the retina overview of the visual system perception &...

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1 Perception & Attention Perception is effortless but its underlying mechanisms are incredibly sophisticated. Biology of the visual system Representations in primary visual cortex and Hebbian learning Object recognition Attention: Interactions between systems involved in object recognition and spatial processing 2 Perception & Attention Some motivating questions: 1. Why does primary visual cortex encode oriented bars of light? 2. Why is visual system split into what/where pathways? 3. Why does parietal damage cause attention problems (neglect)? 4. How do we recognize objects (across locations, sizes, rotations with wildly different retinal images)? 3 Overview of the Visual System Hierarchies of specialized visual pathways, starting in retina, to LGN (thalamus), to V1 & up: optic chiasm temporal temporal nasal LGN V1 V2,V4... V2,V4... right field left field 4 Two Streams: Ventral “what” vs. Dorsal “where” V1 V2 V3 V4 TEO TF TE PO V3A MT FST MST VIP PG PG TE V1 p m m d 5 The Retina Retina is not a passive “camera” Key principle: contrast enhancement that emphasizes changes over space & time. - + - - - - + + + + a) On-center b) Off-center + - - + - + - + - + retinal output ganglion cells 6 LGN of the Thalamus A “relay station”, but so much more. Organizes different types of information into different layers. Performs dynamic processing: magnocellular motion processing cells, attentional processing. On- and off-center information from retina is preserved in LGN

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  • 1Percep

    tion&

    Atten

    tion

    Percep

    tioniseffo

    rtlessbutits

    underly

    ingmech

    anism

    sare

    incred

    ibly

    sophisticated

    .

    •Biologyofthevisu

    alsystem

    •Rep

    resentatio

    nsin

    prim

    aryvisu

    alcortex

    andHeb

    bian

    learning

    •Object

    recognitio

    n

    •Atten

    tion:Interactio

    nsbetw

    eensystem

    sinvolved

    inobject

    recognitio

    nan

    dsp

    atialprocessin

    g

    2Percep

    tion&

    Atten

    tion

    Somemotiv

    atingquestio

    ns:

    1.Whydoes

    prim

    aryvisu

    alcortex

    encodeorien

    tedbars

    oflig

    ht?

    2.Whyisvisu

    alsystem

    split

    into

    what/

    where

    path

    way

    s?

    3.Whydoes

    parietal

    dam

    agecau

    seatten

    tionproblem

    s(neg

    lect)?

    4.How

    dowereco

    gnize

    objects

    (across

    locatio

    ns,sizes,ro

    tations

    with

    wild

    lydifferen

    tretin

    alim

    ages)?

    3Overv

    iewoftheVisu

    alSystem

    Hierarch

    iesofsp

    ecializedvisu

    alpath

    way

    s,startingin

    retina,to

    LGN

    (thalam

    us),to

    V1&

    up:

    opticchiasm

    temporal

    temporal

    nasal

    LGN

    V1

    V2,V

    4...

    V2,V

    4...

    right fieldleft field

    4TwoStream

    s:Ven

    tral“w

    hat”

    vs.Dorsal

    “where”

    V1

    V2V

    3V

    4T

    EO

    TF

    TE

    PO

    V3A

    MT

    FS

    T

    MS

    T

    VIP

    PGP

    G

    TE

    V1

    p

    mm

    d

    5TheRetin

    a

    Retin

    aisnotapassiv

    e“cam

    era”

    Key

    prin

    ciple:

    contrast

    enhan

    cemen

    tthat

    emphasizes

    chan

    ges

    over

    space

    &tim

    e.

    −+−

    −−

    −+

    +

    +

    +

    a) On−

    centerb) O

    ff−center

    +

    −−

    +−

    +

    − +

    +

    retinal

    outputgan

    glio

    ncells

    6LGN

    oftheThalam

    us

    A“relay

    station”,b

    utso

    much

    more.

    •Organ

    izesdifferen

    ttypes

    ofinform

    ationinto

    differen

    tlay

    ers.

    •Perfo

    rmsdynam

    icprocessin

    g:mag

    nocellu

    larmotio

    n

    processin

    gcells,atten

    tional

    processin

    g.

