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ImageNet classification with deep convolutional neural networks. In: Advances in neural information processing systems; 2012. p. 1097\u20131105."},{"key":"ref10","doi-asserted-by":"crossref","unstructured":"Taigman Y, Yang M, Ranzato MA. Deepface: Closing the gap to human-level performance in face verification. In: The IEEE Conference on Computer Vision and Pattern Recognition; 2014. p. 1701\u20131708.","DOI":"10.1109\/CVPR.2014.220"},{"issue":"23","key":"ref11","doi-asserted-by":"crossref","first-page":"8619","DOI":"10.1073\/pnas.1403112111","article-title":"Performance-optimized hierarchical models predict neural responses in higher visual cortex","volume":"111","author":"DLK Yamins","year":"2014","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"25025","DOI":"10.1038\/srep25025","article-title":"A specialized face-processing model inspired by the organization of monkey face patches explains several face-specific phenomena observed in humans","volume":"6","author":"A Farzmahdi","year":"2016","journal-title":"Scientific Reports"},{"issue":"3","key":"ref13","doi-asserted-by":"crossref","first-page":"e0150980","DOI":"10.1371\/journal.pone.0150980","article-title":"Neural Tuning Size in a Model of Primate Visual Processing Accounts for Three Key Markers of Holistic Face Processing","volume":"11","author":"C Tan","year":"2016","journal-title":"PloS one"},{"issue":"6583","key":"ref14","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1038\/381607a0","article-title":"Emergence of simple-cell receptive field properties by learning a sparse code for natural images","volume":"381","author":"BA Olshausen","year":"1996","journal-title":"Nature"},{"issue":"1394","key":"ref15","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1098\/rspb.1998.0303","article-title":"Independent component filters of natural images compared with simple cells in primary visual cortex","volume":"265","author":"JH van Hateren","year":"1998","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"1","key":"ref16","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1038\/4580","article-title":"Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects","volume":"2","author":"RP Rao","year":"1999","journal-title":"Nature Neuroscience"},{"issue":"7","key":"ref17","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1162\/089976600300015312","article-title":"Emergence of phase-and shift-invariant features by decomposition of natural images into independent feature subspaces","volume":"12","author":"A Hyv\u00e4rinen","year":"2000","journal-title":"Neural Computation"},{"issue":"7225","key":"ref18","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1038\/nature07481","article-title":"Emergence of complex cell properties by learning to generalize in natural scenes","volume":"457","author":"Y Karklin","year":"2009","journal-title":"Nature"},{"issue":"8","key":"ref19","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1038\/90526","article-title":"Natural signal statistics and sensory gain control","volume":"4","author":"O Schwartz","year":"2001","journal-title":"Nature Neuroscience"},{"issue":"8","key":"ref20","doi-asserted-by":"crossref","first-page":"2119","DOI":"10.1162\/NECO_a_00310","article-title":"Multinomial Bayesian learning for modeling classical and nonclassical receptive field properties","volume":"24","author":"H Hosoya","year":"2012","journal-title":"Neural Computation"},{"issue":"5","key":"ref21","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/j.jphysparis.2013.01.001","article-title":"A three-layer model of natural image statistics","volume":"107","author":"MU Gutmann","year":"2013","journal-title":"Journal of Physiology-Paris"},{"issue":"29","key":"ref22","doi-asserted-by":"crossref","first-page":"10412","DOI":"10.1523\/JNEUROSCI.5152-14.2015","article-title":"A Hierarchical Statistical Model of Natural Images Explains Tuning Properties in V2","volume":"35","author":"H Hosoya","year":"2015","journal-title":"Journal of Neuroscience"},{"issue":"6755","key":"ref23","doi-asserted-by":"crossref","first-page":"788","DOI":"10.1038\/44565","article-title":"Learning the parts of objects by non-negative matrix factorization","volume":"401","author":"DD Lee","year":"1999","journal-title":"Nature"},{"issue":"1","key":"ref24","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.cviu.2005.09.012","article-title":"Learning generative visual models from few training examples: An incremental bayesian approach tested on 101 object categories","volume":"106","author":"L Fei-Fei","year":"2007","journal-title":"Computer Vision and Image Understanding"},{"issue":"5761","key":"ref25","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1126\/science.1119983","article-title":"A cortical region consisting entirely of face-selective cells","volume":"311","author":"DY Tsao","year":"2006","journal-title":"Science"},{"issue":"11","key":"ref26","doi-asserted-by":"crossref","first-page":"4302","DOI":"10.1523\/JNEUROSCI.17-11-04302.1997","article-title":"The fusiform face area: a module in human extrastriate cortex specialized for face perception","volume":"17","author":"N Kanwisher","year":"1997","journal-title":"The Journal of Neuroscience"},{"issue":"3","key":"ref27","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1152\/jn.1987.57.3.835","article-title":"Visual properties of neurons in area V4 of the macaque: sensitivity to stimulus form","volume":"57","author":"R Desimone","year":"1987","journal-title":"Journal of Neurophysiology"},{"key":"ref28","first-page":"07","article-title":"Labeled faces in the wild: A database for studying face recognition in unconstrained environments","author":"GB Huang","year":"2007"},{"key":"ref29","unstructured":"Huang G, Mattar M, Lee H. Learning to align from scratch. Advances in neural information processing systems. 