文獻綜述后一稿

視知覺學習的文1.1視知覺學習的文1.156、空間位置等）以及復雜刺激（879（izan行為短暫變化。知覺學習不同于其他類型學習：不同于陳述性記憶的學習（declarativelearning）需要一個中間的鞏固儲存器（如海馬11,12（specificity皮層如V1和V2。而認知學習、策略選擇、程序性學習一般被認為是一種高級的（generalization程緩慢，一般需較長時間的訓練；（4）RamachandranBraddick13，他們訓45RamachandranBraddick13，他們訓45o傾斜線條組成的立體圖形，經(jīng)過一段時間的練習，被試的操作能力得以大大提高，但如果立體圖形的傾斜線條變?yōu)?35o，被試的能力回到訓Piggio等人在游標（vernier）任務(wù)上也發(fā)現(xiàn)類似的結(jié)果14，他們要求被試判斷呈90o，要求判斷右邊的線4-9,13-20，1Sowden17圖1：朝向和位置特異性學習的刺激和結(jié)果示..表1：知覺學習具有朝向特異性的研訓練和變朝向差遷移果學習任表1：知覺學習具有朝向特異性的研訓練和變朝向差遷移果學習任出注立體Ramachandran&Braddick,Fiorentini&Berardi,1980Poggioetal.,1992Cristetal.,1997Fahle,1993Fahle,Schoupsetal.,無45o和135o不能遷無0o和90o不能互相遷游標辨光柵朝向0o和90o不能互相遷無大于45度很少遷Karni&Sagi,無質(zhì)地檢texture視覺搜對比度檢Ahissar&Hochstein,1996,Sigman&Gilbert,2000Sowdenetal.,全圖4,5,8,9,15,17,18,21-2930,31表2：知覺學習具有位置特異性的研究匯總，引學習任出注立體Ramachandran,無/沒有測量表2：知覺學習具有位置特異性的研究匯總，引學習任出注立體Ramachandran,無/沒有測量Cristetal.,1997Ahissar&Hochstein,無/立體pop-out無/經(jīng)過訓練的被試有Fahle,1994;Fahle,Edelman&Poggio,1995Beard,Levi,&Reich,Karni&Sagi,游標辨無質(zhì)地檢texture圖形辨部無Nazir&O’Regan,1990;Dill&Fahle,1997Sowdenetal.,無亮度對比度檢無線段朝向Shui&Pashler,無光柵朝向Schoupsetal.,無/Juttner&Rentschler,Rentschleretal.,Ball&Sekuler,Gaborpatch部中線相似位置可以遷運動方向部中線相似位置可以遷空間相位Berardi&Fiorentini,無/27實驗讓被試遮蓋一眼用單眼4-6,26,33,34單細胞記錄研究發(fā)現(xiàn)初級視覺皮層細胞的感受野在視野中只有大約單細胞記錄研究發(fā)現(xiàn)初級視覺皮層細胞的感受野在視野中只有大約1o的直徑，V2V4細胞感受野的直徑會有幾度，而前下顳葉（anteriorinferotemporal，AIT）20o36。因此這種對刺激朝向、位置的特2.2比較容易（SOA較長或目標元素與背景元素的朝向差別較大圖圖2：訓練任務(wù)的難度與特異性的....或除中央注視點外的任意位置。目標元素和背景元素的朝向差別決定了任務(wù)難度。d..-30o）能夠遷移到鄰近位置（0.7o或2.1o），而右圖對困難任務(wù)（2pos-16o）表現(xiàn)出很強的置特異性8AhissarHochstein的結(jié)果，Jeter等人指出確切地說應(yīng)該是“遷移任務(wù)的難度（而非訓練任務(wù)的難度）決定了知覺學習的特異性”37。他們訓練刺激（-35o或55o）的朝向偏差區(qū)分，分困難和容易任務(wù)（偏差分別為±5o和±12o，圖3a。訓練階段刺激始終呈現(xiàn)在以注視點為中心的某對角線上，測試階段刺激的位置換到另一對角線了，同時朝向也改變了（3b。結(jié)果發(fā)現(xiàn)不管訓練容易或困難任務(wù)，均能遷移到簡單任務(wù)上（圖3c，而遷移不到困難任務(wù)上（圖圖3：遷移任務(wù)的難圖3：遷移任務(wù)的難度與特異性的a.實驗刺激。困難和容易任務(wù)的測試刺激與被試內(nèi)在的參照刺激（-35o55o）聲（上圖）和高噪聲（下圖）條件下的結(jié)果一致；d.不管訓練任務(wù)容易還是困難，學習Liu和Weinshall發(fā)現(xiàn)即使是先前發(fā)現(xiàn)存在特異性的學習中，其實也具有遷38。他們讓被試進行隨機點運動方向辨別的學習，任務(wù)分容易和困難的（運外，容易任務(wù)的學習能普遍遷移到其他運動方向的容易任務(wù)上，這與Ahissar和Jeter等人的結(jié)果一致8,37。研究同時發(fā)現(xiàn)在某一方向充分訓練容易任務(wù)，再在另一（insight3.3.1早期階段：早期選擇理論和回饋網(wǎng)絡(luò)模型（Recurrent20,3.3.1早期階段：早期選擇理論和回饋網(wǎng)絡(luò)模型（Recurrent20,39theoryHebbian突觸學習的計算模型中，AdiniTsodyks提出一個對比度辨別學習的回饋網(wǎng)絡(luò)模型（RecurrentModel）41。