    •On-an

    doff-cen

    terinform

    ationfro

    mretin

    aispreserv

    edin

    LGN

  • 7Prim

    aryVisu

    alCortex

    (V1):

    EdgeDetecto

    rs

    V1combines

    LGN

    (thalam

    us)

    inputsinto

    orien

    teded

    gedetecto

    rs:

    −+−

    −−

    −+

    +

    +

    +

    a) On−

    centerb) O

    ff−center

    +

    −−

    +−

    +

    − +

    +

    −−

    −−

    +−−

    −−

    +−−

    −−

    +−−

    −−

    +

    +−

    •Edges

    differ

    inorien

    tation,size

    (spatial

    frequen

    cy),an

    d

    positio

    n.

    •Forcoheren

    tvisio

    n,need

    todetect

    vary

    ingdeg

    reesofall

    these.

    8Prim

    aryVisu

    alCortex

    (V1):

    EdgeDetecto

    rs

    V1combines

    LGN

    (thalam

    us)

    minputsinto

    orien

    teded

    ge

    detecto

    rs :

    −+−

    −−

    −+

    +

    +

    +

    a) On−

    centerb) O

    ff−center

    +

    −−

    +−

    +

    − +

    +

    −−

    −−

    +−−

    −−

    +−−

    −−

    +−−

    −−

    +

    +−

    V1

    edgedetector

    •Edges

    differ

    inorien

    tation,size

    (spatial

    frequen

    cy),an

    dpositio

    n.

    •Forcoheren

    tvisio

    n,need

    todetect

    vary

    ingdeg

    reesofall

    these.

    9Prim

    aryVisu

    alCortex

    (V1):

    Topograp

    hy

    LR

    LR

    4 2−3 blobs

    orientations

    occularity

    hypercolumns

    Pinwheel

    Hyperco

    lumn:Fullset

    ofcodingforeach

    positio

    nPinwheel

    can

    arisefro

    mlearn

    ingan

    dlateral

    connectiv

    ity:nothard

    -wired

    !

    10Rero

    utin

    gofVisu

    alInfo

    toAudito

    ryCortex

    •Sharm

    a,Angelu

    cci&

    Sur(2000),N

    ature

    Rero

    uted

    fibers

    from

    Retin

    a→

    audito

    rythalam

    us(M

    GN)→

    A1

    •Ifvisu

    alproperties

    arelearn

    ed,th

    eysh

    ould

    dev

    elopin

    A1.

    11Rero

    utin

    gofVisu

    alOrien

    tationModules

    inA1

    Ba-d

    :Orien

    tationmap

    s,dark

    -highact

    forgiven

    orien

    tation

    (botto

    m

    right).

    C:co

    mposite

    map

    oforien

    tationpreferen

    ces

    D:red

    dots=pinwheel

    centers

    12Visu

    alBeh

    aviorAfter

    Rero

    utin

    gRightVisu

    alField

    vonMelch

    ner,

    Pallas

    &Sur(2000)

  • 13Visu

    alAcu

    ityAfter

    Rero

    utin

    g

    →Solearn

    ingis

    powerfu

    l,butso

    isev

    olutio

    n!

    14A

    Questio

    n

    What

    mak

    esvisu

    alcortex

    visu

    alcortex

    ?Whydoes

    itrep

    resent

    orien

    tedbars

    oflig

    ht?

    15Prim

    aryVisu

    alRep

    resentatio

    ns

    Key

    idea:

    Orien

    teded

    gedetecto

    rscan

    dev

    elopfro

    mHeb

    bian

    correlatio

    nal

    learningbased

    onnatu

    ralvisu

    alscen

    es.

