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    THE EFFECTOF THE GEOMETRICFORMAND MEANING

    OFTHE STIMULUS ON THE CONFIGURATION OF THE

    VISUALEVOKED RESPONSE

    by

    SHERRILL JEANSWIFT PURVES

    B.Sc, McGill University,1967

    M.D. UniversityofB r i t i s h Columbia,1971

    A THESISSUBMITTED INPARTIAL FULFILMENTOF

    THEREQUIREMENTS FOR THE DEGREE OF

    DOCTOROF PHILOSOPHY

    i n

    THEDEPARTMENT OF PHYSIOLOGY

    Weaccept this thesisascxsnforining

    totherequired standard

    THE UNIVERSITYOFBRITISHCOLUMBIA

    May,1976

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    In pre sen t in g th is th es is in p ar t ia l fu l f i l me nt of the requirements for

    an advanced degree at the U n i v e r s i t y of B r i t i s h Colum bia, I agree that

    the Li br ar y sh a ll make it fr ee ly a va il a bl e for r efere nce and study.

    I fu rt he r agree tha t perm iss io n for ext ens ive copying of thi s t he s i s

    f o r s c h o l a r l y purp oses may be gr an te d by the Head of my Department or

    by hi s re pr es en ta t i ve s . I t is understood that copying or pu bl ic at io n

    of t hi s th es is for f i n a n c i a l gai n sh al l not be allo wed withou t my

    w r i t t e n p e r m i s s i on .

    Depa rtment

    The U n i v e r s i t y of Br i t ish Columbia

    2075 Wesbrook Place

    Vancouver, Canada

    V6T 1W5

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    ABSTRACT

    Inthevisual system i t has been established that certain physical

    parameters of thestimulusincluding intensity, focus,and sizeof

    theelements of apatternhavesignificant effectson the configuration

    ofthe evoked response as recorded from theoccipital region. While

    claimshavebeenmadei n a fewpublications,tothisdate thequestion

    ofwhether anypartof the evoked response i saffectedby higher

    perceptualprocessesratherthan onesmoredirectly related tostimulus

    inputhas not been resolved.

    Evoked responses tofourgeometric shapes (a square,c i r c l e , e l and

    omega)wererecorded frommultiple scalp locationsunder two experimental

    conditions; i n the f i r s tthe shapeswerepresented atrandom intervals

    and inrandomorder with nomeaningassigned tothem. In the second

    theywerepresented i n a fixedrhythmic sequence, and two of the shapes

    occasionally failed to appear intheir usualtime interval i n the

    sequence. The subject"wasrequiredtorapidly signal such omissions by

    a buttonpress. The cerebral electrical activityduring this time

    intervalof the expected, but omittedstimuluswas recorded and

    separatelyaveraged, and these responseswere called emittedpotentia ls.

    Thereweresignificant,differencesbetweenthe evoked responses (in the

    occipital region) to the square and e l shapes andbetweenthose to the

    c i r c l e andomegashapes. These differencesweredemonstrated by three

    measurementtechniques: performance of thediscriminant functions

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    computed bySWDAi ncla ssi fyi ng single t r i a l responses, aratio

    s t a t i s t i c calledXasdescribedbyJohn,Herrington&Sutton (1967),

    and amplitude differences i n the N 2 and P 2 components.

    Theemitted potentials recorded fromthevertex, containedasmall

    and variable early negativecomponent and anobvious late positive

    component similarto the oneseeni n theresponses to thepresent square

    ande lduringthesecondexperimental condition. Manipulationof the

    waveformsfromthesingle t r i a l s priortoaveraging showed that this

    negativecomponent was nottime-lockedto the peak.

    Theevokedpotential differences foundwere believed to be due to

    twoclasses ofvariables; thephysical char acteristi csof thestimulus

    (the contoursin thecentral1.5 of thevisual field)and task-related

    changesi n the meaning of thestimulus; andthese affected earlierand

    laterpartsof the waveform respectively.

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    TABLE OFCONTENTS:

    Page

    t

    Abstract i i

    TableofContents . i v

    L i s tofFigures v i i

    L i s tofTables v i i i

    L i s tofAbbreviations i x

    Acknowledgement x

    I. Introduction 1

    II. Review of theLiterature 5

    2.1 Therelationshipofscalp-recordedevokedresponsesto theunderlying brai na c t i v i t y 5

    2.2 PatternVER's i n thestudyofvision 7

    2.3 Evokedresponsesi n thestudyofpsychologicalvariablesand meaning 14

    2.4 Suitniaryof theliterature review 20

    III. Statementof theproblemf orinvestigation 22

    IV. Methods 24

    4.1 Dataacquisitionmethods 24

    4.2 Dataanalysismethods 33

    V. Results 38

    5.1 Effectofdifferent experimental conditions

    ontheevokedresponse 38

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    5.2 Effectofdifferentstimulus shapes

    ontheevoked responses 42

    5.2.1 ResultsofVisu al analysis 43

    5.2.2 ResultsofStepwise discriminantanalysis 50

    5.2.3 ResultsusingtheXdescript or

    computations 60

    5.2.4 Summary of theeffecto f the

    different stimulus shapeson theevoked responses 64

    5.3 Emitted responses 67

    5.3.1 Averaged emitted responses 67

    5.3.2 Averaged "shifted"emitted responses 72

    5.3.3 Control studies: motor potentials,

    EOGcontributions,andtopography . . . 76

    5.3.4 Summary offindingsfor omitted

    stimuli 79

    VI. Discussion 81

    6.1 Studiesofevoked responsesandstimulusmeaning 81

    6.2 Techniques fortheanalysisofevokedresponses 83

    6.2.1 Visu al analysi s 85

    6.2.2 Xvalues 85

    6.2.3 Stepwisediscriminantanalysis 86

    6.3 Topographicaldistributionof theevoked

    responses 89

    6.3.1 Locationofshape effects 89

    6.3.2 Distributionofresponsecomponents.. 90

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    v i

    6.4 Intershape differences related to physical

    propertiesof the stimuli 93

    6.5 Effectsof stimulusmeaning 99

    6.5.1 Present stimuli 99

    6.5.2 Omitted stimuli 101

    VII Summary& Conclusions 105

    Bibliography 110

    AppendixA. Computerhardware and data

    acquisitionprogram (VER03) 118

    AppendixB. Stepwise Discriminant Analysis 120

    AppendixC. Termsfrom experimental psychology .. 124

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    vii.

    LISTOF FIGURES

    Figure Page

    1. Electrodeplacements 25

    2. Dataacquisitionsystem 27

    3. Stimulusshapes 29

    4. Experimentalparadigms 30

    5. Effectofexperimental conditions forcircleand omega . 39

    6. Effectofexperimental conditions forsquareande l.... 41

    7. Effectofstimulusshapeon theevokedresponses 44

    8. Intersubjectvariability; occipitalVER toe l for

    13 subjects 45

    9. Effectofstimulus size 52

    10. Distributionoflatency pointschosenby the4-groupdiscriminant functions 57

    11. Distributionoflatency pointschosenby the2-groupdiscriminant functions 62

    12. DistributionofXvalues 65

    13. Averagedemitted potentialsat C infivesubjects 68

    14. Averagedemitted potentialsatmultiple recording sites

    inonesubject 69

    15. "Shifted"averagedemitted potentialsat C zinfivesubjects 73

    16. "Shifted"averagedemitted potentialsatmultiplerecordingsitesin twosubjects 74

    17. Responsestoomitted stimuliwithEOGcontrol 77

    18. Responsestoomitted stimuliinfrontal regions withabsentbutton press 78

    19. Flowchartof VER03program(AppendixA) 119

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    v i i i

    LISTOFTABLES:

    TABLE

    I. Latencies of thecomponentso f occipital

    evokedresponses to the fourshapes 48

    II. Amplitudesof thecomponentsof occipital

    evokedresponses to the fourshapes 50

    III. Performanceof the "four-shape" discrinunant

    functions 54

    IV. Percentagesof t r i a l s incorrectly classified

    separated in to pa ir s ofshapesconfused witheachother 56

    V. Performanceof the "two-shape"discriminant

    functions.One subject, five pairs ofshapes 59

    VI. Performanceof the "two-shape" discrinunantfunctions. Fiv e subjects, two pai rs of

    shapes 61

    VII. Mean A values foreachsubject 63

    VIII. Amplitude and latency of P^ forresponses

    to omitted and present stimuli 70

    IX. Amplitude and latency of N 2 forresponses

    to omitted and present stimuli 71

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    LISTOF ABBREVIATIONS

    EEG Electroencephalogram

    EOG Electro-oculogram

    EP Evoked potential

    ER Evoked response

    SWDA Stepwise Discriminant Analys

    VER VisualEvokedResponse

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    X

    The author gratefullyacknowledgesthe assistance inthis research

    workprovided by anumberof people, especially

    Dr.MortLow whoseadvice,encouragement,and t i r e l e s s personal

    involvement i n the projectwereinvaluable fo r me

    MichaelBakerfor hisinterest and a b i l i t y i n the design and writ ing

    ofthecomputerprogramsf or thePDP-11system

    JaniceGallowayand RichardFergusonfortheir technicalassistance

    with the experimental apparatus and the subjects

    JimMcEwenand JohnDoylefromtheDept.of E l e c t r i c a l Engineering

    for their helpful instructioni n the use of the UBC Computing

    Centre F a c i l i t i e s

    Dr. Michael SchulzerfromtheDept.ofMathematicsfor advice on

    s t a t i s t i c a l testing and

    my committeemembersi n theDept.of Physiology; Drs.Tony Pearson,

    HughMcLennanand PeterVaughanfortheir consideration, suggestions

    and criticismso f the pro jec t.

    Financial support was provided by the Medical ResearchCouncil of

    Canadathrough an MRC Fellowship to the author fromSeptember 1972

    u n t i lDecember1975 and MRC GrantMT-3313.

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    Chapter I

    INTRODUCTION

    To attempt tounderstandtheenormouscomplexityof neural transactions

    within thehumanbra in i s a formidable challenge to invest igato rs i n a l l

    of the disciplines of neu rol ogi cal research. One of themethodsof

    directly studying the neural processes i n the brai n, i s to record the

    electrical activity occurring inresponse tosensory stimu lat ion . Two

    classes of electrical activity can be recorded; theminutely localized

    potentials of singlenerve c e l l s and themoredi ff us e and slowly changing

    potential fieldsdetectable i n the v i c i n i t y of lar ge populations of

    neurons. The terms"evokedpot ent ial s" and "evokedresponses" are

    coimonlyusedtodenotethe sig nal s of the latter class of cerebral

    e l e c t r i c a l a c t i v i t y that are recorded followin g stim ulation of sensory

    receptors or afferentpathways.

