The model

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The model

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Title: The model


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The model
good
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Cortical Circuitry
Feedforward intracortical connections V1 (II/III)
to V2 (IV)
Feedback intracortical and subcortical
connections V2 (VI) to V1 (VI) V1 (V) to SC, V1
(VI) to LGN, V2 (V) to SC, Pons, Striatum
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Histology of Cerebral Cortex 2
  • Pyramidal neurons are large and complex
  • Similar orientation
  • Process input from many sources

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Stuff and things
  • Retina, lateral geniculate nucleus and primary
    visual cortex produce rich information about
    local points,
  • but properties of more global objects are not
    represented (stuff versus things).

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Color and motion
  • Need to determine what goes together to represent
    a thing. How does it move? What is its true
    color? This takes place in extrastriate cortex.
    Farah begins consideration of global (rather than
    local) image perception with color and motion.

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Color and motion areas
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Color
  • Color perception begins with wavelength detection
    in the retina. Color contrast is enhanced by
    center-surround receptive fields in retina and
    LGN. Double opponent effects occur in blob cells
    of Layers 2 and 3 in visual cortex and
    color-selective responses continue in the thin
    cytochrome oxidase stripes of V2 and are
    projected to V4. Up to V4, responses are
    wavelength-selective rather than color selective.
    To be color selective means context can influence
    color response.

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Border of V1/V2 with blobs and stripes
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Color constancy
  • Our perception of the color of an object is based
    on the light reflected back to us from that
    object. That light depends on both the spectral
    reflectance (true color) AND the spectral
    composition of the incident light (light that
    bathes the object). We perceive color accurately
    because we can take the color of the incident
    light into account. This ability is called color
    constancy. The larger context helps us here.

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Mondrian colors
  • Land (who invented the Land camera or Polaroid)
    showed that Mondrian colors are perceived
    accurately if the whole thing is bathed in the
    same light but if one light is used on a patch
    and another light on the next patch then we cant
    see color accurately.

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  • In the rosy light of dawn, for instance, a yellow
    lemon will reflect more long-wave light and
    therefore might appear orange but its
    surrounding leaves also reflect more long-wave
    light. The brain compares the two and cancels out
    the increases.

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V4 had color constancy
  • Zeki, recording from visual pathway neurons in
    monkeys, showed that while areas up to V2 only
    see light from a patch and cant compensate for
    the context light overall, neurons in V4 are able
    to compensate for the incident light and
    accurately report the color!

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V4 receptive fields support color constancy
  • These neurons have large receptive fields, even
    extending into the other visual field.
  • The surrounds are inhibitory and sensitive to the
    same wave length as the center thus if the same
    wavelength is everywhere, it is discounted or
    dismissed (psychologically speaking) as ambient
    light.
  • This is consistent with evidence that a split
    brain patient has trouble with color constancy
    for stimuli that cross the midline since he/she
    cant know what the overall light is on both
    sides at once.

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Cerebral achromatopsia
  • Lost of color vision or color blindness when
    acuity, motion, depth perception and object
    recognition are good is called achromatopsia.
  • In some cases the other visual functions are
    transiently affected too or pattern recognition
    may also be a problem. Cases involving artists
    are particularly dramatic.
  • Unilateral lesions can create loss in only one
    hemifield hemiachromotopsia.

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"... as soon as he entered, he found his entire
studio, which was hung with brilliantly colored
paintings, now utterly grey and void of color.
His canvases, the abstract color paintings he was
known for, were now greyish or black and white.
His paintings--once rich with associations,
feelings, meanings--now looked unfamiliar and
meaningless to him. At this point the magnitude
of his loss overwhelmed him. "He had spent his
entire life as a painter now even his art was
without meaning, and he could no longer imagine
how to go on. Oliver Sacks, The Case of the
Colorblind Painter, 1995      In An
Anthropologist On Mars, p.6
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  • Achromatopsia is produced by lesions on the
    inferior surface of temporo-occipital regions,
    (lingual and fusiform gyri).

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Other color related disorders
  • color anomia problem in producing the names of
    colors
  • color agnosia, loss of knowledge about colors
    (hard to define, cant learn paired associates
    where one word is a color and the other is a name
    or number)
  • impaired color-object association, cant tell
    the typical colors of things.

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Color anomia
  • Color anomia, damage to the temporal segment of
    the left lingual gyrus, prevents you from
    naming colors even though you can match and
    discriminate between colors non-verbally. So,
    there seems to be a very specific locus where
    color perception gets coded as color language,
    but the perception can go on normally even when
    the ability to encode the perception as language
    is destroyed

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Neuroimaging and color
  • Positron emission tomography (PET) and
    event-related potentials (ERPs) are consistent
    with lesions. PET showed activation in
    lingual/fusiform region when colored Mondrians
    rather than gray scale Mondrians were presented.
  • When attending to color, rather than multiple
    stimuli, activation was seen in collateral sulcus
    (between lingual and fusiform gyri) and also
    dorsolateral occipital cortex (new area).