他們的實驗發(fā)現(xiàn)中央刺激的對比度辨別只在周邊有線索刺激的條件下才能發(fā)生（Yu3，423.2中層階段：通道重新賦權(quán)模型（Rewighting習的機制16,43-45，信號增強可以表現(xiàn)為神經(jīng)元數(shù)目增加、反應(yīng)增益增強4443,45,46DosherLu的工作，提出了知覺學習模板模型（PTMModel）4247，認為知覺學習可能存在三種不同機制：（1）信號增強（內(nèi)部噪音減少導致在零或低噪聲下的閾限降低（圖4b（2）外部噪聲去除（模板匹配能力增強導致高外部噪聲的閾限降低（圖4c（3）乘法噪聲降低（增益控制增加有外部噪聲條件下都能觀察到閾限降低（4d。他們的實驗驗證了前兩種，基于此結(jié)果提出“通道重新賦Model生在早期視覺表征區(qū)域（V生在早期視覺表征區(qū)域（V1），如果基本的視覺通道發(fā)生改變，那么學習V1圖4：知覺模板模型和知覺學習三種機制的a.知覺模板模型；b-d.學習前后對比度檢測閾值與添加噪音強度關(guān)系的變化。..低（增益控制增強），在所有外部噪聲條件下都能觀察到閾限降低圖5：“通道重新賦權(quán)模圖5：“通道重新賦權(quán)模型”（RewightingModel）示意3.3高級到低級：反轉(zhuǎn)層次理論（Reverse-hierarchytheoryAhissar和Hochstein根據(jù)實驗數(shù)據(jù)（圖2）（Reverse-hierarchytheory,RHT3.4高級階段：Mollon&Danilova樣描述的：“Indirectly,viathecentralsignalstowhichhehasaccessthesubjectbelearningaboutopticalfeaturesofhisretinalimage;belearningaboutopticalfeaturesofhisretinalimage;aboutthelocaltopographyofhisreceptormosaic;andaboutthespecificwiringofindividualneuronswithinhisvisual50,4.控（如“頓（insight）覺學習中起重要作用52,53。注意可能不僅僅起到調(diào)控作用，它可能就是知覺學習的機制。Goldstone提出知覺學習的機制之一就是注意權(quán)重的改變54，它可以通過增加分配在重要的知覺維度和特征上的注意和55Karni和Sagi提出56，知覺學習可分為“快速學習”（fastlearning）和“慢速學 圖7：質(zhì)地檢測任務(wù)（Texture圖7：質(zhì)地檢測任務(wù)（Texturediscriminationtask）學習.變短。c.快速學習在最初一天內(nèi)的幾組訓練中就表現(xiàn)出來，而慢速學習則持續(xù)知覺學習中反饋起到什么作用？ShuiPashler3之間也不會互相遷移61，表明高級視覺信息加工以及自上而各自的學習（disruption。Seitz2005年報道了一種特異于任務(wù)對知覺學習4.1.4多刺激學習—— 63。隨后我們提出了知覺學習的“刺激4.2知覺學習不需要注意的參與也能發(fā)生。Watanabe55,4.2知覺學習不需要注意的參與也能發(fā)生。Watanabe55,65報告了一種“任learninglearning8b能力的提高66圖8：無注意條件下的..5.（15.1675.1.1SchoupsOrban等人訓練猴子進行光柵朝向辨別68，行為學結(jié)果顯示朝向?qū)W習存在位置和朝向特異性（9a圖9：光柵朝向辨別行為和神經(jīng)元圖9：光柵朝向辨別行為和神經(jīng)元.位置（綠色），被動呈現(xiàn)刺激的位置（紅色，passive）。行為學結(jié)果表明學習存在置和朝向特異性。b.來自V1的第2～3層，5～6層數(shù)據(jù)的匯總68V1V2感小但是結(jié)果還是顯著的。他們沒有從生理學水平（至少在V1或V2）發(fā)現(xiàn)神經(jīng)并不一4,28,68?？赡艿脑蛴校海?）盡管使用相似的實驗參數(shù)，如訓練朝向、網(wǎng)膜位置，Ghose3～8o，而Schoups文章訓練后的值僅有0.6～1.2o，有沒有可能是因為其中包含了策略性的學learninggaborYangMausellGhose70YangMausellGhose70RaiguelOrban7169。相反地，Yang等人265.1.2Crist，LiGibert的工作也發(fā)現(xiàn)訓練并沒有改變初級視覺皮層的感受野特性（如位置、大小、朝向選擇性皮層放大系數(shù)60。有趣的是，他effect訓練任務(wù)（兩分任務(wù)）V1神經(jīng)元的背景調(diào)節(jié)（contextualeffect）圖10：訓練兩分任務(wù)（圖10：訓練兩分任務(wù)（bisection）改變背景調(diào)節(jié)（contextual（11上V1神經(jīng)元的活動。