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A critical step towards this goal is the development of successful computational encoding models. For ganglion cells in the vertebrate retina, the development of satisfactory models for responses to natural visual scenes is an ongoing challenge. Standard models typically apply linear integration of visual stimuli over space, yet many ganglion cells are known to show nonlinear spatial integration, in particular when stimulated with contrast-reversing gratings. We here study the influence of spatial nonlinearities in the encoding of natural images by ganglion cells, using multielectrode-array recordings from isolated salamander and mouse retinas. We assess how responses to natural images depend on first- and second-order statistics of spatial patterns inside the receptive field. This leads us to a simple extension of current standard ganglion cell models. We show that taking not only the weighted average of light intensity inside the receptive field into account but also its variance over space can partly account for nonlinear integration and substantially improve response predictions of responses to novel images. For salamander ganglion cells, we find that response predictions for cell classes with large receptive fields profit most from including spatial contrast information. Finally, we demonstrate how this model framework can be used to assess the spatial scale of nonlinear integration. Our results underscore that nonlinear spatial stimulus integration translates to stimulation with natural images. Furthermore, the introduced model framework provides a simple, yet powerful extension of standard models and may serve as a benchmark for the development of more detailed models of the nonlinear structure of receptive fields.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1009925","type":"journal-article","created":{"date-parts":[[2022,3,8]],"date-time":"2022-03-08T13:48:17Z","timestamp":1646747297000},"page":"e1009925","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":21,"title":["Simple model for encoding natural images by retinal ganglion cells with nonlinear spatial integration"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5391-7213","authenticated-orcid":true,"given":"Jian 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Schwartz","year":"2011","journal-title":"J Gen Physiol"},{"key":"pcbi.1009925.ref023","doi-asserted-by":"crossref","first-page":"338","DOI":"10.1016\/j.jphysparis.2012.12.001","article-title":"Features and functions of nonlinear spatial integration by retinal ganglion cells","volume":"107","author":"T Gollisch","year":"2013","journal-title":"J Physiol Paris"},{"key":"pcbi.1009925.ref024","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1038\/s41593-018-0284-0","article-title":"Stimulus- and goal-oriented frameworks for understanding natural vision","volume":"22","author":"MH Turner","year":"2019","journal-title":"Nat Neurosci"},{"key":"pcbi.1009925.ref025","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1167\/11.6.11","article-title":"Dependence of the retinal ganglion cell\u2019s responses on local textures of natural scenes","volume":"11","author":"X Cao","year":"2011","journal-title":"J 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dynamic nonlinear stimulus integration in the retina","volume":"73","author":"D B\u00f6linger","year":"2012","journal-title":"Neuron"},{"key":"pcbi.1009925.ref036","doi-asserted-by":"crossref","first-page":"8713","DOI":"10.1038\/s41598-019-45048-8","article-title":"Functional characterization of retinal ganglion cells using tailored nonlinear modeling","volume":"9","author":"Q Shi","year":"2019","journal-title":"Sci Rep"},{"key":"pcbi.1009925.ref037","doi-asserted-by":"crossref","first-page":"e1006291","DOI":"10.1371\/journal.pcbi.1006291","article-title":"Inferring hidden structure in multilayered neural circuits","volume":"14","author":"N Maheswaranathan","year":"2018","journal-title":"PLOS Comput Biol"},{"key":"pcbi.1009925.ref038","doi-asserted-by":"crossref","first-page":"5411","DOI":"10.1073\/pnas.94.10.5411","article-title":"The structure and precision of retinal spike trains","volume":"94","author":"MJ Berry","year":"1997","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1009925.ref039","doi-asserted-by":"crossref","first-page":"2200","DOI":"10.1523\/JNEUROSCI.18-06-02200.1998","article-title":"Refractoriness and neural precision","volume":"18","author":"MJ Berry","year":"1998","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref040","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1080\/713663221","article-title":"A simple white noise analysis of neuronal light responses","volume":"12","author":"EJ Chichilnisky","year":"2001","journal-title":"Network"},{"key":"pcbi.1009925.ref041","doi-asserted-by":"crossref","first-page":"2277","DOI":"10.1152\/jn.00928.2005","article-title":"Functional organization of ganglion cells in the salamander retina","volume":"95","author":"R Segev","year":"2006","journal-title":"J Neurophysiol"},{"key":"pcbi.1009925.ref042","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1017\/S0952523898152021","article-title":"Responses