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PET Imaging
Upper row Control PET scans (resting while
looking at static fixation point is subtracted
from looking at a flickering checkerboard
stimulus positioned 5.5 from fixation
point). Middle row Subtraction produces a
somewhat different image for each of 5
subjects. Bottom row The 5 images are averaged
to eliminate noise, producing the image at the
bottom.
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PET Identification of Inferior Occipital Region
Activated by Color
Activation produced by staring at colored
stimuli. Panel A shows the blood flow images
before subtraction. Panel B show activation after
subtracting responses to the gray stimuli. Panel
C depicts statistical significance of the
responses. White is highest significance. Panel D
shows the location of the most significant
responses in a sagittal, coronal, and axial view
(Courtesy of Frackowiak and Zeki).
Multicolor abstract display (top) and version of
the same display in shades of gray (bottom) used
as stimuli
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Event related potentials
ERPs are a non-invasive method of measuring brain
activity. Weak electrical fields, representing
the activity of neural populations within the
brain, can be detected at the scalp using
electrodes connected to an amplifier. The
amplifier enhances the electrical signal so that
it can be reliably recorded. This signal is
time-locked to an event, such as the presentation
of a stimulus (like a word or picture) or
production of response (like a button press),
then averaged to reveal changes in brain activity
specifically associated with different aspects of
cognitive processing.  The temporal precision
of ERPs is superior to all other currently
available neuroimaging techniques.
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  • ERPs were examined using checkerboard patterns in
    color after adapting to same color or different
    color. There is a different ERP response to
    different colors in lingual and fusiform gyri and
    in dorsolateral occipital cortex. Also using
    electrical stimulation in those same places could
    alter color perceptions.

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Control issues
  • Another study found the classic area
    (lingual/fusiform area) and also widespread
    activation in other regions, but may not have
    been well controlled. They used color random
    noise patterns, judging if mostly red, versus
    black and white random noise, judging if mostly
    white. One versus 4 colors one versus 2 for
    black and white also stimuli were not matched
    for luminance.

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A good fit for V4 as the color area, but some
cautions
  • In monkey and man color areas are different in
    location, monkey V4 is high up on lateral
    surface. In man color center is on the inferior
    surface and more medial. But that may be ok.
  • V4 also inputs to object or form areas of
    temporal lobe and cells in V4 have some response
    to form.
  • Finally, monkeys with V4 lesions can relearn
    color discrimination tasks, but have permanent
    problems with form discrimination. Same form
    tasks ok in human achromatopsics. Need to do
    imaging studies with monkeys.

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V4 Color Caveat
  • V4 has been suggested to be the color center by
    Zeki. It is more involved with perception of
    colors than other areas, but it may not be both
    necessary and sufficient for color perception or
    have no other role than color perception

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And please let Mom, Dad, Rex, Ginger, Tucker, me
and all the rest of the family see color.
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Motion perception
  • Local measurement of motion is also ambiguous.
    You need to look at global indices (multiple
    local pieces of information). In early areas,
    most neurons respond better to moving than
    stationary stimuli (habituation).
  • M cells are optimized for movement and provide
    input to first direction selective cells (striate
    visual cortex, 4B). They project to the middle
    temporal area (MT) and thick cytochrome oxidase
    stripes of V2 that then project to MT.

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From local to globalINTEGRATION
Small receptive fields (input) LOCAL information
  • (output) GLOBAL perception of motion

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Aperture problem
  • With local view cant tell which way edge is
    really moving when it passes through the local
    field, as several patterns of motion look the
    same to a small window.
  • Need to combine information. Plaid patterns have
    been used to test for cells that can extract
    global, rather than local, motion.

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The motion center MT
  • V1 cells respond to local or component motion of
    plaid while MT cells can also respond to pattern
    or global motion. MT cells have larger receptive
    fields. MT projects to the medial superior
    temporal area (MST), which has cells with even
    larger receptive fields and more complex motion
    detectors like flow field properties of shinking,
    enlarging, rotation and translation.

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Compare psychophysics and cells
  • Discriminate dot motion when varying proportions
    of dots are moving consistently and others
    randomly. More consistent dots, the easier the
    discrimination.
  • Monkey performance and performance of direction
    selective cells in MT was more or less the same.
    If task is at threshold for making the correct
    discrimination, when the cells are correct, the
    monkey is correct.

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Necessary and sufficient
  • If you are working with random dots and stimulate
    a column of cells for a particular direction then
    the monkey will judge that that is the direction
    the dots are moving. Thus MT activity causes
    motion perception.
  • Newsome lesioned the MT with ibotenic acid.
    Monkeys were impaired in motion discrimination in
    the contralateral hemifield, but not for color,
    acuity or depth. (Recovery issues?)

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