    A stimulus i n i t i a t e s asequenceof physiologicalevents,whichare the

    substrates fo r i t s perception, as well as processes leading to an overt

    behavioral response. Therefore, analys is of the e l e c t r i c a l activity

    occurringbetween stimulus and response should provide clues to the

    natureand anatomicalloca tion ofthese physiologicalevents.

    Since the late1950's, there has beena largeamountof data published

    aboutthe eff ec t of awidevar iet y of stimulus parameters on the evoked

    responsesrecorded fromhumansubjects. The stimulus parameters

    investigatedhave included bothphysical characteristicssuchas intensity,

    1

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    2.

    durationandsize,andpsychological-contextualonessuchasexpectation,

    task-relevance andstimulusmeaning. Withrespectto thelat ter,

    conflicting reportshaveappearedi n theliterature. Inspiteofthe ir

    importance forthedevelopmentoftheoriesofinformation processing,

    somewidely publ iciz ed findingson theeffectofstimulusmeaningon the

    evokedpote ntia l configurationhavebeen poorly substantiated.

    Theuse ofpatterned visual stimulitoe l i c i t evokedresponsesi s arecent

    developmenti nthis f i e l d ofstudy, becauseof therelativetechni cal

    complexity of thestimulus presentationapparatus (ascomparedto the

    simple stroboscopic flash thatwasusedpreviouslytostimulatethevis ual

    system). Withpatterned stimulitheexperimentercanprovokeandexamine

    the physiologicaleventsinvolvingmorecomplexaspectsofvisu al

    perceptionandrela ted cogn itive processes thanwasformerly possible

    using onlytheflas h.

    Anotherrecentadvancei n thestudyofcogn itiv e processes rel ate dto

    processingofsensory information, appearsi n somerecent reportsof the

    recordingof aconsistentwaveform fromthescalpi n theintervalwhen a

    subjectexpectsbuti snotac tu al ly presented withastimulus. The

    responsesrecorded i n theintervalwhen nostimulushasbeen presented

    are called emitted potentials,todistinguishthemfromtheevoked

    potentials e l i c i t e d bystimuli thatareact ua ll y presented. Thestudyof

    these emitted potentialwaveformsmayprovide important information

    concerning bra in processes without inte rferen cefromtheactivity directly

    relatedto thestimulus input.

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    3.

    In recordingsmadefrom thescalp, the evoked response i s ofmuchsmaller

    amplitude (l-20yv) than the ongoingelectroencephalographic (EEG)

    activitywhich i s 10 - 100 yv i n amplitude. In order to isolate and

    studythe activity relatedto the stimulus, repeated responses are averaged

    startingat the time of stimuluspresentation. The event-related activity

    which, i spresumedt o be relatively constant in the repeated t r i a l si s

    enhancedand the background non-event related EEG i scancelledby this

    method. Thenumberof t r i a l sper average hasvariedwidely indiffer ent

    reports,ranging from 10 to2000ormorerepetitionsof the stimulus.

    The variance of these t r i a l s included i n the average i srarelyever recorded.

    The resulting averaged evoked responses can beexamined. Thepeaksare

    labelledand their latencyand amplitude can bemeasured. Whensimple

    stimulusvariablessuch as luminance arealtered,theeffect i susual ly

    only seen i n one o f thepeaksorcomponentsof the evoked response and i s

    f a i r l yeasy to observe andmeasure. Howeverwhenchangesaremadei n

    complexstimulusvariablessuch asmeaning,morethan onecomponentin

    the waveformmay be alteredand i tbecomesquite d i f f i c u l t toquantify

    differencesi n thewaveformsthatcan be relatedto stimulusvariable s.

    Ifthe evoked EEG a c t i v i t y i sdigitizedand thentreatedas aseriesof

    discreetobservations,somequite differentmethods,other than averaging

    can be used to approach thisproblem. One suchmethodi scall ed

    discriminant analysis; i t i s a technique based onmultivariate statistical

    theories thatdetermines thevariables (or time points)thatbest account

    for thedifferencesbetweentwo ormoresetsof observations (waveforms).

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    4.

    It requires verycomplexcomputationsand mustbe ca rr ie d out on a large

    computer. (AprogramcalledBMD:07MStepwiseDiscriminant Analysis from

    theUCLAseries i s availabl e inmostcomputingprogram libraries).

    Inorder to investigate the natureand anatomical loca tion of the

    (cortical) events that underlie v is ua l perception, a studywas designed

    usingcomplexpatterned vi su al sti mu li (geometricshapes)todemonstrate

    what,i f any, differences i n the evokedpot ent ial configuration could be

    attributed to differences in stimulus meaningor to differences i n the

    physical properties of the stimuli. The experimentaldesign also included

    aparadigm to record the emittedpotential a c t i v i t y thatappeared during

    the intervalwhenthe subject expected but did not see one of the geometric

    shapes. A l l of the recordedwaveformsweredigitized and stored as single

    t r i a l s so that a variety of techniques, includingaveragingand discriminant

    analysis could be usedtomeasureany differencesbetweenthem.

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    5.

    Chapter II

    REVIEWOF THE LITERATURE

    2.1 The rela tionship of scalp recordedevokedresponsesto theunderlying brain activity.

    A f a i r l y comprehensivereview of the experimental evidenceand current

    theories isprovided by Kiloh,McComas&Osselton (1972)for the origin

    of spontaneous a c t i v i t y and byRegan (1972)for the o ri gi n of both

    spontaneousand evokedEEG a c t i v i t y . The availa bledataabout the origins

    of scalp-recorded e l e c t r i c a l potentials are fragmentaryand do not yet

    precisely define the sources of eitherspontaneousEEG a c t i v i t y or evoked

    potentials.

    Early reports suggested that the 0-60 Hz. "slow" a c t i v i t y in the EEG

    recorded at the c o r t i c a l surface i s the re su lt of asummationof action

    potentials of c o r t i c a l c e l l s . Otherslatershowedconclusively that neuronal

    action potentialswerenot e ssential and suggested that thewavesof the

    surface EEG resultfroma summationof the exc itatory or inhib ito ry

    postsynaptic potentialsdeveloped by the somaand larg er dendrites of the

    c o r t i c a lpyramidal c e l l s (Li &Jasper,1953). This suggestion received

    further supportfrom studies including intracellular recordings of cortical

    neurons (Jasper & Stefanis, 1965; Creut zfeldt,Watanabe& Lux, 1966) that

    showeda close timerelationshipbetweenre pet it ive postsynaptic potent ials

    andmacropotentials recordedon the surface of the ove rly ing cortex.

    Insomemorerecent workE l u l (1968)demonstratedthe existence of slow,

    spontaneousmembranepotential shifts i n c o r t i c a lneurons,in thesame

    frequencyrangeas the EEG. On the basis of theseobservationshe suggested

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    6.

    that the EEG may r e f l e c tthese slowerpotentials ratherthanbeing a

    temporalsummationof fas tera c t i v i t y , as the e a r l i e r studies of post

    synaptic dend ritic pot ential s had implied.

    The re la ti onsh ip of the EEG and evokedpotentiala c t i v i t y recorded from

    the scalp to the e l e c t r i c a l a c t i v i t y recorded fromthe c o r t i c a l surface

    has beeninvestigated by anumberofworkers. Geisl er & Gerstein (1961)

    showedbothexperimentally (inmonkey)and i n theo reti cal formulations that

    amajor attenuation of e l e c t r i c a lpotentialamplitudeoccurs in the highly

    conductivelayers of the CSF and dura but not at the skull or scalp.

    Cooper,Winter,Crow&Walter (1965)in studies onhumanpatientswith

    simultaneousrecording from the scalp and implanted c o r t i c a l electrodes,

    foundthat only "widely synchronized" cort ical activi tywas not obscured

    at the scalp and that theamountof attenuationbetweencortex and scalp

    dependeduponthe size of the area of cortex involved in the activity.

    In a study of the EEG a c t i v i t y followingsensory (flash) stimulation,

    Heath& Galbraith (1966),using c o r t i c a l and scalp electrodes i n the primary

    visual area showedthat an earlycomponent (65msec.)was evident at the

    samelatency i nboth c o r t i c a l and scalp recx)rdings; but for later

    (100 to 300msec.)peakstherewerelatency differenc esbetweenthe response

    componentsrecorded at the two electrodes. In explanation of thesewaveform

    differences, theysuggested that a c t i v i t y generated i nnon-primary cortex

    spreadto the o c c i p i t a l scalp lead but not to the c o r t i c a l lead. Hence

    they concluded forVER'sto flash, that the earlycomponentsin the response

    recorded fromthe s calp electrode directly overtheprimaryvisual cortex

    are generatedonly in the immediately underlying brain area, while the

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    7.

    laterresponsecomponentsrepresentsummationof a c t i v i t y frommore

    extensive (including both primary and non-primarycortex as faraway

    as thetemporal area)brain regions.

    Vaughan (1966)reported similar findings i n a patient explored with cortical

    electrodesduring an intracranial surgicalprocedure. However,in studying

    the distributionofevokedresponseto flash i n the c o r t i c a l regions

    adjacent to the calcarinefissure, he found theresponsedifferedmarkedly

    as the recording electrode was movedanteriorly fromthe occipital pole.

    He concluded thatmostof the scalp VER usually recorded in the occipital

    regions origin ated i n the occipitalpole of the cortex (foveal representation)

    and not inmoreanter ior portions of the striate cortex. This findingi s

    consistentwith other authors' observations (Devoe,Ripps&vaughan, 1968;

    Harter, 1970) that the VER pri ma ri lyreflects foveal stimulation.

    2.2 PatternVER'si n the study of vision.

    The responserecorded i n theoccipital region e l i c i t e d by spatially

    structured stimulus fields i s quitedifferent fromthe one evokedby a flash.

    Thismajordif feren ce was f i r s t described bySphelmann(1965)who showed

    thata patterned stimulusevokesaresponseofmuchgreater amplitude

    and containingpeaksof differentlatency and polarity than a flash of

    comparative l i g h t intensity.