of ganglion cells to contrast steps in the light-adapted retina of the tiger salamander","volume":"15","author":"DA Burkhardt","year":"1998","journal-title":"Vis Neurosci"},{"key":"pcbi.1009925.ref043","doi-asserted-by":"crossref","first-page":"14859","DOI":"10.1523\/JNEUROSCI.0723-12.2012","article-title":"Mapping a complete neural population in the retina","volume":"32","author":"O Marre","year":"2012","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref044","doi-asserted-by":"crossref","first-page":"e1003143","DOI":"10.1371\/journal.pcbi.1003143","article-title":"Inferring nonlinear neuronal computation based on physiologically plausible inputs","volume":"9","author":"JM McFarland","year":"2013","journal-title":"PLOS Comput Biol"},{"key":"pcbi.1009925.ref045","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1007\/s00422-008-0252-y","article-title":"Modeling convergent ON and OFF pathways in the early visual system.","volume":"99","author":"T Gollisch","year":"2008","journal-title":"Biol Cybern"},{"key":"pcbi.1009925.ref046","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/j.semcdb.2020.04.010","article-title":"What the salamander eye has been telling the vision scientist\u2019s brain","volume":"106","author":"F Rozenblit","year":"2020","journal-title":"Semin Cell Dev Biol"},{"key":"pcbi.1009925.ref047","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1016\/j.visres.2015.12.007","article-title":"Physiological and morphological characterization of ganglion cells in the salamander retina","volume":"119","author":"J Wang","year":"2016","journal-title":"Vision Res"},{"key":"pcbi.1009925.ref048","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1016\/j.neuron.2013.11.029","article-title":"The projective field of retinal bipolar cells and its modulation by visual context","volume":"81","author":"H Asari","year":"2014","journal-title":"Neuron"},{"key":"pcbi.1009925.ref049","article-title":"Dataset\u2014Mouse retinal ganglion cell responses to natural images and spatial-integration analysis","author":"D Karamanlis","year":"2021","journal-title":"G-Node"},{"key":"pcbi.1009925.ref050","doi-asserted-by":"crossref","first-page":"4462","DOI":"10.1523\/JNEUROSCI.20-12-04462.2000","article-title":"Functional architecture of synapses in the inner retina: segregation of visual signals by stratification of bipolar cell axon terminals","volume":"20","author":"SM Wu","year":"2000","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref051","doi-asserted-by":"crossref","first-page":"6807","DOI":"10.1523\/JNEUROSCI.4206-07.2008","article-title":"A retinal circuit that computes object motion","volume":"28","author":"SA Baccus","year":"2008","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref052","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1038\/nature07140","article-title":"Spatio-temporal correlations and visual signalling in a complete neuronal population","volume":"454","author":"JW Pillow","year":"2008","journal-title":"Nature"},{"key":"pcbi.1009925.ref053","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1088\/0954-898X_15_4_002","article-title":"Maximum likelihood estimation of cascade point-process neural encoding models","volume":"15","author":"L. Paninski","year":"2004","journal-title":"Network"},{"key":"pcbi.1009925.ref054","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1038\/18678","article-title":"Anticipation of moving stimuli by the retina","volume":"398","author":"MJ Berry","year":"1999","journal-title":"Nature"},{"key":"pcbi.1009925.ref055","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1016\/S0896-6273(01)00322-1","article-title":"Predicting every spike: a model for the responses of visual neurons","volume":"30","author":"J Keat","year":"2001","journal-title":"Neuron"},{"key":"pcbi.1009925.ref056","first-page":"1369","article-title":"Deep learning models of the retinal response to natural scenes","volume":"29","author":"LT McIntosh","year":"2016","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1009925.ref057","doi-asserted-by":"crossref","first-page":"14829","DOI":"10.1523\/JNEUROSCI.2815-13.2015","article-title":"A convolutional subunit model for neuronal responses in macaque V1","volume":"35","author":"B Vintch","year":"2015","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref058","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.cub.2016.11.040","article-title":"Neural circuit inference from function to structure","volume":"27","author":"E Real","year":"2017","journal-title":"Curr Biol"},{"key":"pcbi.1009925.ref059","doi-asserted-by":"crossref","first-page":"1964","DOI":"10.1038\/s41467-017-02159-y","article-title":"Multiplexed computations in retinal ganglion cells of a single type","volume":"8","author":"S Deny","year":"2017","journal-title":"Nat Commun"},{"key":"pcbi.1009925.ref060","first-page":"8535","article-title":"From deep learning to mechanistic understanding in neuroscience: the structure of retinal prediction","volume":"32","author":"H