{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T18:49:48Z","timestamp":1792090188498,"version":"4.4.0"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1014856","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000}}],"reference-count":133,"publisher":"Public Library of Science (PLoS)","issue":"10","license":[{"start":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T00:00:00Z","timestamp":1791763200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"CIHR Vanier Scholarship"},{"name":"NSERC Discovery Grant","award":["RGPIN-2022-04216"],"award-info":[{"award-number":["RGPIN-2022-04216"]}]},{"DOI":"10.13039\/501100017657","name":"Fondation de l'H\u00f4pital G\u00e9n\u00e9ral de Montr\u00e9al","doi-asserted-by":"publisher","award":["Center for Digital Therapies Grant"],"award-info":[{"award-number":["Center for Digital Therapies Grant"]}],"id":[{"id":"10.13039\/501100017657","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>\n                    In the geniculostriate pathway of the human visual system, neuronal projections carry signals from a particular retinal locus in parallel from one anatomical area to the next. Imprecision in the fidelity of these projections would place constraints on the ability of the system to perform tasks requiring positional information. We investigated the impact that \u201cspatial scrambling\u201d between stages would have on visual performance. We consider two stages in a simple canonical model of the early visual cortex where scrambling might occur: either the input to the first orientation-tuned mechanisms (analogous to V1 simple cells), or the output from those mechanisms. These are referred as \u201csubcortical\u201d (SCS) and \u201ccortical scrambling\u201d (CS). We developed a wavelet decomposition and resynthesis algorithm to mimic these effects, and measured human performance in letter identification affected by the two types of scrambling. Our results showed SCS and CS have distinguishable effects on both perceived noisiness of letters and letter identification threshold. Comparing human performance against a suite of pre-trained and custom convolutional neural networks (CNNs) that were trained on the scrambled stimuli, relative efficiency (calculated from the ratio of human:CNN thresholds) is higher for CS than SCS. However, in modelling human inefficiency by reducing the proportion of wavelets available to the CNNs, humans are\n                    <jats:italic>less<\/jats:italic>\n                    efficient in CS than SCS. These differences in efficiencies show humans are better at processing orientation redundant stimuli (CS) than orientation noisy stimuli (SCS). We hypothesize this reflects differences in integration properties at the input and output stages of simple cells in the cortex.\n                  <\/jats:p>","DOI":"10.1371\/journal.pcbi.1014856","type":"journal-article","created":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T17:36:51Z","timestamp":1791826611000},"page":"e1014856","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Identification of letters distorted by physiologically-inspired spatial scrambling"],"prefix":"10.1371","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-1930-8912","authenticated-orcid":true,"given":"Raffles Xingqi","family":"Zhu","sequence":"first","affiliation":[{"name":"McGill Vision Research","place":["Montreal, Quebec, Canada"],"department":["Department of Ophthalmology & Visual Sciences"]},{"name":"Research Institute of the McGill University Health Centre","place":["Montreal, Quebec, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3830-6254","authenticated-orcid":true,"given":"Alex S.","family":"Baldwin","sequence":"additional","affiliation":[{"name":"McGill Vision Research","place":["Montreal, Quebec, Canada"],"department":["Department of Ophthalmology & Visual Sciences"]},{"name":"Research Institute of the McGill University Health Centre","place":["Montreal, Quebec, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,10,12]]},"reference":[{"issue":"3","key":"pcbi.1014856.ref001","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1016\/j.neuron.2012.01.010","article-title":"How