    16TheModel:

    Sim

    ulatin

    goneHyperco

    lumn

    •Natu

    ralvisu

    alscen

    esare

    prep

    rocessed

    bypassin

    gthem

    (separately

    )throughlay

    ersofon-cen

    teran

    doff-cen

    terinputs

    •Hidden

    layer:

    edgedetecto

    rsseen

    inlay

    ers2/

    3ofV1;

    Lay

    er4(in

    put)just

    represen

    tsunorien

    tedon/offinputs

    likeLGN

    (butcan

    bemodulated

    by

    attentio

    n)

    17TheModel:

    Sim

    ulatin

    goneHyperco

    lumn

    •Heb

    bian

    learningonly

    •KWTA

    inhib

    competitio

    nforsp

    ecialization(see

    Ch4)

    18[v1rf.p

    roj]

  • 19TheRecep

    tiveField

    s0

    12

    34

    56

    78

    910

    1112

    13

    Red

    =on-cen

    ter>

    off-cen

    ter,Blue=off-cen

    ter>

    on-cen

    ter

    20

    2122

    23Somedifferen

    ces,butpinwheels

    stillem

    erge

    24Percep

    tionan

    dAtten

    tion

    1.Whydoes

    prim

    aryvisu

    alcortex

    encodeorien

    tedbars

    oflig

    ht?

    Correlatio

    nal

    learningbased

    onnatu

    ralvisu

    alscen

    es.

    Refl

    ectsreliab

    lepresen

    ceofed

    ges

    innatu

    ralim

    ages,w

    hich

    vary

    in

    size,positio

    n,orien

    tationan

    dpolarity.

    →model

    showshow

    docu

    men

    tedV1properties

    canresu

    ltfro

    m

    interactio

    nsbetw

    eenlearn

    ing,arch

    itecture

    (connectiv

    ity),an

    d

    structu

    reofen

    viro

    nmen

    t.

  • 25Percep

    tionan

    dAtten

    tion

    1.Whydoes

    prim

    aryvisu

    alcortex

    encodeorien

    tedbars

    oflig

    ht?

    Correlatio

    nal

    learningbased

    onnatu

    ralvisu

    alscen

    es.

    2.How

    dowereco

    gnize

    objects

    (across

    locatio

    ns,sizes,ro

    tations

    with

    wild

    lydifferen

    tretin

    alim

    ages)?

    3.Whyisvisu

    alsystem

    split

    into

    what/

    where

    path

    way

    s?

    4.Whydoes

    parietal

    dam

    agecau

    seatten

    tionproblem

    s(neg

    lect)?

    26TheObject

    Reco

    gnitio

    nProblem

    Problem

    :Reco

    gnize

    object

    regard

    lessof:locatio

    n,size,ro

    tation.

    Sam

    e

    Diff

    Thisishard

    becau

    sedifferen

    tpattern

    sin

    samelocatio

    ncan

    overlap

    alot,w

    hile

    thesam

    epattern

    sin

    differen

    tlocatio

    ns/

    sizes/rotatio

    ns

    cannotoverlap

    atall!

    2728 G

    radual

    Invarian

    ceTran

    sform

    ations(Fukush

    ima,’80)

    Increasin

    grecep

    tivefield

    sizeen

    ables:

    Conjunctio

    noffeatu

    res(to

    form

    more

    complex

    objects);

    and

    Collap

    singover

    locatio

    ninform

    ation(“sp

    atialinvarian

    ce”)

    29 Grad

    ual

    Invarian

    ceTran

    sform

    ations(Fukush

    ima,’80)

    ifdid

    spatial

    invarian

    cein

    onefell

    swoop:bindingproblem

    -can

    ’ttell

    Tfro

    mL

    30 Grad

    ual

    Invarian

    ceTran

    sform

    ations(Fukush

    ima,’80)

    Goal:

    Units

    atthetopofthehierarch

    ysh

    ould

    represen

    tcomplex

    object

    features

    inalocatio

    nan

    dsize

    invarian

    tfash

    ion

    (alsowan

    tben

    efits

    oftop-downam

    plifi

    cation,pattern

    completio

    n,d

    istributed

    repsetc)

  • 31TheModel

    LGN

    _On

    LGN

    _Off

    V1 V2

    V4/IT

    Output

    V1=orien

    tedlin

    e(ed

    ge)

    detecto

    rs,hard

    -coded

    V2units

    encodeconjunctio

    nsofV1ed

    ges

    across

    asu

    bset

    ofsp

    ace

    Each

    V4unitpay

    satten

    tionto

    allofV2

    32TheObjects

    01

    23

    4

    56

    78

    9

    1011

    1213

    14

    1516

    1718

    19

    Each

    object

    ispresen

    tedat

    multip

    lelocatio

    ns,sizes

    Netw

    ork’sjobisto

    activate

    theap

    propriate

    Outputunit(0-19)

    for

    eachobject,reg

    ardless

    oflocatio

    nan

    dsize

    33[objrec.p

    roj]

    34

    3536

  • 3738

    3940

    4142

  • 4344

    Gen

    eralization

    •Can

    thenetw

    ork

    gen

    eralizeto

    unseen

    view

    sofstu

    died

    objects?