Tanaka","year":"2019","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1009925.ref061","first-page":"1692","article-title":"Bayesian spike-triggered covariance","volume":"24","author":"IM Park","year":"2011","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1009925.ref062","first-page":"2454","article-title":"Spectral methods for neural characterization using generalized quadratic models","volume":"26","author":"IM Park","year":"2013","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1009925.ref063","doi-asserted-by":"crossref","first-page":"1661","DOI":"10.1162\/NECO_a_00463","article-title":"Learning quadratic receptive fields from neural responses to natural stimuli","volume":"25","author":"K Rajan","year":"2013","journal-title":"Neural Comput"},{"key":"pcbi.1009925.ref064","doi-asserted-by":"crossref","first-page":"11003","DOI":"10.1523\/JNEUROSCI.3305-05.2005","article-title":"Prediction and decoding of retinal ganglion cell responses with a probabilistic spiking model","volume":"25","author":"JW Pillow","year":"2005","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref065","doi-asserted-by":"crossref","unstructured":"Lindsey J, Ocko SA, Ganguli S, Deny S. A unified theory of early visual representations from retina to cortex through anatomically constrained deep CNNs. 7th International Conference on Learning Representation (ICLR). New Orleans, LA: OpenReview.net; 2019. Available: https:\/\/openreview.net\/forum?id=S1xq3oR5tQ","DOI":"10.1101\/511535"},{"key":"pcbi.1009925.ref066","doi-asserted-by":"crossref","first-page":"1202","DOI":"10.1126\/science.1225266","article-title":"A large-scale model of the functioning brain","volume":"338","author":"C Eliasmith","year":"2012","journal-title":"Science"},{"key":"pcbi.1009925.ref067","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1167\/14.12.22","article-title":"Retina-V1 model of detectability across the visual field","volume":"14","author":"C Bradley","year":"2014","journal-title":"J Vis"},{"key":"pcbi.1009925.ref068","doi-asserted-by":"crossref","first-page":"7548","DOI":"10.1523\/JNEUROSCI.0413-14.2014","article-title":"Nonlinear spatial integration in the receptive field surround of retinal ganglion cells","volume":"34","author":"D Takeshita","year":"2014","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref069","doi-asserted-by":"crossref","first-page":"1900","DOI":"10.1038\/s41467-021-22042-1","article-title":"Linear and nonlinear chromatic integration in the mouse retina","volume":"12","author":"MH Khani","year":"2021","journal-title":"Nat Commun."},{"key":"pcbi.1009925.ref070","doi-asserted-by":"crossref","first-page":"1692","DOI":"10.1016\/j.neuron.2021.03.015","article-title":"Nonlinear spatial integration in retinal bipolar cells shapes the encoding of artificial and natural stimuli","volume":"109","author":"HM Schreyer","year":"2021","journal-title":"Neuron"},{"key":"pcbi.1009925.ref071","doi-asserted-by":"crossref","first-page":"1206","DOI":"10.1016\/j.neuron.2013.09.030","article-title":"A tale of two retinal domains: near-optimal sampling of achromatic contrasts in natural scenes through asymmetric photoreceptor distribution","volume":"80","author":"T Baden","year":"2013","journal-title":"Neuron"},{"key":"pcbi.1009925.ref072","doi-asserted-by":"crossref","first-page":"e1003289","DOI":"10.1371\/journal.pcbi.1003289","article-title":"Dynamical adaptation in photoreceptors","volume":"9","author":"DA Clark","year":"2013","journal-title":"PLOS Comput Biol"},{"key":"pcbi.1009925.ref073","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1113\/jphysiol.2009.179036","article-title":"Cones perform a non-linear transformation on natural stimuli","volume":"588","author":"D Endeman","year":"2010","journal-title":"J Physiol"},{"key":"pcbi.1009925.ref074","doi-asserted-by":"crossref","first-page":"e2001210","DOI":"10.1371\/journal.pbio.2001210","article-title":"A novel mechanism of cone photoreceptor adaptation","volume":"15","author":"MHC Howlett","year":"2017","journal-title":"PLOS Biol"},{"key":"pcbi.1009925.ref075","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1113\/jphysiol.1987.sp016443","article-title":"The receptive-field spatial structure of cat retinal Y cells","volume":"384","author":"C Enroth-Cugell","year":"1987","journal-title":"J Physiol"},{"key":"pcbi.1009925.ref076","doi-asserted-by":"crossref","first-page":"11277","DOI":"10.1523\/JNEUROSCI.2982-08.2008","article-title":"Y-cell receptive field and collicular projection of parasol ganglion cells in macaque monkey retina","volume":"28","author":"JD Crook","year":"2008","journal-title":"J Neurosci"},{"key":"pcbi.1009925.ref077","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1016\/j.preteyeres.2018.06.003","article-title":"The dynamic receptive fields of retinal ganglion cells","volume":"67","author":"S Wienbar","year":"2018","journal-title":"Prog Retin Eye Res"},{"key":"pcbi.1009925.ref078","doi-asserted-by":"crossref","first-page":"e1004425","DOI":"10.1371\/journal.pcbi.1004425","article-title":"Spike-Triggered Covariance Analysis Reveals Phenomenological Diversity of Contrast Adaptation in the 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