    The type of pattern presentationusedto e l i c i t the "patternvisualevoked

    response" i s important. The stimulus may be abriefpresentation of a

    patternon a blank background; i t may be achangeof patternfrom one

    formto another (called a "patternreversal" i n the literature) or i t may

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    8.

    be illuminationof adarkpatternby aflash. Thetheoreticaladvantage

    ofapattern reve rsaltypeofstimulationi sthat therei s nochangei n

    the t o t a l luminancereaching thesubject's eye, onlyachangei n

    distributionofpattern structure withinthevisual f i e l d . Consequently

    someworkers (Halliday&Michael, 1970)havesuggestedthattheresponse

    to pattern re ver sal contains only "pattern specific"components,and not

    any toluminancechangeasdoestheflashVER.

    However,thepattern rever salmethodi susableonly for certaintypesof

    symmetricalpatterns suchascheckerboardsorgratings. I t hasalsobeen

    shownbyEstevez&Sprekreijse (1974)thatthepotentialgeneratedby

    pattern reversal includescomponentsfound bothi nresponsesto a pattern

    appearanceand apattern disappearance (analgousto an "on" and"off "

    response), andhencei snot asuncomplicatedasoriginally suggestedby

    Halliday.

    The thirdmethodofpresenting thepatterned stimulus (byillurninatinga

    darkpatternwithastroboscopic flash) requires le ss elaborate instrument

    ationand hasbeenused i nmany studies includingtheoriginalone of

    Sphelmann (1965). The VERproducedbythis typeofstimulation consists

    ofasummationof theresponsestoboththelargeluminancechangeproduced

    bytheflashand to thepatterned stimulus.

    For a l lof theabovestimulationtechniquesthestimuliaresuff iciently

    widelyspacedi ntimethatthenervous systemhasbeenregardedasreturning

    tothe samei n i t i a l statebetweensuccessive stimuli,andhencethese

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    9.

    responsesaresometimescalled "transient"responsestodistinguishthem

    from another typeofevoked pote ntia l described i n theliterature calle d

    a "steady state"evoked response. (SeeRegan(1975)fora complete

    descriptionofthisphenomenon). Theterm "steadystate"i semployedto

    indicate thatthestimuliarepresentedatsucharapid rate (e.g.10 - 60 Hz.)

    thatthec o r t i c a lresponsestoindi vidual st imuli overlap. Thesteady state

    responsei sthen separated fromthenon-event relatedbackground EEGbecause

    i t i scomposedofcomponents (or harmonics) whosefrequencies areexactlythe

    same as ormultiplesof thestimulus frequency. AFourier analysisi s usedto

    performtherequired computations and theamplitudeandphaseof thefrequency

    componentscan bedisplayedandmeasured.

    The techniquehas theadvantage ofspeed becausethestimulican bepresented

    somuch morerapidlythan theycan beforthestudyoftransient evoked

    responses. Regan(1974)hassuggested thatthepropertiesof thecomponents

    ofthesteadystateresponseseemtodependonwhetherthestimulation

    frequency i s(a)near10 Hz., (b) 13 - 25 Hz., or (c) 45 - 60 Hz. and has

    foundthatsomediseasesinvolvingthevisualsystemmayaffectthese

    frequencybandsdifferently. Thesteady statetechniquehasalsobeenused

    by Campbell &Kulikowski(1972)i nstudiesof therelationshipof a subject's

    a b i l i t yto see asinewavegrating patternof lowcontrastto thesizeof the

    evoked response e l i c i t e d by the samegrating pattern. Theyseparately tested

    thesubject's thresholdbyhavinghimreportwhen thegrati ng pattern

    disappearedas thecontrastwasdecreased,andthenbyrecordingtheevoked-

    responseto arapid pattern rev ersa lof thegratingatdiffe rent levelsof

    contrast. Theyfoundgood agreementbetweenthethreshold determinedby the

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    10.

    subject's verbal report and the contrast level atwhichthe evoked response

    could no longer be detected. The steady state patternevokedresponse

    thusprovides an objective indi cator of visual function and perception.

    Transient and steadystateevokedresponsesgivecomplementary insights into

    brain function. Van Hof (1960)and Tweel& Verduynlunel (1965)showedthat

    i t i s not possib le to pred ict the parametersof the steadystateresponses

    from the characteristics of the transientresponses. This i s not surp risi ng

    i nviewof the very different stimulation techniques, but consequently

    results obtained with one methodare not directlycomparablewith those

    obtained with the other.

    The remainderof the literature on patternVER'sto be reviewed i sconcerned

    with the transienttypeofresponse, as i s the experimental work described

    i n thisthes is.

    The individualpeaksof the t ransi ent pattern VER have been studied using

    topographicalmappingtechniques in an attempttodetermine the area of

    cortex inwhichthesecomponentsoriginate. Jeffreys(1971)and Je ffre ys &

    Axford (1972a& b) foundthat the f i r s t two peaksof the VER occurring

    65 - 80msec,and 90 - 110msec,af te r presentation of acheckerboard pattern

    (they called thesepeaks "C I" and "C I I" respec tively)weregre atly

    influenced by the retinal lo ca tion of the stimulus. The polarity of these

    componentsreversedwhentheupperrather thanthe lower f i e l d of vision

    was stimulated, and theamplitude of C IIchangeddependingupon the

    distance of the recording electrode from the calcarinefissure inboth the

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    11.

    horizontaland vertical planes. Basedon these and other observations of

    responsesto right and l e f t half f i e l d stimu latio n, Je ff re ys & Axford

    concluded th at the two peakshad spatially separate sources; the striate

    cortex was the source of C I and the ext ra -s tri at e cortex on the outer

    surface of the occipital lobes was the source of C I I . Ha ll id ay & Michael

    (1970)described a sing lecomponentat 100 msec,i n the response to a

    reversingcheckerboard stimulus that alsoshowedpolarity reversaldepending

    on the part of the visual f i e l d that was stimulated. Theysuggested that

    thiscomponentoriginated i nextra-striatecortex i n the occipital pole

    aboveandbelowthe calcar inefissure. A similarcomponentin the responses

    to a reversi ng pattern of red lightswas described by Purves& Low (1975)

    using separate stimul ati on ofupperand lowervisual fields. The polarities

    and latenc ies of thecomponentsdescribed by a l l of these groupsaresomewhat

    differentprobably due i n part to differences i n the luminanceof the stimulus

    and i n the techniques of stimulus presenta tion. The lack of standardized

    stimulation and recording techniquesmakesthe find ings of the different

    investigators d i f f i c u l t tocompare.

    Datafromintracellular recording in the visual cortex in cats andmonkeys

    (Hubel& Weisel, 1965 &1968); Bishop, Coombs&Henry,1971) haveshown

    thatmanyc e l l s i nthis brain region aremuchmoresen si ti ve to contoured

    thanto unstructured light stLmuli. Suchworkhas r esul ted i n important

    advancesi n the understanding of the c e l l u l a r physiology of the occipital

    lobe and of the fu nctio nal substrates of vision. Datafrom the study of

    VER'srecorded i nawakehumansubjects (see theabove observation,

    localizing the origin of these pattern specific responsecomponents to

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    12.

    striateandextra-striatecortex)inturn,haveprovided supportive

    evidenceof theexistenceofthese patternsensitivemechanismsinthi s

    regionandtheymayproveto be animportant toolfor further investigation

    of visual perceptual processes inman.

    In additionto thestudiesconcernedwiththetopographicalmappingof the

    responsecomponents,therehave beenmanyothers thathavedemonstrated

    the effectof avarietyofpatterned stimulusparameterson theevoked

    c o r t i c a l response.

    Clynes&Kohn(1967)reportedone of theearliest studies using stimulus

    patterns other thantherelatively simpleones (checkerboards andstripes)

    thatwereusedbySphelmann (1965). Theystudiedthespatial distribution

    oftheevokedpotentialsto alargenumberofvisu al stimuliincluding

    complexpatternsandcoloursandfound that therewere "spatially independent

    components"thatwerepeculiartoeachtypeofstimulus foragiven individual.

    Theyalso noted that whilethelatenciesofthesecomponentswere very

    constant, their amplitudes werequite variab le,a finding replicatedi n

    manylater studies. Thechanging polaritiesandamplitudes of theresponse

    componentstodiffe rent stimuliwhichthey reporteddosuggestthepossib ility

    ofchangingsourcesandsinksofcurrenti n thec o r t i c a l a c t i v i t yevokedby

    the different stimuli. Howevertheir ratheridiosyncratic electrode

    placements (inarosette patternaroundtheoccipitalregions),theiruse

    ofbi pola r recording deriv ations,andtheir largenumberofstimulus types

    makesi t impossibletocomparetheir resultstothoseof anylat er

    investigators,or to make anydefiniteinterpretationsofthemeaningof

    their results.

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    13.

    Rietveld, Tordoir,Hagenouw,Lubbers & Spoor (1967)and Harter &White

    (1968)presentedmoredetailed studiesof the responses to a checkerboard

    patternandshowedthatthe amplitude of the response from100 to200msec,

    post-stimuluswas sensitive to thesizeo f the checks i n the pattern.

    Theyshowedthati ncontrastto theflashevoked responses i n which the

    luminance of the stimulus i s an important determinant of thelatencyof

    thecomponents, (Creutzfeld &Kuhnt,1967; Tepas, Guterus & Klingman, 1974),

    thelatencyof thecomponentsi n thepatternresponse did notchangeover a

    widerange of luminances. Theysuggestedthat i twas the presence of the

    intersecting contrastbordersthatwas essential for the patterned type of

    evoked c o r t i c a lresponse. How sharplythe stimuluspatternwas focused was

    alsofound tohavea significant effecton the VER amplitude i n these

    studies. Manysubsequent reports (Arden & Lewis,1973; Harter & White,

    1970; John, 1974) havedemonstrated thatthe amplitude of the VER to

    patterns consistingofdifferent sizedchecks i s auseful tool i n the

    determinationof therefractive error i nc l i n i c a l ophthalmology. The

    patient'sresponse i s oflargestamplitude f or thesmallest sizeof checks

    on which he can focus clearly.

