EEG Effects of Time Intervals in Natural Scene Perception

HielkePrins(intern),I.I.A.Groen(supervisor),H.S.Scholte(coaccessor) InstituteforInterdisciplinaryStudies,UniversityofAmsterdam

Abstract The way we process natural visual input has been adapted to our daily environment
through experience and evolution. Since lowlevel statistics have proven to be helpful in scene classificationandobjectidentification,moreefficientprocessingofthesestatisticsmayhavebeenthe targetoftheseadaptations.Anincreasingamountofresultsfurthermoreindicatesthatbrainplasticityis notputonholdafterearlybraindevelopmentbutcontinuestoplayaroleineverydaylearning. Encouragedbyresultsfromapreviousexperiment,wehypothesizemeasurableadaptionofthevisual mindtoourvisualenvironmentaftersinglepresentationsofnaturalstimuli.Totestthishypothesis,we measured changes in the correlation between the lowlevel statistics of the stimuli and the electroencephalographic(EEG)responseevokedinthebrain.Weinvestigatedthetimescaleofthese changes andconcludethatadaptation ofthebraintonatural imagestatistics is bestsupportedfor overnightintervals.

OUTLINE
Introduction.........................................................1 Methods...............................................................2 Results.................................................................5 Discussion...........................................................7

INTRODUCTION
Our perceptual systems are thought to have adaptedtothestatisticalpropertiesofthesignals towhichtheyareexposed,bothduringlongterm evolution and early brain development (Simoncelli & Olshausen 2001). The visual impressions of our natural environment are amongstthesignals thataremostvitalforour survival,bothasaspeciesandasanindividual. An efficient way to adapt to our daily visual environment is to exploit the most important featuresofencounterednaturalstimulitoguide furtherprocessingasearlyaspossible (Geisler 2008).Lowlevelstatisticsofnaturalstimulican provide important clues about their complexity and texture similarity without the need for specificobjectidentificationprocesses.

Inlinewiththisidea,ithasbeenshown(Scholte etal.2009) thatdeflectionsintheearlyevent related potential (ERP) correlate strongly with theparametersofaWeibulldistributionthatisfit totheedgehistogramsofnaturalimages.Edge histograms are a representation of the contrast distributions in visual scenes. The two free parameters of a fitted Weibull distribution (named beta and gamma) seem to describe a structured space with clearcut single objects against an empty background in the lowerleft quadrantandclutteredtexturelikepicturesinthe upperrightone(Figure1). The correlation between early ERP and the Weibullstatisticsofstimulihasbeenexploitedto accuratelypredictwhichofthepresentedscenes participantswereprocessingduringaparticular trial (Ghebreab et al. 2009). These predictions werebasedontheoutcomesofacomputational model of the output of the lateral geniculate nucleus (LGN) to the early visual cortex. Comparison between predictions made by this model and the participant's ERP response revealed a clear effect of the number of presentations per stimulus on the explained variation in early deflections in the ERP. This 1/10

& DiCarlo 2006) and may involve various mechanisms ondifferenttimescales (C.Gilbert 1994;C.D.Gilbertetal.2001). Since Ghrebreab et al. (unpublished) did not control the interval between successive repetitionsofthestimuli,itisdifficulttoderive conclusionsabouttheunderlyingmechanismsof observedadaptationsfromtheirresults.Inorder tofindthemostimportantofthesemechanisms, wedesignedanexperimenttodetermineatwhat timescalesadaptionofthebraintonaturalstimuli is most apparent. Concretely, the aim of the currentexperimentistoinvestigatetheamount ofincreaseinexplainedvarianceoversuccessive repetitions while manipulating the interval betweenthem. Constraintsintheirexperimentaldesignlimitthe interstimulus intervals to a maximum of approximately 30 minutes. Since Ghrebreab foundasignificantincreaseinexplainedvariance forrepetitionswithinthisinterval,weexpected to find a similar effect for the smallest of our interstimulusintervals.However,ourresultshint atanadaptationthatismostpronouncedwhen theinterstimulusintervalspansafullday.

Figure 1:Therelationbetweenclutterinnaturalscenes and their graylevel Weibull statistics. The amount of clutterinthescenescorrespondwiththewidthandshape of the corresponding Weibull fit on their contrast histogramsWidthandshapeofWeibulldistributionsare determinedbytheirbetaandgammaparameters(a). BetaandgammaparametersofaWeibulldistributionform atwodimensionalspacewithpicturesofclearcutobjects againstanuniformbackgroundinthelowerleftquadrant and more texturelike scenes in the topright one (b). FigurecopiedfromGroenetal.(submitted).

