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• W2022061511 abstract "The two eyes of an individual routinely differ in their optical and neural properties, yet percepts through either eye remain more similar than predicted by these differences. Little is known as to how the brain resolves this conflicting information. Differences in visual inputs from the two eyes have been studied extensively in the context of binocular vision and rivalry [1Blake R. Wilson H. Binocular vision.Vis. Res. 2011; 51: 754-770Crossref PubMed Scopus (190) Google Scholar], but it remains unknown how the visual system calibrates and corrects for normal variability in image quality between the eyes, and whether this correction is applied to each eye separately or after their signals have converged. To test this, we used adaptive optics to control and manipulate the blur projected on each retina, and then compared judgments of image focus through either eye and how these judgments were biased by adapting to different levels of blur. Despite significant interocular differences in the magnitude of optical blur, the blur level that appeared best focused was the same through both eyes, and corresponded to the ocular blur of the less aberrated eye. Moreover, for both eyes, blur aftereffects depended on whether the adapting blur was stronger or weaker than the native blur of the better eye, with no aftereffect when the blur equaled the aberrations of the better eye. Our results indicate that the neural calibration for the perception of image focus reflects a single ‘cyclopean’ site that is set monocularly by the eye with better optical quality. Consequently, what people regard as ‘best-focused’ matches the blur encountered through the eye with better optics, even when judging the world through the eye with poorer optics. The two eyes of an individual routinely differ in their optical and neural properties, yet percepts through either eye remain more similar than predicted by these differences. Little is known as to how the brain resolves this conflicting information. Differences in visual inputs from the two eyes have been studied extensively in the context of binocular vision and rivalry [1Blake R. Wilson H. Binocular vision.Vis. Res. 2011; 51: 754-770Crossref PubMed Scopus (190) Google Scholar], but it remains unknown how the visual system calibrates and corrects for normal variability in image quality between the eyes, and whether this correction is applied to each eye separately or after their signals have converged. To test this, we used adaptive optics to control and manipulate the blur projected on each retina, and then compared judgments of image focus through either eye and how these judgments were biased by adapting to different levels of blur. Despite significant interocular differences in the magnitude of optical blur, the blur level that appeared best focused was the same through both eyes, and corresponded to the ocular blur of the less aberrated eye. Moreover, for both eyes, blur aftereffects depended on whether the adapting blur was stronger or weaker than the native blur of the better eye, with no aftereffect when the blur equaled the aberrations of the better eye. Our results indicate that the neural calibration for the perception of image focus reflects a single ‘cyclopean’ site that is set monocularly by the eye with better optical quality. Consequently, what people regard as ‘best-focused’ matches the blur encountered through the eye with better optics, even when judging the world through the eye with poorer optics. In Experiment 1, we used an adaptive optics system [2Marcos S. Sawides L. Gambra E. Dorronsoro C. Influence of adaptive-optics ocular aberration correction on visual acuity at different luminances and contrast polarities.J. Vis. 2008; 8: 1-12Crossref Scopus (69) Google Scholar] to completely correct for the blur within each eye and then present varying amounts of blur (described as the ratio of the peak aberrated image intensity from a point source compared to the maximum attainable intensity using an ideal optical system limited only by diffraction over the system’s aperture) corresponding to defects measured from real observers (see supplementary material). The magnitude of retinal image blur varies substantially both across observers and between the two eyes of the same observer, showing only a weak correlation between the two eyes (r = 0.441, p = 0.052; Figure 1D). The perceived-best-focus — the blur level that appears neither too sharp nor too blurred — also varied across subjects, but it was nearly identical for a given subject regardless of whether the judgment was made with the right or left eye (r = 0.984, p < 0.001; Figure 1E). These judgments corresponded closely to the individual’s native blur, and in subjects with significant (> 30%) differences between their eyes, did not differ from the blur level dictated by the better eye quality (–0.03 ± 0.05; p = ns), but were substantially sharper than predicted by the worse eye (0.097 ± 0.074; t(6) = 3.47, p = 0.013). These results are consistent with previous reports [3Artal P. Chen L. Fernandez E.J. Singer B. Manzanera S. Williams D.R. Neural compensation for the eye’s optical aberrations.J. Vis. 2004; 4: 281-287Crossref PubMed Google Scholar, 4Sawides L. de Gracia P. Dorronsoro C. Webster M.A. Marcos S. Vision is adapted to the natural level of blur present in the retinal image.PloS One. 2011; 6: e27031Crossref PubMed Scopus (59) Google Scholar] that observers perceive as best-focused the image blur that they are chronically exposed to, but reveal for the first time that this calibration is the same through either eye and determined by the eye with better optics. Judgments of image focus could reflect a learned criterion (for example, our own blur is what we are used to seeing) or how sensitivity to blur is calibrated (for example, in neural contrast sensitivity). To test these alternatives, in Experiment 2 we measured changes in perceived focus after brief adaptation to blurred or sharp images, and probed which blur level did not produce an aftereffect, again testing each eye independently. Adapting to blur causes a subsequent test image to appear too sharp, while over-sharpened adaptors instead make images appear blurrier [5Webster M.A. Georgeson M.A. Webster S.M. Neural adjustments to image blur.Nat. Neurosci. 