結(jié)果發(fā)現(xiàn)即使刺激相似，但執(zhí)行任務(wù)不同，V1神經(jīng)元的反應(yīng)特性也可以被特異地動態(tài)改變（圖圖11：任務(wù)調(diào)控神經(jīng)元反圖11：任務(wù)調(diào)控神經(jīng)元反..5.1.374最近，LawGold75，研究者訓練猴子進行視覺運動MTIPS區(qū)的神經(jīng)元活動，MT主要表征解決任務(wù)的74最近，LawGold75，研究者訓練猴子進行視覺運動MTIPS區(qū)的神經(jīng)元活動，MT主要表征解決任務(wù)的LIPMT信號改變的（invariance）的機制，感受野對物體位置也具有選擇性，物體不變性并非與生俱來的，經(jīng)過訓練的8082（168,69，即V455.2ERP的研究報道了知覺學習引起早5.2ERP的研究報道了知覺學習引起早期成分的變87-90，SkrandiesERP幅值增大，潛伏期減少，這些變化主要發(fā)生91。DingSong92，行為學表現(xiàn)出朝向特異性，腦電數(shù)據(jù)在頂區(qū)的N1（140ms左右）波幅表現(xiàn)出朝向遷移P2（200ms左右）波幅上表現(xiàn)出朝向特異性。雖然特異性學習5.310年知覺學習的腦成像研究隨著技術(shù)手段的成熟而發(fā)展起來，目前的研imaging，fMRI顱磁刺激TMS（TranscranialmagneticstimulationTMSPE（Positronemission5.2.1Schiltz1999PET技術(shù)比較被試進行朝向分辨訓練前后的血流水外的區(qū)域，如右距狀溝（rightcalcarinesulcus，左舌回（leftlingualgyrus，左中枕（leftmiddleociitlgyrusSchwartz2002fMRI2794結(jié)果發(fā)現(xiàn)在早期視覺皮層等網(wǎng)膜拓撲區(qū)域（如V1、V2），訓練眼相較于非訓練Furmanski2004fMRIV195BOLDFurmanski2004fMRIV195BOLD信號的變化。訓斜朝向誘發(fā)的信號增強了V1（2fMRNeary2005TMS96。Mukai2007fMRI工作提出根據(jù)大腦視覺和注意相關(guān)區(qū)域的活動Brodman18、19區(qū)）FG（fusiformgyrus）及一些跟注sulcusFE（frontalIPSFEF圖12：“學習組”和“無學習圖12：“學習組”和“無學習組”數(shù).（ROI.93。針對這一矛盾的結(jié)果，Yotsumoto93。針對這一矛盾的結(jié)果，Yotsumoto水平（13A，post3pre相比圖13：訓練位置引起腦區(qū)活動變化指.sulcusSP（superiorgyrusMF（middlegyrus5.3.2傳統(tǒng)觀點認為，復雜圖形的表征在相對高級的大腦區(qū)域完成。Sigman2005年的工作表明視覺學習可能會導致人們對復雜物體的表征從高級皮層區(qū)域5.3.2傳統(tǒng)觀點認為，復雜圖形的表征在相對高級的大腦區(qū)域完成。Sigman2005年的工作表明視覺學習可能會導致人們對復雜物體的表征從高級皮層區(qū)域 著性圖形（low-salienceshapes）的學習從V1到高級視覺區(qū)域信號都增強了，學習通過調(diào)節(jié)LOC的聯(lián)系以編碼主要特征。異，發(fā)現(xiàn)訓練提高了對錯覺輪廓的反應(yīng)。行為學結(jié)果表明學習效果可以保持10deBeeck105107？對一些知覺任務(wù)來說，學習在一些107？對一些知覺任務(wù)來說，學習在一些相關(guān)感覺皮層就能發(fā)生6.（mbla108-114。例Ploat2004年的研究報道108，研究者通過訓練成人弱視進行對比度檢測任務(wù)（刺激見14a左下角，訓練提高了對比度檢測能力，使得對比度敏感函數(shù)曲線大大提高14b（crowding）也大大減小，這一結(jié)果提示弱視知覺學習可以從簡單的對比度110。但關(guān)于弱視知覺學習的機制目前尚不清楚，LiLevi112,115圖14：成人弱視訓練a．訓練對比圖14：成人弱視訓練a．訓練對比度檢測任務(wù)（左下角示意圖。訓練前后比較對比度敏感函數(shù)曲線7.差異等原因所導致的。例如，F(xiàn)ine和Jacobs比較了從朝向辨別到圖形識別的一系列心理物理學的研究116，將訓練效果均轉(zhuǎn)換成d'進行比較，發(fā)現(xiàn)綜述參Merzenich,M.M.&Sameshima,K.Corticalplasticityandmemory.綜述參Merzenich,M.M.&Sameshima,K.Corticalplasticityandmemory.CurrOpin3,187-96Gibson,E.J.Perceptuallearning.AnnuRevPsychol14,29-56Yu,C.,Klein,S.A.&Levi,D.M.Perceptuallearningincontrastdiscriminationandthe(minimal)roleofcontext.JVis4,169-82(2004).Schoups,A.A.,Vogels,R.