    •In

    other

    words:Does

    trainingthenet

    toreco

    gnize

    aset

    of

    objects

    inasize/

    locatio

    ninvarian

    tfash

    ionhelp

    itreco

    gnize

    new

    objects

    inasize/

    locatio

    ninvarian

    tfash

    ion?

    •Proced

    ure:

    –Tak

    eanet

    trained

    on18

    objects

    –Train

    with

    2new

    objects

    inonly

    somelocatio

    ns/

    sizes

    –Test

    thenet

    with

    nonstu

    died

    “view

    s”(sizes/

    locatio

    ns)

    of

    new

    objects

    4546

    Gen

    eralization

    •Can

    thenetw

    ork

    gen

    eralizeto

    unseen

    view

    sofstu

    died

    objects?

    yes

    •Approx.75%

    correct

    onnovel

    view

    sfollo

    wingtrain

    ingon10%

    ofpossib

    lesizes/

    locatio

    nsExplan

    ation:Distrib

    uted

    represen

    tationsan

    dHeb

    blearn

    ing!

    •V4rep

    resents

    objectfeatures

    inalocatio

    n/size

    invarian

    tway

    •Each

    object

    activates

    adistrib

    uted

    pattern

    ofthese

    invarian

    t

    feature

    detecto

    rs

    4748

  • 4950

    5152

    5354

    01

    23

    4

    56

    78

    9

    1011

    1213

    14

    1516

    1718

    19

    Yeah

    ,butthese

    objects

    arereg

    ularly

    shap

    ed,straig

    htlin

    es...

    what

    aboutreal

    objects?

  • 55

    56“E

    mer”

    therobotreco

    gnizin

    gobjects..

    Video

    Dem

    o

    57

    O’Reilly,

    CogSci2009

    58why?

    bidirconssu

    pportattracto

    rsacro

    ssmultip

    lelev

    elsofnet

    toam

    plify

    consisten

    t

    info

    59A

    Challen

    ge

  • 60State

    oftheArt

    61Still

    missin

    g...

    Motio

    n

    •Neu

    ronsin

    areaMTvery

    sensitiv

    eto

    motio

    n

    •Lotsofwork

    onhow

    downstream

    areasinteg

    ratemotio

    nsig

    nals

    across

    timeto

    detect

    coheren

    ce(e.g

    .Shad

    len,

    New

    some,etc)

    •Thomas

    Serre

    has

    shownthat

    motio

    nsig

    nals

    very

    reliable

    for

    discrim

    inatin

    gbetw

    eenparticu

    laractio

    ns(eg

    throwinga

    baseb

    all)

    •Should

    beab

    leto

    solveproblem

    via

    bidirectio

    nal

    influen

    ceof

    motio

    ninteg

    rationsig

    nals,o

    bject

    recognitio

    n,an

    dsp

    atialatten

    tion(next)....

    62Percep

    tionan

    dAtten

    tion

    1.Whydoes

    prim

    aryvisu

    alcortex

    encodeorien

    tedbars

    oflig

    ht?

    Correlatio

    nal

    learningbased

    onnatu

    ralvisu

    alscen

    es.

    2.How

    dowereco

    gnize

    objects

    (across

    locatio

    ns,sizes,ro

    tations

    with

    wild

    lydifferen

    tretin

    alim

    ages)?

    Tran

    sform

    ations:

    increasin

    gly

    complex

    featural

    encodings,in

    creasinglev

    elsof

    spatial

    invarian

    ce;Distrib

    uted

    represen

    tations.

    3.Whyisvisu

    alsystem

    split

    into

    what/

    where

    path

    way

    s?

    4.Whydoes

    parietal

    dam

    agecau

    seatten

    tionproblem

    s(neg

    lect)?