    Anumberof other stimulus parametershavebeenshownto be reflectedi n

    theVER configurationbesides focus, check size and luminance as indicated

    above. The overall sizeo f the stimulus for both unstructured(Rietveld,

    Tordoir& Duyff,1965) and structured (Harter,1971; Rietveldet a l . , 1967)

    f i e l d shasbeenfound to be important with amarkeddifferencebetween

    fovealand extra-fovealstimulation. The colourof the stimulus can also

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    14.

    influence the response (Kinney,McKay,Mensch& Luria, 1972; Clynes &

    Kohn,1967; Shipley, Jones & Fry, 1965) as can i t s or ie nt at io n

    (Kulikowski, 1974; Yoshida,Iwahara &Nagamura,1975) althoughthere i s

    somedispute aboutthe l a t t e r (Kakigi,Miyazaki &Mori,1972). Responses

    have beendemonstratedboth to on- and of f- se t of diff use (White &Easton,

    1966) and patterned l i g h t (Harter, 1971). The specific findings inthese

    reports that are relevant to the presentstudy w i l l be considered i nmore

    detail i n the Discussion Section.

    2.3 Evokedresponsesin the study of psychological variables and meaning.

    A very largenumberof publications i n the evokedpotentialliteraturehave

    originated i n laboratories of physiologic al or experimentalpsychology.

    A review of the relevantworksof nece ssity includes termsand phrases,

    the precise def ini tio ns ofwhichare outside the scopeof th is thes is.

    InappendixC a discussion of the terms "psychological variables" and

    "stimulus meaning"has been included toprovidesomebackground for the

    readerunfamiliarwiththese concepts, and to c l a r i f y th ei r use i n th is text.

    The concept of stimulus meaningi s also considered i n relat ionship to the

    experimental findings of th is di ss er ta ti on i n section 6.5 of the

    Discussion chapter.

    A greatmanyreports of the effec ts of a va riet y of psychological variables

    on the evoked potential configuration are reviewedby Regan (1972) and

    Beck (1975). These studies can be divided into two categories, those that

    are concernedwith the entireevokedresponse configuration and those that

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    15.

    are primarily concerned with a late positive componentof theevoked response

    (also called P^ or"P3QQ"becausei t appears as a prominent positive peakat approximately 300msec,after the stimulus).

    Chapman&Bragdon (1964)i n one of theearlieststudies of theeffect of

    psychologicalmeaningon theevokedresponse not specifically concerned with

    the component,comparedevokedresponses to task relevant (blank) and

    irrelevant (patterned) stimuli and found that the responses to task relevant

    stimuliwerelarger and differentthan those to the irrelevant sti muli.

    Howeverthey f a i l e d to consider that therewould have beenadifferencebetween

    the responses to theirblank and patternedstimulianywaywhethertheywere

    taskrelevant or not. L i f s h i f t z (1966)and Begleiter, Gross &Kissin (1971)

    i n similar studies claimed that the VER to pleasant,repulsive and neutral

    categorieso fvisual stimuliweredifferent for each category, and that the

    effectdisappearedwhenthe stimulus was blurredso as to be unrecognizable.

    John, Herrington & Sutton (1967)i n a widely citedstudy onmeaning found

    thatthe VER was different fordifferent geometric formsof equal area and

    similar fo r versions of thesamegeometric form of unequal area. They

    concluded thatthis evidence suggested thewaveform of theevoked response

    reflects atleast i n part the symbolicmeaningof the stimulus. Austt,

    Buno&Vanzulli (1971)i n a similar study claimed th at both the occipital

    visual evoked response and the visualevokedresponse recorded at the

    vertex arerelated to the processing of sensory information. The occipital

    response was saidto bemorespecificforvisu al stimuli than the one from

    the vertex and was modified by changesi nvisualperception, significance

    ofthe stimulus and theprogramthe subject was performing.

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    16.

    Buchsbaum& Fedio (1969) i n a study of the visualevokedresponse to

    differentpatterns, includingsomewords,also found differences i n the

    VER waveformsthatwererelated tomeaning. Becauseof their relevance to

    the findings of the present study these l a t t e r three studiesw i l l be

    considered in furtherdetail i n the Discussion section,

    Begleiter& Platz (1969) showedthat the responseevokedby a particular

    visualstimulus could be modified by c l a s s i c a l conditioning. A negative

    peakat 155

    msec,to 160

    msec,latency was enhancedwhenthe stimulus was

    conditionedby asso ciation with a loud trainof clicks and thisenhancement

    was reversiblewith extinc tion and reconditioning. In threemorerecent

    reports (Begleiter, Porjesz, Yerre &Kissin, 1973; Porjesz & Begl eiter,

    1975; Begleiter & Porjesz, 1975) of the effect of the subject'sexpectancy

    or perception of a stimulus, these authors haveshownthat the amplitude of

    the vertex VER to a flashofmediumintensitywas higher i f the subject

    expected orthought he saw a brightflash and significantly smaller i f he

    expected orthoughthe saw a dim flash. Thesefindings are reminiscent of

    John, Shimokocki&Bartlett's (1969) and Ruchkin&John's, (1966) findings

    of "neural read-out frommemoryduring generalizat ion" i n cats . These

    investigatorstrai ned cats to discriminatebetweentwo stimulus flicker

    frequencies. Theyfound that theevoked potential i n various regions of

    the brain to a stimulus of an intermediate flicker frequency closely

    resembledthe responseto the appropriate signalfor the behavior which

    was displayed on that t r i a l ; the configuration of theresponseevoked by

    the intermediate stimulus was differentdependingonwhetherthe catmade

    the behavioralresponseassociated with the high or low frequency flicker.

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    17.

    The authors concluded thatthis const itu ted evidence fo r relea se of a

    neural process representing previous experience and provided somefurt her

    support fo r the s t a t i s t i c a l theory of lea rning andmemorysuggested by

    John (1967).

    Investigatorswhoseworkhas centred on the experimental manipulation of the

    latepositive (P^)componentofevokedpotentialshavesuggested a large

    numberof p ossi ble psychologi cal variab les that may relate to thispeak.

    It hasbeenvariablyproposedthat the "P^" i s a physi ol og ic al sig n of

    "deliveryo f task-relevan t information" (Donchin&Cohen,1967),of the

    "resolutionof prior uncertainty" (Sutton, Tueting, Zubin &John,1967) of

    "cognitiveevalu ation of stimulus significance" (Ritter &Vaughan,1969), of

    "a decision regarding the stimulus" (Rohbaugh,Donchin &ErUcsen,1974), of a

    "reactivechangein the state of arousal" (Karlin, 1970),and of "stimulus

    salience" (Jenness, 1972). Someauthors (Karlin, 1970, Naatanen, 1969)

    haveargued a non-cognitive explanation f or a l l of these fin din gs, suggesting

    thatthe dif fer enc e i n theresponsesto the relevan t or attended stimuli i n

    a l l of the experiments can be explained by a "prior-state" hypothesis,

    meaningthat a l l of the f ind ing s could be explained on the basis of achange

    i n the subject's sta te of arousal or expectancy and not by any specific

    parameters of the stimulus i t s e l f . However,Picto n &Hillyard (1974) and

    Ford, Roth, Dirks & Kopell (1973)i nmorerecentworkstudying evoked

    responsecorrel ates of selective attentionhaveshownthat this i s not true .

    A l lhavedemonstrated two separate systemsoperating during selective

    attention (for auditory stimuli); one ind ica ted by increased amplitude

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    18.

    of wavesand invo lvingselectionof a particular input channel and

    thesecondindicated by themorewidespread complexand involving

    complexinformation processing or decisionmakingbased on information

    provided by the stimulus. Harter &Salmon (1972),i n an earlier study,

    had also found a consistent "differencewaveform"with an earlier negative

    and later positivepeakbetweenresponsesto task relevant and task irrelevant

    patterned visual stimuli (although the non-specific"priorstate" of the

    subjectwas not as carefully controlled i nthisworkas i t was i n the later

    studiesthatusedauditory stimuli).

    I t has alsobeenshownthat the componenti s not e l i c i t e d by a l l attended

    and task relevantstimulior emitted stimuli (seebelow)that require

    decision,but only by those forwhichcertain requirements of detection

    confidence and prior subject uncertaintyhave beenmet (Squires,Hillyard &

    Lindsay, 1973; Sutton, Braren, Zubin &John,1965; Tueting, Sutton &

    Zubin, 1971). In a recent report on the P^ componentduring an auditory

    detectiontask withcuedobservation intervals Squires, Squires & Hillyard

    (1975a& b)havefurtherc l a r i f i e d the effect of these two majorfacto rs of

    decision confidence and expectancy. They suggested that the P^ (becauseof

    i t s topographyand sensitivityto the probabilityof the stimulus occurrence

    orabsence)i s thesamep hys iol ogi cal bra in event,whetheri t occurs i n a

    threshold detection task, omitted stimulusparadigm (Picton,Hillyard &

    Galambos,1973) or a single-double stimulus discrimLnation task (Sutton

    et a l . ,1967).

    A few authorshavereported that a c o r t i c a l responsecan be e l i c i t e d by

    omission of an expected stimulus; theseresponsesto stimuli that are

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    19.

    expected but do not occurhave beencalled "emitted potentials" by

    Weinberg,Walter &Crow(1970)and this termhasbeenadoptedbymostother

    investigatorsof thisphenomenon. Sutton, Ruchkin& Tueting (1974) reviewed

    mostof the relevant studies of the componentof the emitted potential

    and the experimental paradigmsusedfor i t s production. Squires etal.(1975b)

    providedsomefurther experimental evidence that c l a r i f i e ssomeof the

    questions on theappearanceof the P^ i n threshold detection tasks raised

    i nthe presentation of Sutton et a l .(1974) and they alsodemonstratedthat

    the essential c r i t e r i a fo r a stimulusabsenceto e l i c i t a P^waveare that

    i t i sclearly recognizable and relatively improbable.

    Therehasbeensomecontroversyaboutthe s p e c i f i c i t y of the emitted response.

    Sutton et a l .(1974)stated that there was no evidence available indicating

    thatthe P^componentof the emitted potential i sspecific for the modality

    or intensity of the stimulus. Ritter,Simson&Vaughan (1974)howeveri n

    another paperfromthesamesymposiumd id providesomeexperimental evidence

    thatthe negativecomponente l i c i t e d by the missing stimulus was different

    forthe two modalities theyused (visualand auditory) but they did not

    investigate s p e c i f i c i t ywithin a modality. Otherauthors who havedescribed

    emitted responses have been investigating effectsof stimulus timing

    (Klinke,Fruhstorfer & Fin kenzell er, 1968) selective attenti on (Picton et a l . ,

    1973) or the subject's expectancy i n a guessing paradigm (Weinberget a l . ,

    1970) andnoneof these experiments provides any evidence on the specificity

    of the response to the modality ormeaningof the omitted stimulus.