&Orban,G.A.Humanperceptuallearninginidentifyingtheobliqueorientation:retinotopy,orientationspecificityandmonocularity.JPhysiol483(Pt3),797-810(1995).27,953-65Fiorentini,A.&Berardi,N.Perceptuallearningspecificfororientationandspatialfrequency.Nature287,43-44(1980).McKee,S.P.&Westheimer,G.Improvementinvernieracuitywithpractice.PerceptPsychophys24,258-62(1978).Ahissar,M.&Hochstein,S.TaskdifficultyandthespecificityofperceptualNature387,401-6Sigman,M.&Gilbert,C.D.Learningtofindashape.NatNeurosci3,264-9(2000).Bruce,V.,Burton,A.M.inPerceptualLearning(eds.Fahle,M.&Poggio,T.)ix-xx(MITPress,Cambridge,MA,USA,2002).Fahle,M.inPerceptualLearning(eds.Fahle,M.&Poggio,T.)ix-xx(MITPress,Cambridge,MA,USA,2002).Fahle,M.Perceptuallearning:specificityversusgeneralization.CurrOpinNeurobiol15,154-60(2005).Ramachandran,V.S.&Braddick,O.Orientation-specificlearninginstereopsis.2,371-6Poggio,T.,Fahle,M.&Edelman,S.Fastperceptuallearninginvisualhyperacuity.256,1018-21Ahissar,M.&Hochstein,S.Learningpop-outdetection:specificitiestostimuluscharacteristics.VisionRes36,3487-500(1996).Folta,K.NeuralfinetuningduringVernieracuitytraining?VisionRes43,1177-85(2003).Sowden,P.T.,Rose,D.&Davies,I.R.Perceptuallearningofluminancecontrastdetection:specificforspatialfrequencyandretinallocationbutnotorientation.VisionResearch42,1249-58(2002).Crist,R.E.,Kapadia,M.K.,Westheimer,G.&Gilbert,C.D.Perceptuallearningofspatiallocalization:specificityfororientation,position,andcontext.JNeurophysiol78,2889-94Ball,K.&Sekuler,R.Aspecificandenduringimprovementinvisualmotiondiscrimination.Science218,697-8(1982).Fahle,M.Visuallearninginthehyperacuityrangeinadults.GerJOphthalmol2,83-6Ramachandran,V.S.Learning-likephenomenainstereopsis.Nature262,382-4O'Toole,A.J.&Kersten,D.J.Learningtoseerandom-dotstereograms.Perception21,43Westheimer,G.&Truong,T.T.Targetcrowdinginfovealandperipheral43Westheimer,G.&Truong,T.T.Targetcrowdinginfovealandperipheralstereoacuity.AmJOptomPhysiolOpt65,395-9(1988).Fahle,M.Humanpatternrecognition:parallelprocessingandperceptualFahle,M.,Edelman,S.&Poggio,T.Fastperceptuallearninginhyperacuity.VisionRes35,3003-13(1995).Beard,B.L.,Levi,D.M.&Reich,L.N.Perceptuallearninginparafovealvision.Vision35,1679-90Karni,A.