From Basic Network Principles to Neural Architecture: Emergence of Orientation ColumnsRalph Linsker doi:10.1073/pnas.83.22.8779 1986;83;8779-8783 PNAS This information is current as of December 2006. www.pnas.org#otherarticlesThis article has been cited by other articles: E-mail Alerts. click heretop right corner of the article orReceive free email alerts when new articles cite this article - sign up in the box at the Rights & Permissions www.pnas.org/misc/rightperm.shtmlTo reproduce this article in part (figures, tables) or in entirety, see: Reprints www.pnas.org/misc/reprints.shtmlTo order reprints, see: Notes:Proc.Natl.Acad.Sci.USAVol.83,pp.8779-8783,November1986NeurobiologyFrombasicnetworkprinciplestoneuralarchitecture:Emergenceoforientationcolumns*(modularself-adaptivenetworks/visualsystem/feature-analyzingcells/orientation-selectivecells)RALPHLINSKERIBMThomasJ.WatsonResearchCenter,YorktownHeights,NY10598CommunicatedbyRichardL.Garwin,July7,1986ABSTRACTOrientation-selectivecells-cellsthatarese-lectivelyresponsivetobarsandedgesatparticularorienta-tions-areasalientfeatureofthearchitectureofmammalianvisualcortex.Inthepreviouspaperofthisseries,Ishowedthatsuchcellsemergespontaneouslyduringthedevelopmentofasimplemultilayerednetworkhavinglocalbutinitiallyrandomfeedforwardconnectionsthatmature,onelayeratatime,accordingtoasimpledevelopmentrule(ofHebbtype).Inthispaper,Ishowthat,inthepresenceoflateralconnectionsbetweendevelopingorientationcells,thesecellsself-organizeintobandedpatternsofcellsofsimilarorientation.Thesepatternsaresimilartothe"orientationcolumns"foundinmammalianvisualcortex.Noorientationpreferenceisspeci-fiedtothesystematanystage,noneofthebasicdevelopmentalrulesisspecifictovisualprocessing,andtheresultsemergeevenintheabsenceofvisualinputtothesystem(ashasbeenobservedinmacaquemonkey).Thisseriesofpapersexplores,withreferencetothemam-malianvisualsystem,thestructuresthatemergeinanetworkconsistingofseverallayersofcellswithconnectionsofinitiallyrandomstrength,whichdevelopaccordingtoaHebb-typerulethat"rewards"correlatedactivityofcon-nectedcells.Inpapers1and2(1,2),Ishowedtheemergenceofspatial-opponentandorientation-selectivecellsinalay-eredsystemwithparallelfeedforwardconnectionsonlyandwithrandomspontaneousuncorrelatedactivity(noenviron-mentalinput)inthefirstlayer.Inprimatevisualcortex,orientation-selectivecellsareorganized,priortoanyvisualexperience,intobandedregions("columns"),suchthatthepreferredcellorientationtendstovarymonotonically,but
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