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    20.

    24 Summaryof the literature review.

    It hasbeenestablished that sensory evokedresponsesrecorded from the

    scalp r e f l e c t den dri tic post-synaptic pote nti ala c t i v i t y i n the underlying

    c o r t i c a l c e l l s . I t has alsobeenshownthat only widely synchronized

    cortical activityappearsi n scal p recordingsbecauseof the att enuation

    by the intervening layers ofmeninges,cerebrospinal f l u i d and skull.

    The EEGwaveformrecorded after presentation of a sensory stimulus consists

    (after theon-going,non-eventre lat ed EEG a c t i v i t y hasbeenextracted) of

    a ser ies of well definedpeaksorcomponents. Experimental evidence

    suggeststhat at leastsomeof thesecomponentsoriginate i n different

    c o r t i c a l regions. I t has al sobeenshownthat these differentcomponents

    are se nsi ti ve tochangesi n a var ie ty of stimulus parameters.

    The use of patterned visual stimuli toproduceevokedresponsesi s a

    relatively recentdevelopmentthat allows the study ofmuchmorecomplex

    aspects of stimulusmeaningthandid the oldermethodof stroboscopic

    flashpresentation. Componentsof the pattern VER have beenshown to be

    sensitive to focus, to thenumberof lines per uni t area (contour density)

    and to a les ser extent t o the luminanceof the presented stimulus. Steady

    stateevokedresponses (related i nprinciple to transientevoked responses,

    but e l i c i t e d by a different stimulationtechnique) to grating patterns of

    varying contrast levels and spatial frequencieshave beenshownto begood

    indicatorsof the subject's thresh old fo r these patterns asdetermined by

    separate psychophysical threshold testing.

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    21.

    Thereis also a largebodyof literature concernedlesswith the physical

    parametersof the stimulus andmorewith the ef fe ct of psychological

    variables on theevokedresponseconfiguration; bothwithnon-specific

    variablessuchas attention, and withmorespecific cognitive variables such

    as stimulus meaningor decision-makingbasedon information provided by a

    stimulus. I t has beenshown,forexample,that a late positivecomponent

    of the responsei n any modality i s related to this decisionmaking process,

    but that thewaveappearsi n theevokedresponseonly i f the subject i s

    certainhe detects the "stimulus" and i f the stimulus occurs relatively

    infrequently during the paradigm.

    I t has alsobeenclaimed that theevokedresponseconfiguration i s affecte d

    by the symbolicmeaning of the stimulus. However,the few published studies

    purporting todemonstratesucha relationshiphave failed to define clearly

    either the experimentalchangesmadein stimulus meaningor the effects

    theyproduce on the configuration of theevokedresponse.

    Therehave been a few reports of the recording of c o r t i c a l responses (at

    the scalp) e l i c i t e d by s ti mu li that areexpectedby the subject but do not

    occur. Thesehavebeencalled emitted potentials since they are not evoked

    by an external stimulus butpresumably r e f l e c tsomebrainprocessrelated

    to therememberedstimulus. I t has beensuggestedthat this typeofresponse

    recorded at the scalpmightprovide a usef ulmethod of examiriing brain

    processes rela te d to stimulus rteaningwithoutcmtaminationby activity

    that isdirectlyrelated to the sensory input. To thisdatehowever,

    neither the techniques for obtaining emitted pot ent ial s, nor the configuration

    and topographyof thesewaveformshave beenwelldocumented in the

    literature.

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    22.

    Chapter I I I

    STATEMENT OF THE PROBLEM FORINVESTIGATION

    The questionofwhetheranypartof theresponse evokedbypatterned stimuli

    reflectshigher perceptualratherthan simple sensory processi ss t i l l

    unresolved. In tworecent, comprehensive reviewsof thef i e l d (Ciganek,1975;

    Regan,1972),theclaimsofJohneta l . (1967)weresingledout ascr ucial

    becauseoftheir significance forthestudyofevoked response correlates

    of higher processes. Both reviewers urged thatthesefindings require

    independent validationanddevelopment. Tothisdate therehasbeenno

    reportofsuch studies.

    Thereforeaseriesofexperimentswasdesignedtostudytheeffectof the

    geometric formandmeaningof thestimuluson theconfigurationof the

    visual evoked response. More s p e c i f i c a l l y i twas proposed:

    (1) Toestablishwhethertherearereliableorcons istent

    differencesbetweenevoked responsestogeometricshape

    stimuliand todeterminethespatial (scalp distribution)

    of anydiffere nces.

    (2) Todemonstrate theeffecton theevoked responseof

    assigningameaning(ascuesi n atask)to thediff erent

    shapes.

    (3) Torecordthe"emittedpotentials" occurringwhen asubject

    expectsbutdoesnotactuallyseethese shapes,andthus

    determine i f i t i spossiblet orecordEEGcorrelatesof

    brainprocessesrelatedto themeaningof astimulusi n

    the absenceof thestimulus i t s e l f .

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    23.

    ( 4 ) Toapplyavarietyofanalytical techniquesto theevoked

    potentialsobtainedtoexploremeansofquantifyingany

    differencebetween thecomplexwaveformsrecordedi n the

    differentexperimentalparadigms.

    I n a l lof theexperimentalwork,data collectionandanalysiswere performed

    byad i g i t a lcomputerwherepossible. Singlet r i a l samplesweresavedso

    that'a variableandsmallnumber oft r i a l s couldbeusedi n theaverages

    andsothat alternate analysis techniques suchas astepwise discriminant

    analysis,manipulationofsingle t r i a l s priortoaveragingand computation

    of theratio s t a t i s t i c calledtheAdescriptor (Johneta l . , 1967)could

    becarriedout. (Seesection6.2 for adetailed discussionofthese

    analyticaltechniquesandAppendixB for adescriptionofstepwise

    discrimiiiant analysis.

    Becauseinaveragesofsmallnumbers o ft r i a l s ,anya c t i v i t yo frelatively

    highvoltage presenteveni nonlyone t r i a l affectstheaveragedwaveform

    to adisproportionate extent,anautomatic artefact rejectionroutinewas

    includedi n thedataacquisitionprogram (seeMethods).

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    ChapterIV

    METHODS

    4.1 DataAcquisitionMethods.

    Forthef i r s t part (paradigm1)oftheexperimentt hir tee n subjectswere

    used. Ten ofthesewerepaidwomenstudent volunteers betweentheages

    of19 and28. Theother threeweremembersoftheLaboratory St af f. A l l

    ofthesubjectsweretestedatleastonceandsomeweretested repeatedly

    but alwaysondifferentdays separatedby atleastaweek. At o t a lof

    26runsofparadigm1wereusedfortheanalysi s.

    Fiveofthethirt een subjects, includingtwo ofthestaffmembers,were

    usedforthesecondpartoftheexperiments (paradigm2),i nwhichfrom

    threetotenseparate recording sessions foreachsubjectwerecarriedout.

    The subjectssatalonei nashieldedroomwithlowintensityoverhead

    lighting (.5ft-candles)andwatchedatelevision screen1meterdista nt

    forthepresentationofthestimulus.

    The electrodeswereGrassgold discs applied with Grasselectrode paste

    accordingtotheInternational10-20system (Jasper,1958)forelectrode

    placement. (SeeFig.1fordiagramatic representation). Fora l lof

    paradigm1thestandard locationsof 0 2, 0^ P 4, P , C 4, C 3and C zwereused.

    Inparadigm2theselocationswerealsousedbut in someexperiments

    0 , P , C , F and twolocati ons designated CP 0and CP.locatedhalfway

    Z Z Z Z j 4betweenCj and P^ and C^ and P respectivelywereused. A l l recordingwas

    referredtothecontralateralearor tothel e f tear i n thecaseof

    midline locations.

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    25.

    Fig. 1 Diagramaticrepresentationof thepositionof the

    electrodes on thescalp.

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    26.

    The EEG was recorded on aBeckmanDynograph TypeR with a time constant

    settingof .3 sec andhigh frequencycut-off f i l t e r a t 30 Hz (aset ting

    of 3 on thepoweramplifier f i l t e r ) givingan effectivesystem band width

    of 0.5 -30Hz (3dB points).

    FromtheDynographthe EEG signalson seven channelswereled to A-D

    convertersof the computer (adetailed descript ionof the computer system

    used i n these experiments i sgiveni n Appendix A) and sampled every 2msec,

    for a 512msec,epoch, beginning 56msec,before thestimuluspresentation.

    Single t r i a l s (256pointsper trial) werestoredon thediscof the PDP 11

    computer system and a l ldata was transferredto Dectapes forpermanent

    storageat the end of each paradigm.

    The experimental set-up i sshowni n the diagram i nFig. 2. The experiments

    were a l l controlled i nrealtime by a program calledVER03which ran on the

    PDP 11 system. The flowchartof the program i sgiveni n Appendix A with

    someexplanation. An automaticartefact reject ionroutinewas included.

    This routinechecked each t r i a l sample to determine i f i t exceededsome

    presetamplitude limits, and i f i t di d the t r i a lwas not saved. The program

    thencontinued the experimentu n t i l therequirednumberof t r i a l s had been

    collected i n each group to be sampled. (Thenumberwas 30 t r i a l sof each

    shape i n paradigm 1, and 10 t r i a l so f each shape in paradigm 2). I t was

    found fromobservingthe paperwriteouto f the EEG a c t i v i t y that this

    technique effectively excluded any t r i a l swheretherewas eye blink

    or subjectmovement. Insomeexperiments the EGG (electro-oculogram) was

    recordedon one o f the seven channels to check f or any event-related slow

    eyemovementswhich might nothavebeen rejectedby the system.

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    D a t a A c q u i s i t i o n S y s t e m .

    Fig. 2 Experimental se t up.

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    The stimuliwere the fourdifferentgeometricshapesshownin Fi g. 3.

    They included two familiarshapes, (one with corners and one without),

    a square, and a c i r c l e , and two unfamilia r and le sseasilynamedshapes

    calledhere the e l and theomega. Theywerea l lshownaswhiteoutli nes

    on a blackbackground. A l l had approximately equal length of contrasting

    border. Theywerepresented as slidesby aKodakCarousel pr ojecto r.