2012;p. 764\u2013772."},{"key":"ref30","article-title":"Pattern recognition and machine learning (information science and statistics)","author":"CM Bishop","year":"2006"},{"issue":"23","key":"ref31","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 Research"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1162\/NECO_a_00843","article-title":"Learning Visual Spatial Pooling by Strong PCA Dimension Reduction","volume":"28","author":"H Hosoya","year":"2016","journal-title":"Neural Computation"},{"key":"ref33","first-page":"695","article-title":"Estimation of Non-Normalized Statistical Models by Score Matching","volume":"6","author":"A Hyv\u00e4rinen","year":"2005","journal-title":"The Journal of Machine Learning Research"},{"issue":"6","key":"ref34","doi-asserted-by":"crossref","first-page":"1450","DOI":"10.1109\/TNN.2002.804287","article-title":"Face recognition by independent component analysis","volume":"13","author":"MS Bartlett","year":"2002","journal-title":"IEEE Transactions on Neural Networks"},{"issue":"01","key":"ref35","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1017\/S0952523809090051","article-title":"Efficient coding correlates with spatial frequency tuning in a model of V1 receptive field organization","volume":"26","author":"J Wiltschut","year":"2009","journal-title":"Visual Neuroscience"},{"key":"ref36","unstructured":"Yildirim I, Kulkarni TD, Freiwald WA. Efficient and robust analysis-by-synthesis in vision: A computational framework, behavioral tests, and modeling neuronal representations. Annual Conference of the Cognitive Science Society. 2015;."},{"key":"ref37","first-page":"249","article-title":"Qualitative representations for recognition","volume":"2525","author":"P Sinha","year":"2002","journal-title":"Workshop on Biologically Motivated Computer Vision (Lecture Notes in Computer Science)"},{"issue":"2","key":"ref38","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1023\/B:VISI.0000013087.49260.fb","article-title":"Robust real-time face detection","volume":"57","author":"P Viola","year":"2004","journal-title":"International Journal of Computer Vision"},{"issue":"1","key":"ref39","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1073\/pnas.0706079105","article-title":"Face perception in monkeys reared with no exposure to faces","volume":"105","author":"Y Sugita","year":"2008","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"2","key":"ref40","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1037\/0033-295X.98.2.164","article-title":"CONSPEC and CONLERN: a two-process theory of infant face recognition","volume":"98","author":"J Morton","year":"1991","journal-title":"Psychological review"},{"key":"ref41","first-page":"217","article-title":"Sensory communication","author":"HB Barlow","year":"1961"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1068\/p010371","article-title":"Single units and sensation: a neuron doctrine for perceptual psychology?","volume":"1","author":"HB Barlow","year":"1972","journal-title":"Perception"},{"issue":"10","key":"ref43","article-title":"ICA mixture models for unsupervised classification of non-Gaussian classes and automatic context switching in blind signal separation","volume":"22","author":"TW Lee","year":"2000","journal-title":"IEEE Transactions on Pattern Analysis and Machine Intelligence"},{"issue":"5881","key":"ref44","doi-asserted-by":"crossref","first-page":"1355","DOI":"10.1126\/science.1157436","article-title":"Patches with Links: A Unified System for Processing Faces in the Macaque Temporal Lobe","volume":"320","author":"S Moeller","year":"2008","journal-title":"Science"},{"issue":"6","key":"ref45","doi-asserted-by":"crossref","first-page":"1325","DOI":"10.1016\/j.neuron.2016.05.009","article-title":"Anatomical Connections of the Functionally Defined \u201cFace Patches\u201d in the Macaque Monkey","volume":"90","author":"P Grimaldi","year":"2016","journal-title":"Neuron"},{"issue":"3","key":"ref46","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1016\/j.neuron.2012.03.024","article-title":"What Makes a Cell Face Selective? The Importance of Contrast","volume":"74","author":"S Ohayon","year":"2012","journal-title":"Neuron"},{"issue":"6005","key":"ref47","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1126\/science.1194908","article-title":"Functional compartmentalization and viewpoint generalization within the macaque face-processing system","volume":"330","author":"WA Freiwald","year":"2010","journal-title":"Science"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1005667","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2017,8,8]],"date-time":"2017-08-08T00:00:00Z","timestamp":1502150400000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1005667","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2019,10,1]],"date-time":"2019-10-01T11:44:23Z","timestamp":1569930263000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1005667"}},"subtitle":[],"editor":[{"given":"Wolfgang","family":"Einh\u00e4user","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2017,7,25]]},"references-count":47,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2017,7,25]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1005667","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/086637","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,7,25]]}}}