does the brain solve visual object recognition?","volume":"73","author":"JJ DiCarlo","year":"2012","journal-title":"Neuron"},{"issue":"1","key":"pcbi.1014856.ref002","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1113\/jphysiol.1962.sp006837","article-title":"Receptive fields, binocular interaction and functional architecture in the cat\u2019s visual cortex","volume":"160","author":"DH Hubel","year":"1962","journal-title":"The Journal of Physiology"},{"issue":"11","key":"pcbi.1014856.ref003","doi-asserted-by":"crossref","first-page":"1019","DOI":"10.1038\/14819","article-title":"Hierarchical models of object recognition in cortex","volume":"2","author":"M Riesenhuber","year":"1999","journal-title":"Nat Neurosci"},{"issue":"3","key":"pcbi.1014856.ref004","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1109\/TIT.1962.1057716","article-title":"Information capacity of a single retinal channel","volume":"8","author":"D Kelly","year":"1962","journal-title":"IEEE Trans Inform Theory"},{"issue":"6838","key":"pcbi.1014856.ref005","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1038\/35079612","article-title":"Retinal ganglion cells act largely as independent encoders","volume":"411","author":"S Nirenberg","year":"2001","journal-title":"Nature"},{"key":"pcbi.1014856.ref006","volume-title":"Understanding vision: theory, models, and data","author":"Z Li","year":"2014"},{"issue":"1","key":"pcbi.1014856.ref007","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/S1364-6613(00)01817-9","article-title":"A saliency map in primary visual cortex","volume":"6","author":"Z Li","year":"2002","journal-title":"Trends Cogn Sci"},{"issue":"3","key":"pcbi.1014856.ref008","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1017\/S0140525X12000477","article-title":"Whatever next? 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Phantom Forms in Amblyopic Vision and What They Reveal About the Generative Brain. 2025;:2025.06.24.661078.","DOI":"10.1101\/2025.06.24.661078"},{"issue":"15","key":"pcbi.1014856.ref049","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1167\/iovs.66.15.65","article-title":"Letter distortion mapping in amblyopia: spatial patterns, stability, and relationship to visual acuity","volume":"66","author":"H Molaei","year":"2025","journal-title":"Investigative Ophthalmology & Visual Science"},{"issue":"45","key":"pcbi.1014856.ref050","doi-asserted-by":"crossref","DOI":"10.1523\/JNEUROSCI.1111-25.2025","article-title":"Visual Distortions in Human Amblyopia Are Correlated with Deficits in Contrast Sensitivity","volume":"45","author":"F Olianezhad","year":"2025","journal-title":"J Neurosci"},{"key":"pcbi.1014856.ref051","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1016\/0042-6989(93)90188-3","article-title":"Distortions in two-dimensional visual space perception in strabismic 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Comparison of classifier methods: a case study in handwritten digit recognition. In: 1994. 77\u201382.","DOI":"10.1109\/ICPR.1994.576879"},{"key":"pcbi.1014856.ref071","article-title":"Character-level Convolutional Networks for Text Classification","volume-title":"Advances in Neural Information Processing Systems","author":"X Zhang","year":"2015"},{"issue":"10","key":"pcbi.1014856.ref072","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1016\/j.tics.2008.06.006","article-title":"Letter perception: from pixels to pandemonium","volume":"12","author":"J Grainger","year":"2008","journal-title":"Trends Cogn Sci"},{"issue":"9","key":"pcbi.1014856.ref073","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1016\/S0042-6989(02)00045-7","article-title":"The role of spatial frequency channels in letter identification","volume":"42","author":"NJ Majaj","year":"2002","journal-title":"Vision Res"},{"issue":"6479","key":"pcbi.1014856.ref074","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/369395a0","article-title":"The visual filter mediating letter identification","volume":"369","author":"JA