared to measurements obtained previously from RP patients and sighted individuals. RESULTS: The presence of the Argus II implant resulted in artifacts that were localized around the patient's implanted eye and did not extend into cortical regions or white matter tracts associated with the visual system. Structural data on V1 cortical thickness and the retinofugal tract obtained from the two Argus II subjects fell within the ranges of sighted and RP groups. When compared to the RP and sighted subjects, Argus II patients' tactile-evoked cross-modal functional MRI (fMRI) blood oxygen level-dependent (BOLD) responses in V1 also fell within the range of either sighted or RP groups, apparently depending on time since implantation. CONCLUSIONS: This study demonstrates that successful acquisition and quantification of structural and functional MR images are feasible in the presence of the inactive implant and provides preliminary information on functional changes in the brain that may follow sight restoration treatments. TRANSITIONAL RELEVANCE: Successful MRI and fMRI acquisition in Argus II recipients demonstrates feasibility of using MRI to study the effect of retinal prosthesis use on brain structure and function.

Using an achiasmic human visual system to quantify the relationship between the fMRI BOLD signal and neural response.

Achiasma in humans causes gross mis-wiring of the retinal-fugal projection, resulting in overlapped cortical representations of left and right visual hemifields. We show that in areas V1-V3 this overlap is due to two co-located but non-interacting populations of neurons, each with a receptive field serving only one hemifield. Importantly, the two populations share the same local vascular control, resulting in a unique organization useful for quantifying the relationship between neural and fMRI BOLD responses without direct measurement of neural activity. Specifically, we can non-invasively double local neural responses by stimulating both neuronal populations with identical stimuli presented symmetrically across the vertical meridian to both visual hemifields, versus one population by stimulating in one hemifield. Measurements from a series of such doubling experiments show that the amplitude of BOLD response is proportional to approximately 0.5 power of the underlying neural response. Reanalyzing published data shows that this inferred relationship is general.

Visual speech discrimination and identification of natural and synthetic consonant stimuli.

From phonetic features to connected discourse, every level of psycholinguistic structure including prosody can be perceived through viewing the talking face. Yet a longstanding notion in the literature is that visual speech perceptual categories comprise groups of phonemes (referred to as visemes), such as /p, b, m/ and /f, v/, whose internal structure is not informative to the visual speech perceiver. This conclusion has not to our knowledge been evaluated using a psychophysical discrimination paradigm. We hypothesized that perceivers can discriminate the phonemes within typical viseme groups, and that discrimination measured with d-prime (d') and response latency is related to visual stimulus dissimilarities between consonant segments. In Experiment 1, participants performed speeded discrimination for pairs of consonant-vowel spoken nonsense syllables that were predicted to be same, near, or far in their perceptual distances, and that were presented as natural or synthesized video. Near pairs were within-viseme consonants. Natural within-viseme stimulus pairs were discriminated significantly above chance (except for /k/-/h/). Sensitivity (d') increased and response times decreased with distance. Discrimination and identification were superior with natural stimuli, which comprised more phonetic information. We suggest that the notion of the viseme as a unitary perceptual category is incorrect. Experiment 2 probed the perceptual basis for visual speech discrimination by inverting the stimuli. Overall reductions in d' with inverted stimuli but a persistent pattern of larger d' for far than for near stimulus pairs are interpreted as evidence that visual speech is represented by both its motion and configural attributes. The methods and results of this investigation open up avenues for understanding the neural and perceptual bases for visual and audiovisual speech perception and for development of practical applications such as visual lipreading/speechreading speech synthesis.

Correlation of vision loss with tactile-evoked V1 responses in retinitis pigmentosa.