METHODS
Stimuli Atotalof1600imagesofnaturalsceneswhere selected for the experiment. The images had a resolution640x480pixels.Thesetwasdivided into 4 groups of 400 images with an equal standarddeviationinbetagammaspace.Within eachofthesesets200imagesshowedmanmade environments (city landscapes and vehicles) whereasthe200otheroneswherepicturesfrom natural environments (natural landscapes and animals). Figure2showsthedistributionofstimuliinthe foursetsinbetagammaspace.Picturesofman made environments (red circles) had a lower average gamma value ( =0.9877 , 2/10

mayindicateadaptationofthebraintothelow level statistics of the stimuli (Ghrebreab et al., unpublished). A growing body of evidence supports a continuing role for brainplasticity during perceptual learning in adult human brains (Karmarkar&Dan2006),includingshortterm adaptiontonaturalimagestatistics(Schwarzkopf et al. 2009). Adaption seems to take place throughout the whole process of target recognitioninclutterednaturalscenes (Kourtzi

Experimental design Four different interval conditions were defined ranging from relatively small interstimulus intervals to overnight rehearsal (Table 1). The foursetsof400stimuliwhereeachassignedto one of these conditions. The stimuli where offeredinblocks of40images takenfromthe sameset(Figure3). Distributionofthestimuliblockswithineachof thesessionswascounterbalancedoverduration ofthesessionsbytestingforequalvariancein the block distributions across the interval conditions. We did this by generating random block designs whose maximal block intervals mettheconstraintsinTable1,untilanumberof 32designsexceededathresholdof p=0.95 on aBarlett'stest.Withineachblockthe40images werepresentedinrandomorder.
Set RepetitionInterval I 48minutes II 1220minutes III >30minutes IV 2324hours NumberofStimuli 2*200 2*200 2*200 400

Figure2:Distributionoftheimagesinbetagammaspace foreachofthefourstimulisets.Stimulishowingnatural environments are shown in green, stimuli of manmade environmentsinred.

=5.719410 ) than those of natural ones (green circles, =1.1883 , =0.5447104 ). Thetwotypesdonotdifferinaveragebetavalue ( =0.013 )althoughdistributionofthesevalues differsslightlybetweenmanmade( =0.1951 ) andnaturalenvironments( =0.2495 ).
4

Determining the Weibull values Coordinates ofthestimuliplotin Figure2 are determinedbyaweightedsumofthebetaand gamma values for three different color layers (graylevel 0.5, redgreen 0.25, blueyellow 0.25). A three parameter Weibull distribution (Formula1)wasfitonthecontrastdistributions intheselayers. p f = ce
f

Table1:Conditionsandthenumberofuniquestimuli per condition. Stimuli in the first three sets are repeatedduringasinglesessionanddifferbetween thetwosubsequentsessions.Thefourthsetcontains the same stimuli during both sessions, effectively creatingadelayofapproximately24hours.

(1)

Parameter c is a normalization constant converting the frequency distribution in a probabilitydistribution.Parameter represents theoriginofthedistributionandisnormalized out by subtracting the lowest contrast value in eachpicture.Remainingfreeparameters,thebeta and gamma values, determine the scale and shapeoftheWeibulldistribution (Scholteetal. 2009).

After every three blocks participants were allowed to have a break. They performed 2 sessionsof1600trialsin2subsequentdays,at approximately the same time of the day. Participants wereassignedtwoofthebalanced designs,oneforeachofthesesession. Participants A group of 16 participants (2044 years) were recruited to participate in the experiment and dividedin4groupsof4participants.Eachgroup was assigned a different set of stimuli per interval condition. Participants gave written informedconsentpriortoparticipationandthe 3/10

Figure 3:Exampleofabalanceddistributionofstimuli blocksthroughoutasessionforeachoftheconditionsin Table1.Ineachrow(intervalcondition),stimuliblocks containingthesamestimulihavethesamecolor.