    The imagesfromthe projectorweremonitoredby a televisioncamera and

    were transmitted to the T.V. monitor placed one meteri n front of the

    subject. An electromagnetic shutter (Gerbrant) on the lens of the

    projector controlled the brief 20msec,presentation of the stimulus.

    Advancementof the carousel and the shutteropeningwerecont rol led by

    thecomputer.

    The size of the shapeson the T.V. screen was approximately 6 cm. by 6 cm.;

    therefore the shapessubtendedan angle of 3.6 degrees at the centre of

    the subject's visual f i e l d . The luminancemeasuredat the screen was

    10 ft-candle s fo r thebackgroundand approximately25 ft -candles for a l l

    four stimuli. The subject was instructed to fixate visually on a point on

    the centre of the screen throughout the recording period.

    Forparadigm1 (see Figure 4 for a diagramatic representation), the four

    stimuliwerepresented i n arandomsequenceat from 3-5 sec. in te rv al s

    u n t i l 30 t r i a l s ofeachshapewerecol lec ted . The number30 was chosen

    becausei t was the lowestnumberper averagethatproducedconsistent

    evokedpotentialwaveformsand was the highestnumberof si ngle trials

    i twouldbe p ossibl e to store in thecomputermemoryduring a sin gleexperiment.

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    29.

    Fig.3 StiraulijsShapes 1.circle 2.square 3.omega 4.el

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    Stimulus presentation

    20 msec

    fc==l-*H

    56

    msec*

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    31.

    The subjectswereinstructedt owatchthepresentationof thediffer ent

    shapesand totrytostay alertandattentive. Total time f or paradigm1,

    includingone or twobrief interruptions,wasfrom10 to 15minutes. The

    ongoingEEG wasmonitoredon thepaper writeoutby theexperimenterand

    when theAlpha rhythmbegantodominatetherecordor anyTheta activity

    appeared,theexperimentwasinterruptedand thesubjectaskedtorefocus

    her attention afterabrief rest.

    Inparadigm2 (seeFigure4foradiagramaticrepresentation), five shapes,

    a c i r c l e , e l ,cross, squareand omegawerepresentedi n the sameordera t

    regular intervalsof 1.4sec. Inapproximatelyonethirdof thesequences

    eitherthesquareor thee lwasomitted (theshutterdidnotopen). The

    experiment continuedu n t i lthe EEGduringtenomissionsofeach shapewas

    collectedandstored. Withnorejectionsdue toartefact this required

    63 sequencesoffiveshapes,or 315stimuli, which tooklhminutes.

    Ten responsestoeachof thefourshapes usedi nparadigm1 whenthey d id

    appearwerealso saved.

    The subjectwasinstructedtolearnthesequenceoffive shapes, thenwas

    told that occasionallythesquareore l would f a i ltoappear. She wasgiven

    two buttons (handheldthumb-switches) andinstructed thati fthesquare

    f a i l e d toappearat theexpected time,shemustpresstheleft-handbutton

    andwhen thee lwasemittedshemustpresstheright-hand button. Failure

    torespondcorrectlywithin400msec,of when thestimulus shouldhave

    appearedi n therhythmic sequence,orpressingthebuttonwhen thestimulus

    did appear resultedi n awarning tone being sounded (controlledby the

    computer).

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    The rhythmicpresentation sequence was not broken by the warning tone.

    Button presseswererecorded as squarewavepulses on one of the seven

    datachannels sothatthe exacttimingof the responsecouldbecompared

    to the EEG a c t i v i t y . I talsoprovided a way of counting thenumberof

    correctresponses at the end of the experiment. The subjectswere allowed

    to practise this f a i r l y d i f f i c u l t taskpriorto therecordingsession and

    a l lbecamequiteproficienti nmakingthecorrectresponsewithin400msec,

    in from70 - 100%of the t r i a l swith the missing stimulus, and i nmaking

    veryfew responseswhenthe stimulus was present.

    At the end of an experimentalsessionthe datawereaveraged, 30 t r i a l s per

    average for paradigm 1,and 10 t r i a l s per average for paradigm 2and the

    averageswereimmediatelywrittenout. A l l of the data (assingle trials)

    was thentransferredto Dectapes f orpermanentstorage andcould be

    accessedfrom there by aprogramcalledGASPf orfurther analysis.

    Q

    Calibrationof the dataacquisitionsystem was doneat the beginning of

    each experiment by insertinga5 Hz. 50yv squarewavesignalfrom a Grass

    calibrator intoone pairof inputs at the headboard. A l l seven recording

    channelswerecalibratedusingthispairofelectrodes. The VERprogram

    was then run and a writeout ofsomesingle t r i a l smade. The gainswere

    adjustedon the EEG amplifiers sothatthe50 yv signal i n a single t r i a l

    appeared with an amplitude of25mm. on theplotter. Sincea l l averaged

    EEG signalson the f i n a l plotweretruemeans (ratherthansums)a50 yv

    signalappeared with a25mm. amplitude i n the final plotwhetheri t was

    the signalof a single t r i a l or the signal i n the averaged response.

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    4.2 DataAnalysisMethods.

    The differencesbetweenthe evokedpotentialwaveformse l i c i t e d by the

    different shapeswereanalyzed by a va ri et y of techniques. They include

    visual inspection foridentification andmeasurementof the prominent

    peaksof thewaveformsand twomorecomplexquantitativeapproaches.

    The simpler of the two i s the computationof a descriptors t a t i s t i c called X

    that was described by Johnet a l .(1967). This s t a t i s t i c gives a

    quantitativemeasureof how different two waveforms are and the formula

    andmethodsusedtocomputei t are givenbelow.

    The secondmethodof determining the dif fer enc esbetweenthe responsesi s

    called discriminant anal ysis , a techniquewhichi sbasedon multi varia te

    s t a t i s t i c a l theory. A desc riptio n of the technique and the assumptions

    onwhichi t i sbasedi s given i nAppendixB. The discriminant ana lys is

    performedon this experimental data was carried out by theBMD:07Mprogram

    i n the UBC computingcenter. The input data fo r theprogramconsi sts of

    two ormoregroupsof observations (inthis experimental workeach "group"

    was ashapeand eachobservation was the EEG a c t i v i t y recorded during a

    single presentation of thatshape). Eachobservation i n turn consists of

    anumberof variableswhichi nthis studywerethe digitized values of the

    EEG voltage at different timepoints.

    The programproceedstochoosea subset of the var iab les (i .e . voltages

    at different timepoints) and cal cul ate s a discriminant function using these

    variables thatw i l l discriminatebetweenthe differentgroupsthatwere

    entered in to theprogram. Thismathematical functio n can be evaluated fo r

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    34.

    anewobservation (asingleEEGsample)andi tw i l l provideapredict ion

    ofwhich group (shape) thisnewobservationwasmost l i k e l ytohavecome

    from. Themethodtheprogramusestosel ect the varia blesand tocompute

    thisdiscrimin ant function i s described i nAppendixB. Certainparameters

    for thisprocessmustbesetatthe timethe data i s enteredandthese are

    also given in thatappendix. Anotherre late d, butmuchsimpler program

    calledUBC:CLASSevaluates the discriminant function fo rnewdata.

    In ordertosubmittheEEGdatatotheUCLABMD:07Mprogram (runonthe

    UniversityIBM 370computer) the selecteddigitized t r i a l s (one channelat

    a time)weretransferredfromDectapetoeitherpapertapeormagnetictape

    on thePDP 11system. Duringthe transfe r the digitized t r i a l representation

    was reduced from256pointsto 67pointsbyomitting the f i r s t26points

    (whichrepresented the52msecs.ofthe pre-stimulus bas eli ne) ,then taking

    the averageofeachofthe67setsof 3poin ts t hat followedandomitting

    the l a s t29points. Thedata fromonechannelat atimewerethentransferred

    to f i l e sonthe Uni versi tyIBM 370system. Each67point single t r i a l

    representation includeda 5character label indicatingwhichshapei tcame

    from. The BMD:07Mprogramwasrun for ei th er2 or 4groups (each group

    consistingof 30t r i a l s e l i c i t e d by adifferent shape). On a fewruns

    60 t r i a l s per groupwere included but the separationofthegroupsbythe

    programdi d not appearto beimprovedbythis increase in thenumberof

    observations per group. Thenumberofvaria ble s per observationwas 67 as

    indicatedin the previous paragraphandtheprogramselected6 or 7 ofthese

    (i.e.6 or 7latency points)asthe b asisofthe discriminant function.

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    35.

    P i l o t studiesshowedthat allowing theprogram to run throughmorethan

    6 or 7steps (i. e. tochoosemorevariablesthanthis) did not improve

    the discrimination and so the programwas instructed to stopatthisnumber

    of steps. This discriminant function producedby each run of the program

    was stored i n aseparate f i l e and thenamesof the variab les i t selected

    wererecorded separately (e.g. the 5th, 21st, 35th,42nd, 51st and 60th

    variable)..

    For test ing one of thesefunctions, datafroma l l of the experiments for

    thesamesubject fo r thesamechannel that had not beenused fo r input to

    theBMD:07Mprogramthatdetermined the function,wereput i n a sing le f i l e .

    The programUBC:Classwas run on this f i l e with the discrhntlnant function

    and aformat statement indicatingwhichwere the six or seven variables

    (latency points) to be selected fromthe responsefor thi s partic ular

    function as it s input. The r es ul ts of th is testing wereexpressed as

    percentagesof correc tlyc l a s s i f i e d t r i a l s and i n the case of the 4group

    choices, thewrongly c l a s s i f i e d t r i a l swerealsogrouped as towhichshape

    theyweremistakenfor and thesepercentageswere tabulated.

    Tocomputethe X rati os as described by Johnet al .(1967)averages of

    10 t r i a l swereprepared on thePDP-11systemwiththeGASPprogram. These

    weretransfer red bymagnetictapeto the IBM 370 wherea Fortr an program

    computed the X values for the 4averagesgiven i tfrom the following formula:

    = ( dl , 3 + d2,4 } / ( dl , 2 + d3,4 )

    whered.. i s the absolute value of the rootmeansquare (r.m.s.) difference

    between waveformsi and j . /"r.m.s.= x. /nwherex. i s the i sample

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    36.

    of thewaveformwith the valueof thebaselineascomputedfromthef i r s t

    3points(18msec.)of theresponsesubtracted fromi t V Thesubscript s

    1and 2denotethe tworeplicatedresponsesto oneshapeand 3 and 4

    denotethereplicatedresponsesto thesecond shape.