Neuroimaging studies have shown that the visual cortex of visually impaired humans is active during tactile tasks. We sought to determine if this cross-modal activation in the primary visual cortex is correlated with vision loss in individuals with retinitis pigmentosa (RP), an inherited degenerative photoreceptor disease that progressively diminishes vision later in life. RP and sighted subjects completed three tactile tasks: a symmetry discrimination task, a Braille-dot counting task, and a sandpaper roughness discrimination task. We measured tactile-evoked blood oxygenation level dependent (BOLD) responses using functional magnetic resonance imaging (fMRI). For each subject, we quantified the cortical extent of the tactile-evoked response by the proportion of modulated voxels within the primary visual cortex (V1) and its strength by the mean absolute modulation amplitude of the modulated voxels. We characterized vision loss in terms of visual acuity and the areal proportion of V1 that corresponds to the preserved visual field. Visual acuity and proportion of the preserved visual field both had a highly significant effect on the cortical extent of the V1 BOLD response to tactile stimulation, while visual acuity also had a significant effect on the strength of the V1 response. These effects of vision loss on cross-modal responses were reliable despite high inter-subject variability. Controlling for task-evoked responses in the primary somatosensory cortex (S1) across subjects further strengthened the effects of vision loss on cross-model responses in V1. We propose that such cross-modal responses in V1 and other visual areas may be used as a cortically localized biomarker to account for individual differences in visual performance following sight recovery treatments.

Radial-tangential anisotropy of crowding in the early visual areas.

Crowding, the inability to recognize an individual object in clutter (Bouma H. Nature 226: 177-178, 1970), is considered a major impediment to object recognition in peripheral vision. Despite its significance, the cortical loci of crowding are not well understood. In particular, the role of the primary visual cortex (V1) remains unclear. Here we utilize a diagnostic feature of crowding to identify the earliest cortical locus of crowding. Controlling for other factors, radially arranged flankers induce more crowding than tangentially arranged ones (Toet A, Levi DM. Vision Res 32: 1349-1357, 1992). We used functional magnetic resonance imaging (fMRI) to measure the change in mean blood oxygenation level-dependent (BOLD) response due to the addition of a middle letter between a pair of radially or tangentially arranged flankers. Consistent with the previous finding that crowding is associated with a reduced BOLD response [Millin R, Arman AC, Chung ST, Tjan BS. Cereb Cortex (July 5, 2013). doi:10.1093/cercor/bht159], we found that the BOLD signal evoked by the middle letter depended on the arrangement of the flankers: less BOLD response was associated with adding the middle letter between radially arranged flankers compared with adding it between tangentially arranged flankers. This anisotropy in BOLD response was present as early as V1 and remained significant in downstream areas. The effect was observed while subjects' attention was diverted away from the testing stimuli. Contrast detection threshold for the middle letter was unaffected by flanker arrangement, ruling out surround suppression of contrast response as a major factor in the observed BOLD anisotropy. Our findings support the view that V1 contributes to crowding.

A visual or tactile signal makes auditory speech detection more efficient by reducing uncertainty.

Acoustic speech is easier to detect in noise when the talker can be seen. This finding could be explained by integration of multisensory inputs or refinement of auditory processing from visual guidance. In two experiments, we studied two-interval forced-choice detection of an auditory 'ba' in acoustic noise, paired with various visual and tactile stimuli that were identically presented in the two observation intervals. Detection thresholds were reduced under the multisensory conditions vs. the auditory-only condition, even though the visual and/or tactile stimuli alone could not inform the correct response. Results were analysed relative to an ideal observer for which intrinsic (internal) noise and efficiency were independent contributors to detection sensitivity. Across experiments, intrinsic noise was unaffected by the multisensory stimuli, arguing against the merging (integrating) of multisensory inputs into a unitary speech signal, but sampling efficiency was increased to varying degrees, supporting refinement of knowledge about the auditory stimulus. The steepness of the psychometric functions decreased with increasing sampling efficiency, suggesting that the 'task-irrelevant' visual and tactile stimuli reduced uncertainty about the acoustic signal. Visible speech was not superior for enhancing auditory speech detection. Our results reject multisensory neuronal integration and speech-specific neural processing as explanations for the enhanced auditory speech detection under noisy conditions. Instead, they support a more rudimentary form of multisensory interaction: the otherwise task-irrelevant sensory systems inform the auditory system about when to listen.

Visual crowding in V1.