Solomon","year":"1994","journal-title":"Nature"},{"issue":"1","key":"pcbi.1014856.ref075","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1017\/S0952523800004314","article-title":"Role of inhibition in the specification of orientation selectivity of cells in the cat striate cortex","volume":"2","author":"AB Bonds","year":"1989","journal-title":"Vis Neurosci"},{"issue":"5","key":"pcbi.1014856.ref076","doi-asserted-by":"crossref","first-page":"2108","DOI":"10.1152\/jn.00540.2011","article-title":"Cross-orientation suppression in human visual cortex","volume":"106","author":"GJ Brouwer","year":"2011","journal-title":"J Neurophysiol"},{"issue":"9","key":"pcbi.1014856.ref077","doi-asserted-by":"crossref","first-page":"2406","DOI":"10.1364\/JOSAA.14.002406","article-title":"Visual signal detection in structured backgrounds. II. Effects of contrast gain control, background variations, and white noise","volume":"14","author":"MP Eckstein","year":"1997","journal-title":"J Opt Soc Am A Opt Image Sci Vis"},{"issue":"6","key":"pcbi.1014856.ref078","doi-asserted-by":"crossref","first-page":"1710","DOI":"10.1364\/JOSAA.11.001710","article-title":"Human luminance pattern-vision mechanisms: masking experiments require a new model","volume":"11","author":"JM Foley","year":"1994","journal-title":"J Opt Soc Am A Opt Image Sci Vis"},{"issue":"2","key":"pcbi.1014856.ref079","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1017\/S0952523800009640","article-title":"Normalization of cell responses in cat striate cortex","volume":"9","author":"DJ Heeger","year":"1992","journal-title":"Vis Neurosci"},{"issue":"1606","key":"pcbi.1014856.ref080","first-page":"127","article-title":"Spatial and temporal dependencies of cross-orientation suppression in human vision","volume":"274","author":"TS Meese","year":"2007","journal-title":"Proc Biol Sci"},{"issue":"3","key":"pcbi.1014856.ref081","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1016\/j.tins.2022.12.008","article-title":"Using artificial neural networks to ask \u201cwhy\u201d questions of minds and brains","volume":"46","author":"N Kanwisher","year":"2023","journal-title":"Trends Neurosci"},{"issue":"10","key":"pcbi.1014856.ref082","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.1162\/jocn_a_01544","article-title":"Convolutional Neural Networks as a Model of the Visual System: Past, Present, and Future","volume":"33","author":"GW Lindsay","year":"2021","journal-title":"J Cogn Neurosci"},{"issue":"13","key":"pcbi.1014856.ref083","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1167\/18.13.2","article-title":"Deep learning-Using machine learning to study biological vision","volume":"18","author":"NJ Majaj","year":"2018","journal-title":"J Vis"},{"issue":"9","key":"pcbi.1014856.ref084","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1146\/annurev-vision-120522-031739","article-title":"Are deep neural networks adequate behavioral models of human visual perception?","volume":"9","author":"FA Wichmann","year":"2023","journal-title":"Annual Review of Vision Science"},{"issue":"3","key":"pcbi.1014856.ref085","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1038\/nn.4244","article-title":"Using goal-driven deep learning models to understand sensory cortex","volume":"19","author":"DLK Yamins","year":"2016","journal-title":"Nat Neurosci"},{"issue":"1","key":"pcbi.1014856.ref086","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1038\/s41562-021-01244-z","article-title":"Deep neural network models of sound localization reveal how perception is adapted to real-world environments","volume":"6","author":"A Francl","year":"2022","journal-title":"Nat Hum Behav"},{"key":"pcbi.1014856.ref087","doi-asserted-by":"crossref","unstructured":"He K, Zhang X, Ren S, Sun J. Deep Residual Learning for Image Recognition. In: 2015. https:\/\/doi.org\/arXiv:1512.03385","DOI":"10.1109\/CVPR.2016.90"},{"key":"pcbi.1014856.ref088","article-title":"ImageNet Classification with Deep Convolutional Neural Networks","volume-title":"Advances in Neural Information Processing Systems","author":"A Krizhevsky","year":"2012"},{"key":"pcbi.1014856.ref089","unstructured":"Kubilius J, Schrimpf M, Kar K, Rajalingham R, Hong H, Majaj N, et al. Brain-Like Object Recognition with High-Performing Shallow Recurrent ANNs. In: Advances in Neural Information Processing Systems, 2019."