    Analysisof theresponsestothe omittedstimuliwas madesomewhat

    d i f f i c u l tbytherelatively smallnumberofthese responses available.

    The squareore lwasomitted from i t spositioni n thesequence only

    occasionally. Anexperiment consistingofpresentationof atlea st

    320 stimuliprovidedonly20"emitted" responses, ten each forthesquare

    and e l . In theaveragesofonlytent r i a l sthebackgroundEEG(noise)

    levelwassometimeshighenoughto makei td i f f i c u l ttovisuallyseparate

    the componentsofevent-related a c t i v i t y , especially i f thesewere

    relatively smalli n amplitude. However,combining data frommorethanone

    experimenttoprovidemoreresponses fo r averagingintroduced added

    v a r i a b i l i t yto theresponses becauseofchangesi n thesubject's performance

    level, stateofalertnessor i ntechnical elements suchasminor differences

    i n electrodeplacement. Inviewoftheseconsiderations, forpresentat ion

    of theresults, themeasurementsmade byvisual inspection fo r compiling

    tablesVTI andVTIIweremade onaveragesof tent r i a l s from each single

    experiment,butforthei l l u s t r a t i o n sof theresponses30t r i a l speraverage

    wereusedsothattheevoked responsecomponentscouldbemore easily

    visualized.

    Aftertheconventional averageswereproducedandexamined,analternate

    techniquesimilartotheoneusedbyRuchkin (1974)wast r i e d tocompensate

    fortheeffectof avaryingestimateof thetimeofoccurrenceof the

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    37.

    omittedstimulus by thesubject. Its purpose was tohave a l lof the

    peaks from thesingle t r i a l s coincidei n timeprior to averaging, so that

    any activity thatwas time-locked to the P peaks would be enhanced.

    The single t r i a l s at C were f i r s t filtered by a 0 - 6 Hz band-pass

    d i g i t a l f i l t e r tomakethepeakof the slow positive activity clearerfo r

    measurement. The filtered trial swerethenmovedto adisplayscreen.

    Aclear positivewavecouldbe seen i n about 90% of the responsesexamined,

    between240 and 450msec,afterthe stimulus shouldhaveoccurred.

    (If nopeakcouldbe seen inthat interval the t r i a l was notincludedfo r

    the "shifted" average). The latencyof themostpositivepeakwasmeasured

    and thesizeo f the shift along the timeaxis (inmsec.)thatwas required

    tomovethepeakof thelargest positivewaveto an arbitrarily chosen

    centralpoint (350msec.)was calculated. As a simplecontrol, this

    shiftingprocedure was also appliedtosomeresponses from single trials

    elicited by the square and e lwhentheywerepresent. Thisi sfurth er

    discussed i nsection5.3.2.

    TheGASPprogram mentioned included routinesforretrieving the single

    t r i a l s from any experiment from Dectapes, averaging anynumberof responses,

    switchingaround the seven channels i f necessary i n caseswhere different

    channelshadbeenused for a givenscalp location, displayingaverages or

    single t r i a l son thedisplay screen and f orplottingany of these responses

    beforeo raftermanipulation on aprinter-plotter. The program also had

    the capacity fordigital filter ing of data through whatever bandpass was

    required.

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    38.

    Chapter V

    RESULTS

    5.1 Effectof the DifferentExperimental Conditions on theEvokedResponse.

    Before examiningthe effectof the stimulusshapeon the configuration of

    theevokedresponse, i t i s necessary to assess the contribution of the

    differentpresentation conditions of the two experimental paradigmsi n

    order to decidewhetherthe datafromthe two could becombinedfor

    subsequent analysis.

    In the stimulussequenceofparadigm 2, the c i r c l e andomegawerealways

    present and served only as cue stimuli. For the square and e l , on the

    otherhand,since theappearanceof the appropriateshapemeantthat no

    button press was to bemade (detected omissionsweresignalledby pressing

    one of the two switches) and since there was someuncertainty as towhether

    the square or e lwouldappeari n i t s time interval in any given stimulus

    sequence, the presence of these stimulusshapeswas informational!/more

    important for the subject.

    In theevokedpotentialse l i c i t e d by the c i r c l e andomegathere was

    essentiallyno differencebetweenthe responses recorded i n the two paradigms

    at any of the five recordingsitescommonto both experiments. Figure 5

    showsthese responses for one subject in the two paradigmsand i t can be

    seenthat theevokedpotentialsfromparadigm1 and 2 are very similar.

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    39.

    100msec

    Fig.5 Effectofexperimental conditionsfo r thec i r c l eand

    omega. Paradigm1 (notask) responsesare shown bys o l i d linesandparadigm2 (task)responsesare shownby dotted lines. n=30trials/average. (SubjectSC

    Exp. 18A,34,35)

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    In theevokedpotentialse l i c i t e d by the squareand e lhowever, there

    wereconsistent differenc esbetweenthe two paradigms, but only in the

    componentslater than 220msec. Inparadigm 2, but not inparadigm 1,

    abroadpositivecomponentwith amaximumamplitudeof 8 to 18 pv

    (depending on the subject)between 220 and 320msec,post-stimulus was

    seenin the responses e l i c i t e d by the squareand e lshapes. This diff ere nce

    i s i l l u s t r a t e d i n Figure 6 for one of the subjects. Thi s lateposit ive

    waveappearing i nparadigm 2 was la rg es t at C , smaller i namplitude but

    definitely present i n the right and l e f t parietal regions, and small and

    inconsistentlypresent i n the occipital regions. For a given subject this

    positivewavewas of thesameamplitudeand config uratio n fo rboth the

    squareand e l responses. Therewereno amplitudeasymmetriesbetween

    and C^, CP^ a n a- CP^ (notshowni n Fig. 6) and P^ and P .

    Insummary,the effect of the task imposed i nparadigm 2 on theevoked

    responsesto the four present shapeswas as follows:

    1. The effectwas to introduce abroadpositivewavei n theevoked

    responses at C ,whichwas only varia bl y present and of smaller

    2

    amplitude i noccipital regions.

    2. Thiswavewas only seeni n theresponsese l i c i t e d by the square

    and e lshapes and not to the c i r c l e andcmegashapes.

    3. I t was seenonly i ncomponents220msec,or later post-stimulus.

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    41.

    +100 msec

    F i g . 6 E f f e c t o f e x p e r i m e n t a l c c n d i t i o n s fo r the square and e l .

    Paradigm 1 (no task) respon ses ar e shown by d o t t e d l i n e sand paradigm 2 (task) respon ses a r e shown by s o l i d l i n e s .

    n=30 t r i a l s / a v e r a g e . ( Su b je c t MP, Ex p. 7,53,54 )

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    Inview of theabove findings, i nconsideringthe data foranalysisof

    theeffectof thedifferentshapes on the evoked responses, the circle

    andonega t r i a l s from paradigm 1 and 2weremixed. For the square and

    e l shapes there was a differencebetweentheparadigms whichwas maximal

    atC and startedonly 220msec,post-stimulus. Since i t was very small

    in occipital regions theresultsof computerized analysisof the

    differentresponses at the occipital locationsfor these two paradigms

    werealsocombinedforconsiderationi n the sectionto follow.

    5.2 Effectofdifferentstimulus shapes on the evoked responses.

    Consistentdifferencesbetweenthe evoked responses to the four different

    shapeswereobserved. In order to best define and quantify these

    differences,three techniqueswereused. F i r s t ,the data from the

    standard paradigm 1condition (no task) was examinedvisually, the

    componentslabelled and amplitude and latencymeasurementsmadeas

    describedbelow. Secondly, thedigitized single t r i a l s from occipital

    channels of one experimentweresubmitted to theBMD:07MStepwise

    DiscariminantAnalysisProgram; a function todiscriminatebetween shapes

    foreach subject was computedand thentestedon largenumbersof single

    t r i a l s from other experiments with this subject. Finally,the A

    s t a t i s t i c as described by John et a l . (1967)was computedusing

    averages of ten t r i a l s formostof the experiments.

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    5.2.1 Resultsofvis ual analysis.

    For v is ua l analysistheevoked responses fromthirteen subjects (who

    had eachbeen run i nparadigm 1 atleastoncewitho c c i p i t a l , parietal,

    central,andvertex electrodes) wereused. Someof thesubjectswere

    testedup tofive timesi nthisparadigmbut theaveragenumber of

    runspersubjectwas two.

    Thirty t r i a l s foreach stimulus shapewereaveragedtoobtainthe

    VER's forexamination. With subjects thatweretestedmorethan

    oncetheresponseswerefoundto benearly identical fromone

    experimenttoanother. A typicalset ofresponsesto thefour shapes

    in seven channelsfor onesubject i s shown i nFig.7. Theresponses

    showedsomevariationi nlatencyandconfigurationbetweensubjects,

    butdidexhibit es sential lythesameseriesof components fora l l

    subjects. Fig.8 shows theresponseat 0 to the e lshapefora l l

    thirteen subjects.

    Ina fewsubjectsmid (T^ or T ) orpo sterio r (Tj-or T )temporalor

    frontal (F^,F^ or F^) electrodeswereused instead of therightand

    l e f t centrals. Noclearresponsewasseenatmid-temporal or frontal

    locations. Theresponseat theposteriortemporals appeared very

    similartothat seenat 0, and 0.

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    100 msec

    Fig. 7 Evokedresponsese l i c i t e d by thefour di ff er en t stirnulusshapes at the 7electrode locations used i n paradigm 1.

    n=30trials/ aver age. (Subject SC, Exp. 1)

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    45.

    LP

    msec

    Fig. 8 Intersubject v a r i a b i l i t y . Responserecordedfrom

    ^1 - A2 e l i c : i - t e d DY t b ee lshape for each of the 13 subjects.n=30trials /avera ge.

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    For further desc ript ionof theresponsesthelabellingshowni n

    Fig.7 wasperformed. The f i r s tmajorpositivepeakseenatabout

    95 msec., i n most subjects,was calledPp Thiswas sometimes

    precededby adefi ni te small negative peakcalledN . Following P^

    the nextlarge negative peak (whichwas sometimesdoublei nsome

    channels)was calledN,,, thebroaderpositivewavefollowingN 2 was

    calledP 2, and alater positivepeak (wheni toccurred)was called

    P . It wasnotedthattherelative amplitudesofthesecomponents

    variedi n theresponsestodifferentshapes andamongsubjects,but

    i twasusually possibletoidentifythem ina l lof therecorded channels

    for a l l subjects.