2002; 5: 839-840Crossref PubMed Scopus (205) Google Scholar]. By titrating the level of adapting blur, the level that does not alter the blur percepts can be determined, and reveals the stimulus that neural sensitivity is calibrated for [6Webster M.A. Leonard D. Adaptation and perceptual norms in color vision.J. Opt. Soc. Am. A. 2008; 25: 2817-2825Crossref Scopus (69) Google Scholar]. Accordingly, we chose adapting levels to bracket and include the magnitude of blur within each eye, again using adaptive optics to bypass the eye’s optics while projecting the adapting and test images on the retina. Despite large differences between subjects in subjective focus (which varied from 0.094 to 0.412 SR), for each the pattern of aftereffects was again strikingly similar between their eyes (Figure 1F), with an interocular difference in SR of only 0.002 ± 0.002 (and no interocular difference in the magnitude of aftereffects; F = 1.07; and df = 24; p = 0.819). Moreover, for either eye, the blur level at which the aftereffect was nulled again corresponded closely to the better eye, while exposure to the worse eye’s blur or to the average blur of the two eyes caused the previous subjective focus level to appear too sharp. Thus, both the focus judgments and how they were biased by the adaptation were completely determined by the better eye, consistent with a neural calibration matched to the optical quality of the eye with least optical defects. It is well known that in binocular viewing one eye is typically dominant [7Arnold D.H. Grove P.M. Wallis T.S. Staying focused: a functional account of perceptual suppression during binocular rivalry.J. Vis. 2007; 7: 1-8Crossref Scopus (38) Google Scholar], and previous work has shown that a sharper image presented to one eye dominates a blurrier image in the other (for example [7Arnold D.H. Grove P.M. Wallis T.S. Staying focused: a functional account of perceptual suppression during binocular rivalry.J. Vis. 2007; 7: 1-8Crossref Scopus (38) Google Scholar, 8Georgeson M.A. May K.A. Freeman T.C. Hesse G.S. From filters to features: scale-space analysis of edge and blur coding in human vision.J. Vis. 2007; 7: 1-21Crossref PubMed Scopus (59) Google Scholar]). However, our findings are novel and important in showing that this sensory dominance persists to influence perceived focus — a fundamental perceptual judgment — even when the eyes are stimulated separately. These results establish that there is a single ‘cyclopean’ locus of the neural compensation for the eye’s optical defects, calibrating the neural signals carried by either eye but set only by the better eye, and that the perception of focus corresponds to a unique null point in the sensitivity of the underlying neural code. This correspondence also reveals a close correspondence between subjectively neutral percepts (what ‘looks’ focused) and neutral states in the neural code (what stimulus neural sensitivity is adapted to), a link that has rarely been documented but which may reflect a general basis for perceptual norms [9Webster M.A. Adaptation and visual coding.J. Vis. 2011; 11: 1-23Crossref Google Scholar]. For example, the stimulus that appears a neutral gray similarly reflects a sensitivity null in color mechanisms, but one established in the retina [6Webster M.A. Leonard D. Adaptation and perceptual norms in color vision.J. Opt. Soc. Am. A. 2008; 25: 2817-2825Crossref Scopus (69) Google Scholar]. The nature of these visual calibrations is also clinically important for understanding the consequences of interocular differences in optical errors as well as refractive corrections such as monovision which intentionally introduce these differences by focusing one eye for far vision and the other for near [10Schor C. Landsman L. Erickson P. Ocular dominance and the interocular suppression of blur in monovision.Am. J. Optom. Physiol. Opt. 1987; 64: 723-730Crossref PubMed Scopus (124) Google Scholar]. Supported by the European Research Council under the European Union’s Seventh Framework Programme (FP7/2007-2013) / ERC Grant Agreement n° 294099, Spanish Government grant FIS2011-264605 and 7th Framework Programme of the European Community through the Marie Curie Initial Training Network OpAL (OpAL is an Initial Training Network funded by the European Commission under the Seventh Framework Programme (PITN-GA-2010-264605)), and National Eye Institute grant EY-10834. Download .pdf (.32 MB) Help with pdf files Document S1. Experimental procedures, Results and Two Figures" @default.
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• W2022061511 title "A cyclopean neural mechanism compensating for optical differences between the eyes" @default.
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