&Sagi,D.Wherepracticemakesperfectintexturediscrimination:evidenceforprimaryvisualcortexplasticity.ProcNatlAcadSciUSA88,4966-70.(1991).Shiu,L.P.&Pashler,H.Improvementinlineorientationdiscriminationisretinallylocalbutdependentoncognitiveset.PerceptPsychophys52,582-8(1992).Berardi,N.&Fiorentini,A.Interhemispherictransferofvisualinformationinhumans:spatialcharacteristics.JPhysiol384,633-47(1987).Nazir,T.A.&O'Regan,J.K.Someresultsontranslationinvarianceinthehumanvisualsystem.SpatVis5,81-100(1990).Dill,M.&Fahle,M.Theroleofvisualfieldpositioninpattern-discriminationProcBiolSci264,1031-6Dill,M.inPerceptualLearning(eds.Fahle,M.&Poggio,T.)219-231(MITPress,Cambridge,MA,USA,2002).Fiorentini,A.&Berardi,N.Learningingratingwaveformdiscrimination:specificityfororientationandspatialfrequency.VisionRes21,1149-58(1981).Schoups,A.A.&Orban,G.A.Interoculartransferinperceptuallearningofapop-outdiscriminationtask.ProcNatlAcadSciUSA93,7358-62(1996).Hubel,D.H.&Wiesel,T.N.Receptivefieldsofsingleneuronesinthecat'sstriatecortex.JPhysiol148,574-91(1959).ofmonkeys.CurrBiol5,552-63(1995).Jeter,P.E.,Dosher,B.A.,Petrov,A.&Lu,Z.-L.Taskprecisionattransferdeterminesspecificityofperceptuallearning.JournalofVision9,1-13(2009).Liu,Z.&Weinshall,D.Mechanismsofgeneralizationinperceptuallearning.Vision40,97-109Fahle,M.Perceptuallearning:acaseforearlyselection.JVis4,879-90Paz,R.,Wise,S.P.&Vaadia,E.Viewinganddoing:similarcorticalmechanismsforperceptualandmotorlearning.TrendsNeurosci27,496-503(2004).Adini,Y.,Sagi,D.&Tsodyks,M.Context-enabledlearninginthehumanvisualNature415,790-793Dosher,B.A.&Lu,Z.L.Perceptuallearningreflectsexternalnoisefilteringandinternalnoisereductionthroughchannelreweighting.ProcNatlAcadSciUSA95,13988-93Saarinen,J.&Levi,D.M.PerceptualLearninginVernierAcuity:WhatisLearned?VisionResearch35,519-527(1995).Nature402,176-8Li,R.W.,Levi,D.M.&Klein,S.A.PerceptualNature402,176-8Li,R.W.,Levi,D.M.&Klein,S.A.Perceptuallearningimprovesefficiencybyre-tuningthedecision'template'forpositiondiscrimination.NatNeurosci7,178-83(2004).Lu,Z.L.&Dosher,B.A.Perceptuallearningretunestheperceptualtemplateinfovealorientationidentification.JVis4,44-56(2004).Dosher,B.A.&Lu,Z.L.Mechanismsofperceptuallearning.VisionResearch39,3197-221.(1999).Hochstein,S.&Ahissar,M.Viewfromthetop:hierarchiesandreversehierarchiesinthevisualsystem.Neuron36,791-804(2002).Mollon,J.D.