Title: The model
1The model
good
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3Cortical 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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6Histology of Cerebral Cortex 2
- Pyramidal neurons are large and complex
- Similar orientation
- Process input from many sources
7Stuff 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).
8Color 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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11Color and motion areas
12Color
- 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.
13Border of V1/V2 with blobs and stripes
14Color 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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18Mondrian 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.
19- 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.
20V4 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!
21V4 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.
22Cerebral 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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25 "... 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, 1995Â Â Â Â Â In An
Anthropologist On Mars, p.6
26- Achromatopsia is produced by lesions on the
inferior surface of temporo-occipital regions,
(lingual and fusiform gyri).
27Other 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.
28Color 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
29Neuroimaging 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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31PET 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.
32PET 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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34Event 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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37- 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.
38Control 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.
39A 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.
40V4 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
41And please let Mom, Dad, Rex, Ginger, Tucker, me
and all the rest of the family see color.
42Motion 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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44From local to globalINTEGRATION
Small receptive fields (input) LOCAL information
- (output) GLOBAL perception of motion
45Aperture 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.
46The 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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48Compare 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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51Necessary 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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