100to500millisecondtimeepochwithregard tostimulusonset.Thefirst100millisecondsof thisinterval(100to0ms)wastakenasbaseline activity and subtracted from all trials. Ocular correction was applied using an offline algorithm (Gratton et al. 1983). Data were re referencedtotheaverageactivityrecordedatthe earlobes. Trialsinwhichblinksweredetectedwithinthe 50 to 300 millisecond interval were excluded fromanalysis,as participantsmayhavemissed thestimulus.Additionally,trialswithanabsolute amplitude exceeding the 250 V or 250 V range,andtrialswithanamplitudeofmorethen twotimesthestandarddeviation,wereleftoutof theanalysis. The data was currentsourcedensity (CSD) transformedbeforeanalysis.CSDtransformacts as a highpass spatial filter, effectively subtracting volumeconducted common activity duetodistantsources (Kayser&Tenke2006). CSDthusresultsinamorespecificattributionof electroencephalographic activity to nearby electrodes. Analysis of explained variance Reportedexplainedvarianceisalwaysobtained bycomparingtheamplitudefrompreprocessed singletrial ERP's at each time point, with the beta and gamma values from the stimuli presentedinthecorrespondingtrials. Dependingonthepurposeoftheanalysis,ERP's for the comparison are taken from a single subject or averaged over multiple participants. Furthermore, the amount of trials taken into account naturally differs between the overall analysisandtherepetitioncontrastsperinterval condition. The explained variances per interval condition arebasedontheERPamplitudeaveragedover the four participants that were presented the samestimuliineachcondition.Foreachchannel andeachtimepointwithinthefourconditions, estimationofalinearregressionmodelresultsin 4/10

experiment was approved by the local ethical committee. Task Participants had to categorize the stimuli as either manmade or natural, using buttons attachedtotheleftandrightarmsoftheirchair. Orderofthebuttonswascounterbalancedacross participants. Participants were instructed to answerasquickaspossibleandaskedtotrynot tomissanyofthestimuli. Stimuli presentation Trial duration varied between 1495 and 1995 milliseconds,takenfromauniformdistribution. Atthebeginningofeachtrial,afixationcross wasdisplayedfor100milliseconds.Afterwards thestimuliwhereverybrieflypresentedatthe centerofthescreen,for100milliseconds.During theremainingpartofthetrialthefixationcross returned and participants could respond by pressingoneofthebuttons. EEG recording and preprocessing EEG data were recorded at 256Hz using a Biosemi ActiveTwo amplifier (Biosemi Instrumentation, Amsterdam, The Netherlands) from 64 scalp electrodes and 4 perioccular electrodes. Scalp electrodes were positioned accordingtoastandard1020electrodelayout. A50Hznotchfilter,a0.1Hzlowcutoffanda 30Hzhighcutofffilter,wereappliedtothedata using Brain Vision Analyzer (Brainproducts, Munich,Germany).ERP's werecomputedover

Figure4:Timecourseoftheexplainedvarianceacrossall interval conditions for the two presentation separately (left)andtheincreaseduringthesecondrelativetothe firstpresentation.

Figure5:Distributionofmaximalexplainedvarianceover the scalp electrodes between 100 and 160 milliseconds, acrossconditions. Maximalexplainedvarianceforboth presentations show foci around occipital and parietal electrodes(left).Differencesinexplainedvarianceforthe secondrelativetothefirstpresentationareminor(right).

asinglemeasureofexplainedvariance, R2 ,for allremainingstimuliassignedtothatcondition. Where R2 resembles the goodness of fit betweenthemeasuredEEGamplitudesandthe two independent parameters of the model, the betaandgammavaluesofthestimuli. Reportedstatisticalresults arealwaysbasedon trials from individual participants. The ttests performed, compare distributions of individual measuresofthemodelfit R2 betweenthefirst andsecondpresentation. We expected to find the largest differences in explained variance for a group of 7 occipital electrodes(PO3,POz,PO4,O1,Oz,O2andIz) because of their proximity to primary visual regionsofthecortex.