&Danilova,M.V.Threeremarksonperceptuallearning.SpatVis10,51-8.Xiao,L.Q.etal.CompleteTransferofPerceptualLearningacrossRetinalLocationsEnabledbyDoubleTraining.CurrBiol(2008).Dwyer,D.M.Perceptuallearning:completetransferacrossretinallocations.CurrBiol18,R1134-6(2008).Ahissar,M.&Hochstein,S.Attentionalcontrolofearlyperceptuallearning.ProcNatlAcadSciUSA90,5718-22(1993).Rubin,N.,Nakayama,K.&Shapley,R.Abruptlearningandretinalsizespecificityinillusory-contourperception.CurrBiol7,461-7(1997).Goldstone,R.L.Perceptuallearning.AnnuRevPsychol49,585-612Watanabe,T.,Nanez,J.E.&Sasaki,Y.Perceptuallearningwithoutperception.Nature413,844-8(2001).Karni,A.&Sagi,D.Thetimecourseoflearningavisualskill.Nature365,250-252(1993).Karni,A.,Tanne,D.,Rubenstein,B.S.,Askenasy,J.J.&Sagi,D.DependenceonREMsleepofovernightimprovementofaperceptualskill.Science265,679-82(1994).Herzog,M.H.&Fahle,M.TheroleoffeedbackinlearningavernierdiscriminationVisionRes37,2133-41Fahle,M.&Morgan,M.Notransferofperceptuallearningbetweensimilarstimuliinthesameretinalposition.CurrBiol6,292-7(1996).Crist,R.E.,Li,W.&Gilbert,C.D.Learningtosee:experienceandattentioninprimaryvisualcortex.NatNeurosci4,519-25(2001).Li,W.,Piech,V.&Gilbert,C.D.Perceptuallearningandtop-downinfluencesinprimaryvisualcortex.NatNeurosci7,651-7(2004).Seitz,A.R.etal.Task-specificdisruptionofperceptuallearning.ProcNatlAcadSciUS102,14895-900Kuai,S.G.,Zhang,J.Y.,Klein,S.A.,Levi,D.M.&Yu,C.Theessentialroleofstimulustemporalpatterninginenablingperceptuallearning.NatNeurosci8,1497-9(2005).Watanabe,T.etal.Greaterplasticityinlower-levelthanhigher-levelvisualmotionprocessinginapassiveperceptuallearningtask.NatNeurosci5,1003-9(2002).Seitz,A.R.&Watanabe,T.Psychophysics:Issubliminallearningreallypassive?422,36Butts,D.A.&Goldman,M.S.Tuningcurves,neuronalvariability,andsensoryPLoSBiol4,e92Schoups,A.,Vogels,R.,Qian,N.&Orban,G.PractisingorientationidentificationimprovesorientationcodinginV1neurons.Nature412,549-53.(2001).Ghose,G.M.,Yang,T.&Maunsell,J.H.PhysiologicalcorrelatesSchoups,A.,Vogels,R.,Qian,N.&Orban,G.PractisingorientationidentificationimprovesorientationcodinginV1neurons.Nature412,549-53.(2001).Ghose,G.M.,Yang,T.&Maunsell,J.H.PhysiologicalcorrelatesofperceptuallearninginmonkeyV1andV2.JournalofNeurophysiology87,1867-88.