    63Spatial

    Atten

    tion:Unilateral

    Neg

    lect

    Patien

    tcopyinga

    scene

    Self

    portrait,

    copying,

    linebisectio

    ntask

    s:In

    allcases,p

    atientswith

    parietal/

    temporal

    lesionsseem

    toforget

    about1/

    2ofsp

    ace!butthey

    stillsee

    it!

    6465

  • 6667

    6869

    7071

  • 7273

    74Effects

    ofParietal

    Lesio

    nsonPosn

    erTask

    40−

    60−

    80−

    100−

    120−01

    2

    Intact

    Lesioned

    Neutral

    Valid

    Invalid

    •Patien

    tsperfo

    rmnorm

    allyin

    the“n

    eutral”

    (nocu

    e)conditio

    n,

    regard

    lessofwhere

    thetarg

    etispresen

    ted

    •Patien

    tsben

    efitjustas

    much

    ascontro

    lsfro

    mvalid

    cues

    •Patien

    tsare

    hurtmore

    than

    contro

    lsbyinvalid

    cues

    75Possib

    leModels

    +

    Alert

    Interrupt

    Localize

    Disengage

    Move

    Engage

    Inhibit

    Object

    V1

    (features x location)

    Spatial

    Atten

    tionem

    erges

    from

    bidirectio

    nal

    constrain

    tsatisfactio

    n&

    inhibito

    rycompetitio

    n.

    76Sim

    ple

    Model

    Input

    V1

    Spat1

    Spat2

    Obj1

    Obj2

    targ

    cue

    Output

    Object 1 (C

    ue)

    Object 2 (T

    arget)

    77[attn

    simple.p

    roj]

  • 78Posn

    erTask

    Data

    Valid

    Invalid

    Diff

    AdultNorm

    al350

    39040

    Elderly

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    al540

    60060

    Patien

    ts640

    760120

    Elderly

    norm

    alized(*.65)

    350390

    40Patien

    tsnorm

    alized(*.55)

    350418

    68

    79Posn

    erTask

    Sim

    s

    •Themodel

    explain

    sthebasic

    findingthat

    valid

    cues

    speed

    target

    processin

    g,while

    invalid

    cues

    hurt

    •Also

    explain

    sfindingthat

    patien

    tswith

    small

    unilateral

    parietal

    lesionsben

    efitnorm

    allyfro

    mvalid

    cues

    inipsilateral

    field

    butare

    disp

    roportio

    nately

    hurtbyinvalid

    cues.

    •Noneed

    toposit

    “disen

    gag

    e”module!

    •Also

    explain

    sfindingofneglect

    ofcontralateral

    visu

    alfield

    afterlarg

    e,unilateral

    parietal

    lesionswhen

    somestim

    ulusis

    presen

    tin

    ipsilateral

    field

    (“extin

    ction”)

    80More

    Posn

    erLesio

    nFun

    •Retu

    rningto

    patien

    twith

    leftparietal

    lesion...

    •What

    hap

    pen

    sifcu

    esare

    presen

    tedin

    contralateral(affected

    )

    hem

    ifield

    ?(“R

    everse

    Posn

    er”)

    81More

    Posn

    erLesio

    nFun

    Retu

    rningto

    patien

    twith

    leftparietal

    lesion...

    •What

    hap

    pen

    sifcu

    esare

    presen

    tedin

    contralateral(affected

    )hem

    ifield

    ?

    Pred

    ictions:

    •Smaller

    ben

    efitforvalid

    cues

    •Patien

    tssh

    ould

    behurtless

    than

    contro

    lsbyinvalid

    cues.

    82[attn

    simple.p

    roj]

    83Inhibitio

    nofRetu

    rn

    •Typically,targ

    etdetectio

    nisfaster

    ontrials

    with

    valid

    vs

    invalid

    cues

    •H

    owever

    ,ifthecu

    eispresen

    tedforalonger

    time(eg

    .500

    ms),

    perfo

    rman

    ceisfaster

    on

    invalidvsvalid

    trials

    •Can

    explain

    interm

    sofaccom

    modation

    (neu

    ralfatig

    ue)

  • 84[attn

    simple.p

    roj]

    85Sim

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    86Complex

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    87Percep

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