},{"key":"pcbi.1014856.ref090","unstructured":"Simonyan K, Zisserman A. Very Deep Convolutional Networks for Large-Scale Image Recognition. 2015. https:\/\/arxiv.org\/abs\/1409.1556"},{"issue":"10","key":"pcbi.1014856.ref091","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1121\/1.1909408","article-title":"Definitions of d\u2032 and \u03b7 as Psychophysical Measures","volume":"30","author":"Jr Tanner WP","year":"1958","journal-title":"The Journal of the Acoustical Society of America"},{"issue":"2","key":"pcbi.1014856.ref092","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1037\/0033-295X.96.2.267","article-title":"Sequential ideal-observer analysis of visual discriminations","volume":"96","author":"WS Geisler","year":"1989","journal-title":"Psychol Rev"},{"key":"pcbi.1014856.ref093","volume-title":"Detection, estimation, and modulation theory, part i: Detection, estimation, and linear modulation theory","author":"HL Van Trees","year":"1968"},{"issue":"10","key":"pcbi.1014856.ref094","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1167\/14.10.1311","article-title":"The bottleneck in human letter recognition: A computational model","volume":"14","author":"A Ziskind","year":"2014","journal-title":"Journal of Vision"},{"key":"pcbi.1014856.ref095","article-title":"Generalisation in humans and deep neural networks","volume-title":"Advances in Neural Information Processing Systems","author":"R Geirhos","year":"2018"},{"issue":"2","key":"pcbi.1014856.ref096","doi-asserted-by":"crossref","DOI":"10.1214\/aos\/1176344136","article-title":"Estimating the Dimension of a Model","volume":"6","author":"G Schwarz","year":"1978","journal-title":"Ann Statist"},{"key":"pcbi.1014856.ref097","first-page":"199","article-title":"Information Theory and an Extension of the Maximum Likelihood Principle","volume-title":"Springer Series in Statistics","author":"H Akaike","year":"1998"},{"key":"pcbi.1014856.ref098","unstructured":"Berardino A, Laparra V, Ball\u00e9 J, Simoncelli E. Eigen-Distortions of Hierarchical Representations. In: Advances in Neural Information Processing Systems, 2017."},{"issue":"11","key":"pcbi.1014856.ref099","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.1038\/s41593-023-01442-0","article-title":"Model metamers reveal divergent invariances between biological and artificial neural networks","volume":"26","author":"J Feather","year":"2023","journal-title":"Nat Neurosci"},{"key":"pcbi.1014856.ref100","doi-asserted-by":"crossref","unstructured":"Borji A, Itti L. Human vs. Computer in Scene and Object Recognition. In: 2014. 113\u201320.","DOI":"10.1109\/CVPR.2014.22"},{"key":"pcbi.1014856.ref101","doi-asserted-by":"crossref","unstructured":"Dodge S, Karam L. A Study and Comparison of Human and Deep Learning Recognition Performance under Visual Distortions. In: 2017 26th International Conference on Computer Communication and Networks (ICCCN), 2017. 1\u20137. https:\/\/doi.org\/10.1109\/icccn.2017.8038465","DOI":"10.1109\/ICCCN.2017.8038465"},{"key":"pcbi.1014856.ref102","unstructured":"Geirhos R, Meding K, Wichmann FA. Beyond accuracy: quantifying trial-by-trial behaviour of CNNs and humans by measuring error consistency. arXiv. 2020. doi: 10.48550\/arXiv.2006.16736"},{"issue":"33","key":"pcbi.1014856.ref103","doi-asserted-by":"crossref","first-page":"7255","DOI":"10.1523\/JNEUROSCI.0388-18.2018","article-title":"Large-Scale, High-Resolution Comparison of the Core Visual Object Recognition Behavior of Humans, Monkeys, and State-of-the-Art Deep Artificial Neural Networks","volume":"38","author":"R Rajalingham","year":"2018","journal-title":"J Neurosci"},{"issue":"10","key":"pcbi.1014856.ref104","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1167\/12.10.20","article-title":"Zero-dimensional noise: The best mask you never saw","volume":"12","author":"DH Baker","year":"2012","journal-title":"Journal of