(2002).Yang,T.&Maunsell,J.H.TheeffectofperceptuallearningonneuronalresponsesinmonkeyvisualareaV4.JNeurosci24,1617-26(2004).Raiguel,S.,Vogels,R.,Mysore,S.G.&Orban,G.A.LearningtoseethedifferencespecificallyaltersthemostinformativeV4neurons.JNeurosci26,6589-602(2006).improvementinperceptualperformance.Science263,1289-92(1994).Rainer,G.&Miller,E.K.Effectsofvisualexperienceontherepresentationofobjectsintheprefrontalcortex.Neuron27,179-89(2000).Rainer,G.,Lee,H.&Logothetis,N.K.Theeffectoflearningonthefunctionofmonkeyextrastriatevisualcortex.PLoSBiol2,E44(2004).Law,C.T.&Gold,J.I.Neuralcorrelatesofperceptuallearninginasensory-motor,butnotasensory,corticalarea.NatureNeuroscience11,505-13(2008).Logothetis,N.K.,Pauls,J.,Bulthoff,H.H.&Poggio,T.View-dependentobjectrecognitionbymonkeys.CurrBiol4,401-14(1994).Kobatake,E.,Wang,G.&Tanaka,K.Effectsofshape-discriminationtrainingontheselectivityofinferotemporalcellsinadultmonkeys.JNeurophysiol80,324-30(1998).Baker,C.I.,Behrmann,M.&Olson,C.R.Impactoflearningonrepresentationofpartsandwholesinmonkeyinferotemporalcortex.NatNeurosci5,1210-6(2002).DiCarlo,J.J.&Maunsell,J.H.Anteriorinferotemporalneuronsofmonkeysengagedinobjectrecognitioncanbehighlysensitivetoobjectretinalposition.JNeurophysiol89,3264-78(2003).Cox,D.D.&DiCarlo,J.J.Doeslearnedshapeselectivityininferiortemporalcortexautomaticallygeneralizeacrossretinalposition?JNeurosci28,10045-55(2008).Li,N.&DiCarlo,J.J.Unsupervisednaturalexperiencerapidlyaltersinvariantobjectrepresentationinvisualcortex.Science321,1502-7(2008).Ghose,G.M.Learninginmammaliansensorycortex.CurrOpinNeurobiol14,513-8Recanzone,G.H.,Merzenich,M.M.&Jenkins,W.M.Frequencydiscriminationtrainingengagingarestrictedskinsurfaceresultsinanemergenceofacutaneousresponsezoneincorticalarea3a.JNeurophysiol67,1057-70(1992).Recanzone,G.H.,Schreiner,C.E.&Merzenich,M.M.Plasticityinthefrequencyrepresentationofprimaryauditorycortexfollowingdiscriminationtraininginadultowlmonkeys.JNeurosci13,87-103(1993).Dinse,H.R.,Ragert,P.,Pleger,B.,Schwenkreis,P.&Tegenthoff,M.Pharmacologicalmodulationofperceptuallearningandassociatedcorticalreorganization.Science301,91-4(2003).Blake,D.T.,Strata,F.,Churchland,A.K.&Merzenich,M.M.Neuralcorrelatesofinstrumentallearninginprimaryauditorycortex.ProcNatlAcadSciUSA99,10114-9Fahle,M.