In peripheral vision, objects in clutter are difficult to identify. The exact cause of this "crowding" effect is unclear. To perceive coherent shapes in clutter, the visual system must integrate certain local features across receptive fields while preventing others from being combined. It is believed that this selective feature integration-segmentation process is impaired in peripheral vision, leading to crowding. We used functional magnetic resonance imaging (fMRI) to investigate the neural origin of crowding. We found that crowding was associated with suppressed fMRI signal as early as V1, regardless of whether attention was directed toward or away from a target stimulus. This suppression in early visual cortex was greatest for stimuli that produced the strongest crowding. In contrast, the pattern of activity was mixed in higher level visual areas, such as the lateral occipital cortex. These results support the view that the deficiency in feature integration and segmentation in peripheral vision is present at the earliest stages of cortical processing.

Indoor navigation by people with visual impairment using a digital sign system.

There is a need for adaptive technology to enhance indoor wayfinding by visually-impaired people. To address this need, we have developed and tested a Digital Sign System. The hardware and software consist of digitally-encoded signs widely distributed throughout a building, a handheld sign-reader based on an infrared camera, image-processing software, and a talking digital map running on a mobile device. Four groups of subjects-blind, low vision, blindfolded sighted, and normally sighted controls-were evaluated on three navigation tasks. The results demonstrate that the technology can be used reliably in retrieving information from the signs during active mobility, in finding nearby points of interest, and following routes in a building from a starting location to a destination. The visually impaired subjects accurately and independently completed the navigation tasks, but took substantially longer than normally sighted controls. This fully functional prototype system demonstrates the feasibility of technology enabling independent indoor navigation by people with visual impairment.

Rapid and persistent adaptability of human oculomotor control in response to simulated central vision loss.

The central region of the human retina, the fovea, provides high-acuity vision. The oculomotor system continually brings targets of interest into the fovea via ballistic eye movements (saccades). Thus, the fovea serves both as the locus for fixations and as the oculomotor reference for saccades. This highly automated process of foveation is functionally critical to vision and is observed from infancy. How would the oculomotor system adjust to a loss of foveal vision (central scotoma)? Clinical observations of patients with central vision loss suggest a lengthy adjustment period, but the nature and dynamics of this adjustment remain unclear. Here, we demonstrate that the oculomotor system can spontaneously and rapidly adopt a peripheral locus for fixation and can rereference saccades to this locus in normally sighted individuals whose central vision is blocked by an artificial scotoma. Once developed, the fixation locus is retained over weeks in the absence of the simulated scotoma. Our data reveal a basic guiding principle of the oculomotor system that prefers control simplicity over optimality. We demonstrate the importance of a visible scotoma on the speed of the adjustment and suggest a possible rehabilitation regimen for patients with central vision loss.

Crowding during restricted and free viewing.

Crowding impairs the perception of form in peripheral vision. It is likely to be a key limiting factor of form vision in patients without central vision. Crowding has been extensively studied in normally sighted individuals, typically with a stimulus duration of a few hundred milliseconds to avoid eye movements. These restricted testing conditions do not reflect the natural behavior of a patient with central field loss. Could unlimited stimulus duration and unrestricted eye movements change the properties of crowding in any fundamental way? We studied letter identification in the peripheral vision of normally sighted observers in three conditions: (i) a fixation condition with a brief stimulus presentation of 250 ms, (ii) another fixation condition but with an unlimited viewing time, and (iii) an unrestricted eye movement condition with an artificial central scotoma and an unlimited viewing time. In all conditions, contrast thresholds were measured as a function of target-to-flanker spacing, from which we estimated the spatial extent of crowding in terms of critical spacing. We found that presentation duration beyond 250 ms had little effect on critical spacing with stable gaze. With unrestricted eye movements and a simulated central scotoma, we found a large variability in critical spacing across observers, but more importantly, the variability in critical spacing was well correlated with the variability in target eccentricity. Our results assure that the large body of findings on crowding made with briefly presented stimuli remains relevant to conditions where viewing time is unconstrained. Our results further suggest that impaired oculomotor control associated with central vision loss can confound peripheral form vision beyond the limits imposed by crowding.