Vision"},{"issue":"3","key":"pcbi.1014856.ref105","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0150942","article-title":"What Do Contrast Threshold Equivalent Noise Studies Actually Measure? Noise vs. Nonlinearity in Different Masking Paradigms","volume":"11","author":"AS Baldwin","year":"2016","journal-title":"PLoS One"},{"issue":"9","key":"pcbi.1014856.ref106","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1167\/15.9.24","article-title":"Prior implicit knowledge shapes human threshold for orientation noise","volume":"15","author":"JH Christensen","year":"2015","journal-title":"J Vis"},{"issue":"11","key":"pcbi.1014856.ref107","doi-asserted-by":"crossref","DOI":"10.1167\/8.11.9","article-title":"A \u201cdipper\u201d function for texture discrimination based on orientation variance","volume":"8","author":"M Morgan","year":"2008","journal-title":"J Vis"},{"issue":"23","key":"pcbi.1014856.ref108","doi-asserted-by":"crossref","first-page":"3311","DOI":"10.1016\/S0042-6989(97)00169-7","article-title":"Sparse coding with an overcomplete basis set: a strategy employed by V1?","volume":"37","author":"BA Olshausen","year":"1997","journal-title":"Vision Res"},{"issue":"2","key":"pcbi.1014856.ref109","doi-asserted-by":"crossref","DOI":"10.1167\/7.2.5","article-title":"The nature of letter crowding as revealed by first- and second-order classification images","volume":"7","author":"AS Nandy","year":"2007","journal-title":"J Vis"},{"key":"pcbi.1014856.ref110","doi-asserted-by":"crossref","first-page":"108588","DOI":"10.1016\/j.visres.2025.108588","article-title":"The impact of eye movements on amblyopic vision: A mini-review","volume":"230","author":"DM Levi","year":"2025","journal-title":"Vision Res"},{"key":"pcbi.1014856.ref111","doi-asserted-by":"crossref","first-page":"108819","DOI":"10.1016\/j.visres.2026.108819","article-title":"The amblyopic acuity deficit: Identification of letters distorted by spatial scrambling algorithms","volume":"244","author":"RX Zhu","year":"2026","journal-title":"Vision Res"},{"issue":"9","key":"pcbi.1014856.ref112","doi-asserted-by":"crossref","first-page":"5822","DOI":"10.1167\/jov.23.9.5822","article-title":"Plenoptic: A platform for synthesizing model-optimized visual stimuli","volume":"23","author":"L Duong","year":"2023","journal-title":"Journal of Vision"},{"key":"pcbi.1014856.ref113","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.jneumeth.2006.11.017","article-title":"PsychoPy--Psychophysics software in Python","volume":"162","author":"JW Peirce","year":"2007","journal-title":"J Neurosci Methods"},{"key":"pcbi.1014856.ref114","unstructured":"Paszke A, Gross S, Massa F, Lerer A, Bradbury J, Chanan G, et al. PyTorch: An Imperative Style, High-Performance Deep Learning Library. 2019. doi: 10.48550\/arXiv.1912.01703"},{"issue":"7","key":"pcbi.1014856.ref115","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1080\/13506280444000049","article-title":"Inverse discrimination time as a perceptual distance for alphabetic characters","volume":"11","author":"P Courrieu","year":"2004","journal-title":"Visual Cognition"},{"issue":"9","key":"pcbi.1014856.ref116","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1010522","article-title":"General object-based features account for letter perception","volume":"18","author":"D Janini","year":"2022","journal-title":"PLoS Comput Biol"},{"issue":"7","key":"pcbi.1014856.ref117","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1364\/JOSAA.2.001160","article-title":"Uncertainty relation for resolution in space, spatial frequency, and orientation optimized by two-dimensional visual cortical filters","volume":"2","author":"JG Daugman","year":"1985","journal-title":"J Opt Soc Am A"},{"issue":"12","key":"pcbi.1014856.ref118","doi-asserted-by":"crossref","first-page":"2379","DOI":"10.1364\/JOSAA.4.002379","article-title":"Relations between the statistics of natural images and the response properties of cortical cells","volume":"4","author":"DJ Field","year":"1987","journal-title":"J Opt Soc Am A"},{"issue":"2","key":"pcbi.1014856.ref119","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1007\/s11263-006-0026-8","article-title":"Self-Invertible 2D Log-Gabor Wavelets","volume":"75","author":"S Fischer","year":"2007","journal-title":"Int J Comput Vis"},{"key":"pcbi.1014856.ref120","unstructured":"W\u00fcrtz R. Multilayer dynamic link networks for establishing image point correspondences and visual object recognition. 