160millisecondinterval. The differences in total explained variance betweenthefirstandsecondsecondpresentation overallelectrodesareminor.Differencesin R2 range from 0.040 at P10 to 0.029 at CP6. Occipital electrodes are no exception with a rangefrom0.014atOzto0.006atO2(Table2). Largest explained variance for the first ( R2 = 0.5843)andsecond( R2 =0.5732)presentation, over all stimuli is found outside the region of interestatCP4.
Electrode Oz POz PO4 Iz

R2first
0.437 0.384 0.100 0.196 0.364 0.102 0.286

2 R2 second R

t 15 p
2.45 1.32 0.29 0.81 2.03 1.57 0.84 0.986 0.896 0.610 0.785 0.970 0.931 0.792

0.424 0.371 0.086 0.189 0.364 0.104 0.292

0.014 0.013 0.014 0.008 0.000 0.002 0.006

RESULTS
Repetition contrasts across conditions AnalysisacrossconditionswasbasedontheERP overall16participants.AmplitudeoftheERP's was correlated with Weibull statistics from all 1600 stimuli for each one of the two presentationsseparately. The timecourse of the correlation shows a significant peak between 100 and 160 milliseconds (Figure 1). Spatial distribution of peakactivityoverthescalpelectrodesshowsfoci of explained variance around occipital and parietal electrodes (Figure 5). Shown is the maximal explained variance within the 100 to

O1 PO3 O2

Table 2:Explainedvarianceforthesevenoccipital electrodes of interest (first two columns) and their increase over repetitions (third column). Last two columnsreporttheresultsofattestcomparingthe distribution of individual explained variances betweenthetworepetitions.

Totestthesignificanceoftheeffectofasecond presentation for each of the electrodes, we performedapairedttestbetweenthemaximum explained variances within the 100 to 160 millisecondintervalforthetworepetitionsfrom all 16 participants. None of the electrodes 5/10

Figure 6:VarianceinEEGamplitudeexplainedbythebetaandgammaparametersateachtimepointintthetrials.Each coloredlinedepictsregressionresultsforadifferentelectrode.Rowsarethefourdifferentintervalconditions(Table1).Plots foreachconditionarebasedontheaverageERPforagroupof4participants.Firsttwocolumnsshow R2 forthefirstand secondpresentationofthestimuli.Thirdcolumnshowsthedifferenceinexplainedvariancebetweenthetworepetitions.

supportedanincreaseofexplainedvariancewith the number of presentations ( =0.05 ), when disregardingtheintervalcondition.. Repetition contrasts per interval Analysis of all stimuli in the four different intervalconditions simultaneouslymayobscure aneffectofrepetitionwhenimplicitlearningof the contrast distributions takes place only at someofthestudiedtimescales. Wethereforerepeatedtheanalysisofrepetition contrastsperintervalcondition(Figure6).Due tothesmalleramountofobservations,regression analysisperintervalconditionyieldslowerpeak valuesthentheanalysisacrossconditions. Location of the electrodes whose increase in explained variance exceeds the significance threshold( =0.05 )areshownin Figure7.All testsarebasedontheexplainedvarianceforthe first and second presentations of all 16 participants,regardlessofthestimulitheywere

presented within each of the four interval conditions. Forshortstimulusintervals,explaineddifference increasessignificantlyforelectrodesF4,t(15)= 1.98,p=0.033,andPO7,t(15)=2.48,p=0.013. Explainedvariancefortheintermediateandlong intervals does not significantly increase over conditions. Largestincreaseinmaximumexplainedvariance, intermsofthenumberofelectrodesthatsurpass thettestandinrelativeincreaseof R2 ,takes
Figure7:Electrodesfor which a paired ttest supports a larger maximum explained variance during the second than during the first presentation. Peak explained variance is taken within the 100 160msinterval.

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DISCUSSION
Summary of the results Analysis across intervalconditions provides no support for an increase in explained variance between subsequent presentations of the same stimuli. Within the four different interval conditions, the short and overnight conditions yieldsignificantresults.Theovernightcondition supports an increase of explained variance for oneofoccipitalelectrodesofinterest. Within session repetition Differences inexplained variance between first and second presentations for the two shortest interval conditions are minor at the occipital electrodes(Figure9).Significantincreaseswere onlyfoundoutsidetheregionofinterestandonly fortheshortestintervalcondition. Previous research by Ghebreab et al. (unpublished)indicatesthataconsistenteffectof therepeatedexposureontheexplainedvariance does exist for these intervals. A considerable increase in maximum explained variance ( R20.08 )hasbeenfoundbetweenthefirst andsecondpresentation. The maximum explained variance for both presentations varies considerably between participants(Figure10).Furthermore,thestimuli presentedperconditionweredeliberatelyvaried betweengroupsofparticipantsinordertocontrol forstimulidependenteffects.Lackofstatistical powerduetothesmallsamplesizepercondition couldbethereasonthatwefailedtoconfirmthe effect of rehearsal within the two shortest intervalconditions. Forthelongcondition,visualinspectionofthe difference in explained variance between presentationsattheoccipitalelectrodes,indicates a decrease rather than an increase during the secondpresentation(Figure9).Withaminimum interstimulus interval of 30 minutes the long condition will be most affected by decreased attentionduetofatigue. 7/10