    For a l l subjectstheamplitudeof the P 1 component was approximately

    equali noccipitalandparietal recordings and slightly smaller than

    thisi n thecentr al leads.

    The N 2 component in six of thethirt een subjectswas largesti n the

    occipital regions andeit her reversed p ol ar it ycompletelyorbecame

    biphasic (i.e. partofi t reversed i npolarity) anter iorly. The

    reversaltookplace ei the r jus t anter iorto orposteriorto the

    parietal electrodes (asi tdoes i nFig.9). In theremainingseven

    subjects thiscomponentbecameprogressively smaller anter iorto the

    occipital locations rather thanreversingi npolarity,(asi tdoes i n

    Fig.7).

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    The ?2 conponent had abroadconfigurationand thelatencyof this

    peak wasmorevariablebetween thedifferent recording sites than

    was thelatencyof theothercomponents. Insevenofthirte en

    subjects thiswavewas ofhighest amplitudei n theoccipitaland

    parietal regions,and i n six ofthirteen i twas largesti n the

    centralorvertex channels.

    Becausethe components were most clearly definedi n theoccip ital

    regions therightandl e f t occipital channelswereusedto measure

    the latencyandamplitudeof thethreecomponents fora l lof the

    subjects. Theaveragesoft h i r t y t r i a l swereusedforthese

    measurements, althoughi nsomesubjects i twasnotedthattheaverage

    of justtent r i a l s showed the componentsequally well.

    Itwas notedthatthelatencyof thethreecomponents for a

    particular subjectwas quite consistentfora l l fourshapes.

    Table Ishowsthemeanlatenciesof thethreecomponents for the

    thirteen subjects' averageresponsesto thefour different shapes.

    Itcan beseenbyinspectiono f the magnitude of thestandard deviations

    ofthemeanvalues i nthis Table that therewere no significant

    differencesin thelatencyofthesethreecomponents between the

    different shapes. Themeanlatencyfora l l fourshapesanda l l

    thirteen subjects for P- was 95 msec, for N 2 was 148 msec, and for

    P~ was 219 msec.

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    48.

    TABLEI:

    Latenciesof thepeaksof thethree principalcomponentsofthe occipitalevoked responsesto thefourshapespresented

    inparadigm1. Eachvalue (inmsec.) i s themeanand

    standarddeviation for13subjects.

    N.2

    square 95- 17 145- 20 212- 25

    e l 92 - 20 150- 21 225- 18

    circle 101- 23 150- 26 219- 21

    omega 93- 24 148- 25 221- 24

    Mean of 4shapes 95 148 219

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    49.

    Therewerenoconsistent differencesi n thelatencyof thecomponents

    fo r any of theshapesbetweenrightandl e f t occipital regions. I f any

    small differenc ebetweenrightandl e f twasseenforagiven subject,

    themean o f the two wastakenas thevalue used fo r preparing TableI.

    The amplitudeofthese threecomponentswasmeasuredfromthe pre-

    stimulusbase l i n e . Tocomparetheamplitudeof thea c t i v i t yat

    0- and0^thesubjectsweredivid ed accordingtohandedness. There

    wereeight right-handers andfour left-handers (datafromonesubject

    wereomitted becausethe EEGfrom 0^wastechnically inadequate).

    Ameanamplitudedifference foreachof thefourshapeswascomputed

    for eachsubject. For theright-handed groupno meandiffere nces

    between0- and 0 2amplitudesofgreaterthan1 yvwereseenforany

    ofthecomponentsandthesewerenotevenconsistentlyi n the same

    direction. For thefour left-handed subjects therewas aconsisten t

    tendencyforN 2to bes l i g h t l y higher overtherighthemisphere(mean

    differenceof 2 uv)for a l l fourof theshapes.

    Themeanamplitudeofeachof thethreecomponentsrecorded at for

    the fourshapesfor12 of thesubjectsi sshowninTableII(datafrom

    one subjecthad to bediscardedbecauseoftechnicalproblems). The

    amplitudesof the N 2componentof thesquarevs. e l ,c i r c l e vs.omega

    and c i r c l e vs.e l and of the P 2componento f thesquarevs. e l ,

    c i r c l e vs.omega andsquarevs.omegawerea l lfoundto be significantly

    different usingtheScheffe's (1959)methodoftestingmult iple

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    TABLE II :

    Amplitudesof thethreeprincipalcomponentsof theresponses

    at 02to thefourshapes presentedi nparadigm1. Each value

    (inyv) i s the mean andstandarddeviationfor12 subjects.*

    P

    lN

    2P

    2

    square 2.9 2.4 6.4 4.0 6.2 3.1

    circle 3.4 -2.4 5.2 2.8 7.2 4.5

    cmega 3.8 3.1 8.1 3.8 10.5 5.5

    e l 4.8 3.4 10.4 4.8 9.5 5.0

    ByScheffe's(1959)methodoftesting contrasts(asprovidedi n the UBCComputingCenterProgramMFAV),theamplitudesof thecomponentsof thefollowingpairsofshapesweresignificantly different (p= .01):forthe Ncomponentthesquareandel ,thec i r c l eandomega and thec i r c l eande l were a l ldifferent,andfo rthe P~componentthesquareandel ,thec i r c l eand omegaandthesquareand omegapairswere a l ldifferent. For

    the P^component noneo f the6possiblepairsofshapesweresignificantly different.

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    51.

    contrasts (p = .01). Therewereno significant differences (p = .01)

    in the amplitude of the component. I t i s notedthattheScheffe's

    methodi s based on a two way analysisofvarianceand thus i t i s

    essentially testingthedifferencebetweenthe shapes for each of

    the12 subjects.

    Fortwo subjects,paradigm 1 was repeated wi th the T.V. camera lens

    adjustedsothatthe shapes appeared on the viewing screen at

    approximatelyone half theirusual size. The evoked responses to

    thesesmallerstimulus shapeswerenotdifferent (byvisual inspection)

    from the ones evoked by the standardsized shapes. The responses to

    thesmallandlargesquare, e l ,c i r c l e andomegafor one subjectare

    showni nFig. 9.

    5.2.2 Resultsof StepwiseDiscrirninant Analysis.

    Inap i l o t study with theBMD:07Mprogram a sampling of data from a l l

    five subjectsfrom a l l of therecording locations (occipitals, parietals

    and centrals)was t r i e d to determine which ones would bebestfo r

    ccnpletetesting. Using the U s t a t i s t i cof the program as an indicator

    ofhow wellthe groupswereseparated(see,Appendix B) i t was foundthat

    one of theoccipital locationsalways had the lowest U value ( i . e .

    showedthebest separation) for anypairo f groups from agivenexperiment.

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    52.

    Fig. 9 Effectofstimulus size. Responses tostandard sized

    stimuli (visualangleof 3.6) areshownassol id

    lines, those to thehalf-sizedones asdotted lines,

    (subjectML, Exp. 8,21) n=30 trials/average

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    Therefore, fortestingtheabilityofdiscriminant analysistoclas sify

    newdataforeachsubject, onlytheoccipital datawere completely

    testedandarereported here.

    TheworkwiththeBMD:07Mand thetestingof thefunctionsproducedby

    i t (withtheprogramUBC:CLASS)wasdonei n twophases. In thef i r s t

    phase,data collected fromfour subjectsi nparadigm1wereused. Single

    t r i a l s froma l lfour stimulus shapesweresubmittedto theBMD:07Mprogram

    at once,hencetheresulting discriminant functionswereintendedto

    classifyanysingle t r i a l (orobservation)asbelongingto one of the

    fourpossiblegroups (shapes). In thesecondphasethedata fromonly

    twostimulus shapesat atimeweresubmittedto theprogram.

    In TableIIItheresultsoftestingthediscriminant functions

    classifying a l l fourshapesforthefour subjectsondata fromrightand

    l e f t occipital locationsarepresented. Thepercentages giveni n the

    Tablearetheproportionsof thetotalnumberoft r i a l sthefunction

    correctly classified. Theclassificationsmade on thet r i a l s usedto

    computethefunction (i.e.ap o s t e r i o r i c l a s s i f i c a t i o n )wereincluded

    inthepercentages andtotalnumbersoft r i a l s giveni nthis Table.

    120 t r i a l s ,30 ofeach shape,wereusedtocomputeeachfunction.

    Itcan beseen fromtheTable thatthefunctionsclassifiedfrom35 to 44%

    of the waveformsfromthesingle t r i a l s correctly, proportionswhichare

    significantly greater thanthelevelof 25%expected witharandom

    c l a s s i f i c a t i o n (z =3.58f or35%whichi sgreater thanzn 1 =2.57).

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    TABLEII I:

    Performanceof the "Fourshape"Discriminant Functions.

    Percentageof t r i a l s correctly c l a s s i f i e d by theBMD:07M

    StepwiseDiscriminant Analysis technique on data from right

    and l e f t occipital locations. Totalnumberof trials

    c l a s s i f i e d foreach subject i s given in parentheses. A l l

    percentages given are significantly* greater than the 25%

    level expected with arandom c l a s s i f i c a t i o n (p = .01).

    For 35% z = 3.58 whichi s greater than z m = 2.57.

    Subject 0 2 (right) 0-j (left)

    MP (260) 41% 44%

    SC (480) 37% 37%

    VS (240) 39% 40%

    ML (360) 35% 41%

    meanof 4 subjects 38% 40%

    * significancedeterminedby evaluating the test statisticz = p - p//pq/n-wherep i s the observed proportion,

    p i s the expected proportion, q = 1 - p and n i s thesamplesize, z i s considered to be normally distributed.(Bahn, 1972).

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    TABLEII I:

    Performanceof the "Fourshape"Discriminant Functions.

    Percentageof t r i a l s correctly c l a s s i f i e d by theBMD:07M

    Stepwise Discriminant Analysis technique on data from right

    and l e f t occipital locations. Totalnumberof trials

    c l a s s i f i e d foreach subject i s given i n parentheses.

    Subject 0 2(right) O-^left)

    MP (260) 41% 44%

    SC (480) 37% 37%

    .VS (240) 39% 40%

    ML (360) 35% 41%

    meanof 4 subjects 38% 40%

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    55.

    To ascertainwhichof thefourshapesthefunctionswerebest ableto

    identify,thepatternof theincorrect classificationswas studied

    (i.e.whentheresponse fromoneshapewas c l a s s i f i e d