&Skrandies,W.Anelectrophysiologicalcorrelateoflearninginperception.GerJOphthalmol3,427-32Skrandies,W.&Fahle,M.Neurophysiologicalcorrelatesofperceptuallearninginthehumanbrain.BrainTopogr7perception.GerJOphthalmol3,427-32Skrandies,W.&Fahle,M.Neurophysiologicalcorrelatesofperceptuallearninginthehumanbrain.BrainTopogr7,163-8(1994).Shoji,H.&Skrandies,W.ERPtopographyandhumanperceptuallearningintheperipheralvisualfield.IntJPsychophysiol61,179-87(2006).Pourtois,G.,Rauss,K.S.,Vuilleumier,P.&Schwartz,S.Effectsofperceptuallearningonprimaryvisualcortexactivityinhumans.VisionRes48,55-62(2008).Skrandies,W.,Lang,G.&Jedynak,A.Sensorythresholdsandneurophysiologicalcorrelatesofhumanperceptuallearning.SpatVis9,475-89(1996).Ding,Y.,Song,Y.,Fan,S.,Qu,Z.&Chen,L.Specificityandgeneralizationofvisualperceptuallearninginhumans:anevent-relatedpotentialstudy.Neuroreport14,587-90Schiltz,C.etal.Neuronalmechanismsofperceptuallearning:changesinhumanbrainactivitywithtraininginorientationdiscrimination.Neuroimage9,46-62(1999).Schwartz,S.,Maquet,P.&Frith,C.Neuralcorrelatesofperceptuallearning:afunctionalMRIstudyofvisualtexturediscrimination.ProcNatlAcadSciUSA99,17137-42(2002).Furmanski,C.S.,Schluppeck,D.&Engel,S.A.Learningstrengthenstheresponseofprimaryvisualcortextosimplepatterns.CurrBiol14,573-8(2004).Neary,K.,Anand,S.&Hotson,J.R.Perceptuallearningoflineorientationmodifiestheeffectsoftranscranialmagneticstimulationofvisualcortex.ExpBrainRes162,23-34Mukai,I.etal.Activationsinvisualandattention-relatedareaspredictandcorrelatewiththedegreeofperceptuallearning.JNeurosci27,11401-11(2007).Yotsumoto,Y.,Watanabe,T.&Sasaki,Y.Differentdynamicsofperformanceandbrainactivationinthetimecourseofperceptuallearning.Neuron57,827-33(2008).Sigman,M.etal.Top-downreorganizationofactivityinthevisualpathwayafterlearningashapeidentificationtask.Neuron46,823-35(2005).Vaina,L.M.,Belliveau,J.W.,desRoziers,E.B.&Zeffiro,T.A.NeuralsystemsunderlyingKourtzi,Z.,Betts,L.R.,Sarkheil,P.&Welchman,A.E.Distributedneuralplasticityforshapelearninginthehumanvisualcortex.PLoSBiol3,e204(2005).Maertens,M.&Pollmann,S.fMRIrevealsacommonneuralsubstrateofillusoryandrealcontoursinV1afterperceptuallearning.JCognNeurosci17,1553-64(2005).Dolan,R.J.etal.HowthebrainlearnstoseeobjectsandfacesinanimpoverishedNature389,5