Figure 8: Timecourse of the difference in explained variance between second and first presentation of the stimuliintheovernightcondition.

placeintheovernightcondition.Theovernight condition shows a significant increase in maximum explained variance at one of the occipitalelectrodesofinterest(Figure9). Electrode O2 shows an increase during the second(M=0.012,SD=0.0012)asopposedto thefirstpresentation(M=0.015,SD=0.001) thatisfoundsignificant,t(15)=1.80,p=0.046. Three other electrodes show a significant increase in peak explained variance within the 100 to 160 millisecond interval: AF4, t(15) = 1.82,p=0.045,AF8,t(15)=2.32,p=0.017,and T8,t(15)=1.85,p=0.042. Time-course of increase in overnight condition Thespatiotemporaldistributionoftheincrease inexplainedvariancefortheovernightcondition during the 100 to 160 milliseconds interval, revealsafocusofsustainedactivityaroundone oftheoccipitalelectrodes(Figure7). Anincreaseandasubsequentdeclineinrelative explainedvarianceatPOzisclearlyobservable during a relatively long continuous interval. Averaged increase in explained variance over repetitions is significant between 108 and 128 millisecondsafterstimuluspresentation,t(15)= 1.9463,p=0.0353.

These findings may therefore reflect false discoveries. Because of the large number of simultaneous tests, correction for the false discovery rate is appropriate. Our results did not survive correctionformultiplecomparisonsbutanotable increaseintheovernightconditionisconsistent withtheliterature. Overnight rehearsal Theresultssupportaneffectofstimulirepetition for the overnight condition. Average maximal explained variance during the 100 to 160 millisecondspoststimulusinterval,increasesfor mostoftheoccipitalelectrodes(Figure9).More centraloccipitalelectrodes(POz,Oz,O2)showa relativelylargeincreasebetweenrepetitions.For one of these electrodes (POz) the increase in explained variance per time point is found significant during a continuous period of 20 milliseconds. Improving performance in a visual discrimination task is reported to require sleep within 30 hours of training (Stickgold et al. 2000). Selective increase ofexplained variance between repetitions within the overnight condition may indicate that sleep is likewise required for implicit learning of feature distributionsinnaturalstimuli.
Figure10:Distributionoftheexplainedvariancebasedon all 16 participants during first (blue) and second (red) presentationperelectrode.

Figure 9: Mean explained variance for the occipital electrodes of interest during the first (blue) and second (red)presentationofthesamestimuli.

Statistical power and false discoveries Activity measured at frontal and temporal electrodes that show a significant increase, is unlikelytodirectlyreflectprocessingofstimuli basedontheircontrastvalues. Although other processes may differ between firstandsecondpresentationitishardtoseewhy these would correlate with beta and gamma parameters of the contrast distributions in the stimuli,especiallywithinthetimewindowof100 to160millisecondsafterstimuluspresentation.

Evaluation of the results Somecautionisrequiredwheninterpretingthe difference in maximum explained variance betweenrepetitions,aspeaklatenciesmayvary betweenparticipantsandbetweenrepetitions. Oursampleincludesparticipantswithrelatively low values of maximal explained variance (Figure10).Significanceofincreaseinmaximal explained variance, may therefore reflect arbitrary fluctuations over time rather then increasing correlation with the imagestatistics duringprocessingofthestimuli. Despite these remarks, selective increase in explainedvariancefortheovernightconditionat 8/10