1995."},{"issue":"5","key":"pcbi.1014856.ref121","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0267056","article-title":"A psychophysical performance-based approach to the quality assessment of image processing algorithms","volume":"17","author":"DH Baker","year":"2022","journal-title":"PLoS One"},{"issue":"8","key":"pcbi.1014856.ref122","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1167\/10.8.14","article-title":"Spatially extensive summation of contrast energy is revealed by contrast detection of micro-pattern textures","volume":"10","author":"TS Meese","year":"2010","journal-title":"J Vis"},{"issue":"6","key":"pcbi.1014856.ref123","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1152\/jn.1987.58.6.1233","article-title":"An evaluation of the two-dimensional Gabor filter model of simple receptive fields in cat striate cortex","volume":"58","author":"JP Jones","year":"1987","journal-title":"J Neurophysiol"},{"issue":"7825","key":"pcbi.1014856.ref124","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1038\/s41586-020-2649-2","article-title":"Array programming with NumPy","volume":"585","author":"CR Harris","year":"2020","journal-title":"Nature"},{"key":"pcbi.1014856.ref125","volume-title":"Psychophysics: A Practical Introduction","author":"FA Kingdom","edition":"2"},{"issue":"3","key":"pcbi.1014856.ref126","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1038\/s41592-020-0772-5","article-title":"Author Correction: SciPy 1.0: fundamental algorithms for scientific computing in Python","volume":"17","author":"P Virtanen","year":"2020","journal-title":"Nat Methods"},{"key":"pcbi.1014856.ref127","volume-title":"Exploratory Data Analysis","author":"JW Tukey","year":"1977"},{"issue":"10","key":"pcbi.1014856.ref128","article-title":"Contrast sensitivity in natural scenes depends on edge as well as spatial frequency structure","volume":"9","author":"PJ Bex","year":"2009","journal-title":"J Vis"},{"issue":"1","key":"pcbi.1014856.ref129","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1068\/p190079","article-title":"The standard deviation of luminance as a metric for contrast in random-dot images","volume":"19","author":"B Moulden","year":"1990","journal-title":"Perception"},{"issue":"10","key":"pcbi.1014856.ref130","doi-asserted-by":"crossref","first-page":"2032","DOI":"10.1364\/JOSAA.7.002032","article-title":"Contrast in complex images","volume":"7","author":"E Peli","year":"1990","journal-title":"J Opt Soc Am A"},{"issue":"9","key":"pcbi.1014856.ref131","doi-asserted-by":"crossref","first-page":"170285","DOI":"10.1098\/rsos.170285","article-title":"Perception of global image contrast involves transparent spatial filtering and the integration and suppression of local contrasts (not RMS contrast)","volume":"4","author":"TS Meese","year":"2017","journal-title":"R Soc Open Sci"},{"key":"pcbi.1014856.ref132","volume-title":"Deep Learning with Python, Second Edition","author":"F Chollet","year":"2021"},{"key":"pcbi.1014856.ref133","unstructured":"Kingma DP, Ba J. Adam: A method for stochastic optimization. 2014."}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1014856","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014856","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T17:54:50Z","timestamp":1792086890000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014856"}},"subtitle":[],"editor":[{"given":"Jian","family":"Liu","sequence":"first","affiliation":[{"name":"University of Birmingham","place":["UNITED KINGDOM OF GREAT BRITAIN AND NORTHERN IRELAND"]}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,10,12]]},"references-count":133,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2026,10,12]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1014856","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,12]]}}}