multiple occipital electrodes is a remarkable result.Thecomparisonbetweenthefirstandthe second repetition over a longer continuous interval (Figure 8) complements the evidence fromdifferencesinmaximalexplainedvariance, andleadstothesameconclusion. Suggestions for further analysis Although our preliminary results are indicative fortheovernightcondition, analternativepath of analysis may strengthen the support for an increaseinexplainedvarianceoverrepetitions. Ourtrialrejectioncriteriawereconservativein order to preserve as many observations as possiblefortheregressionanalysisandtokeep thebalancedexperimentaldesignintact.Whena lateorcompletelackofresponseindicatesalack of attention, exclusion of trials without a responsemayincreasethesignaltonoiseratio. Despiterelativelylargeamountofmissedtrials for some of the participants, none of these participants nor the missed trials have been excludedfromoursample. For the purpose of the current analysis the discriminationtaskwasincludedtoensurethat participantspayedattentiontothestimulibutis otherwise considered irrelevant. Behavioral results were not taken into account during the analysis. However, given the large individual differences in maximum explained variance, analysisofgoodandbadperformersseparately maybeinformative. Additional support for exclusion of bad performers comes from the results of previous researchinthefrequencydomain.Theamplitude ofoscillationsinthealphabandareshowntobe related to a decrease in performance on visual discrimination task (Hanslmayr et al. 2005). Good performers furthermore showed a significantlylargerP1,adeflectioninearlyERP that is presumably related to our regression analysis of the ERP amplitude and stimuli statisticsinthe100to160millisecondinterval. The same study relates visual discrimination

performancewithincreasedphaselockinginthe alpha band suggesting that related cortical activity may be better coordinated in time (Hanslmayr et al. 2005). In addition, spatial frequencies in stimuli have been shown to differentially affect the EEG alpha and evoked gammaresponses (Frndetal.2007).Evidence frommacaquestudiessuggestthatthedecrease in gamma responses with spatial frequency originatesintheLGN(Tootelletal.1988). Analysis of synchrony between different electrodesorfrequencybands,maythusreveal adaptationsoverrepetitionsthatgoundetectedin the current analysis. Additionally, trialtotrial differencesinstimuluslockedphaseonsetsmay provide new insights in temporal differences between brain responses evoked by subsequent repetitions. Duration of the breaks between blocks varies withinandbetweenparticipants.Forsomeofthe participants durations may also vary between conditions. Resulting variations between block intervals cause variation in intervals between repeated stimuli. Outliers with significantly largerinterstimuliintervalsthanintendedbyour designarecurrentlynotrejectedbeforeanalysis. Weanalyzedtheeffectsofvaryinginterstimuli intervals within bins of considerable width. Especiallywithintheshorterintervalconditions, anyobservedincreaseinexplainedvarianceover repetitionsmaydependonrapidadaptiontothe stimulithatisnotequallydistributedoverthese bins.Includingtheindividualintervalsbetween pairsofidenticalstimuliinourregressionmodel maythereforeprovidedifferentresults.

CONCLUSION
Contrary to previous research we did not find convincing statistical support for increased adaptiontocontrastdistributionsinimagesover repetitions of stimuli. Although not significant for our sample, indications that such an effect 9/10

existsinthepopulationfortheovernightinterval condition are observable in the data. Further analysis and future research with increased statisticalpowerisnecessarytoreachadefinite conclusion.

DependentPlasticityinAdultVisualCortex. Neuron,52(4),pp.577585. Kayser, J. & Tenke, C.E., 2006. Principal componentsanalysisofLaplacianwaveforms as a generic method for identifying ERP generatorpatterns:I.Evaluationwithauditory oddballtasks. ClinicalNeurophysiology,117, pp.348368. Kourtzi,Z.&DiCarlo,J.J.,2006.Learningand neuralplasticityinvisualobjectrecognition. Current Opinion in Neurobiology, 16(2), pp.152158. Scholte, H.S. et al., 2009. Brain responses strongly correlate with Weibull image statistics when processing natural images. JournalofVision,9(4). Schwarzkopf, D.S., Zhang, J. & Kourtzi, Z., 2009.FlexibleLearningofNaturalStatistics in the Human Brain. Journal of Neurophysiology,102(3),pp.18541867. Simoncelli, E.P. & Olshausen, B.A., 2001. Natural image statistics and neural representation. Annual Review of Neuroscience,24,pp.11931216. Stickgold,R.,James,L.&Hobson,J.A.,2000. Visualdiscriminationlearningrequiressleep after training. Nat Neurosci, 3(12), pp.1237 1238. Tootell, R. et al., 1988. Functional anatomy of macaquestriatecortex.V.Spatialfrequency. The Journal of Neuroscience, 8(5), pp.1610 1624.

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