review, the visual and oculomotor systemsdspace.mit.edu/bitstream/handle/1721.1/91561/9-04-fall...v2...
TRANSCRIPT
The visual and oculomotor systems Peter H. Schiller, year 2006
Review,the visual and oculomotor systems
Basic wiring of the visual system
or
left h
e
fix
Primates
Horopter (Vieth-Muller circle)
temporal nasal
chiasmoptic
terminal nuclei
supericolliculi
NOT
LGN m
p
PARIETAL LOBE
FRONTAL LOBE
misphe
re right hemisphere
TEMPORAL LOBE
V1
V2
V3
V4
MT
hemifield left right
hemifield
Retina and LGN
ON OFF ONbipolars
ON OFF
ganglion cells
H
amacrine
AIIamacrine
AII
ON OFF bipolars
pigment epitheliumpigment epithelium
receptors
incoming light
photo-
conesrods
to CNS
cone horizontal
IPL
OPL
rods
to CN
receptors
ON
ON OFF
ganglion cells
H
S
cone horizontal
incoming light
photo-
cones
Visual cortex
Transforms in V1
Orientation
Direction
Spatial Frequency
Binocularity ON/OFF Convergence
Midget/Parasol Convergence
Original Hubel-Wiesel "Ice-Cube" Model
Left Eye Right Eye
Left Eye Right Eye
Swirl Model
Sub-cortical
Cortical
Radical Model
MidgetParasol
Three models of columnar organisation in V1
1 m
m
Figure by MIT OCW.
Striate Cortex Output Cell
Intracortical
Midget ON Midget ON
LEFT EYE Midget OFF Midget OFF RIGHT EYE INPUT INPUT
Parasol ON Parasol ON Parasol OFF Parasol OFF
luminance color
orientation spatial frequency
depth motion
The ON and OFF Channels
The receptive fields of three major classes of retinal ganglion cells
OFF
inhibition
ON
inhibition
ON/OFF
inhibition
Action potentials discharged by an ON and an OFF retinal ganglion cell
Stimulation confined to receptive field center
ON cell
OFF cell
Stimulation of the entire receptive field
ON cell
OFF cell
light spot dark spot
light stimulation condition
time
The midget and parasol channels
MIDGET SYSTEM PARASOL SYSTEM
ON OFFON OFF
neuronal response profile
time
P
Midget
1V
MT
Midget
Mixed
Parasol
w
?
Parasol
LGNM
PARIETAL LOBE TEMPORAL LOBE
2
V4
V2V
Projections of the midget and parasol systems
H
H
H
H
L
L
L
L
Proc
essi
ng C
apac
ity
Spatial Frequency
Temporal Frequency
Parasol SystemMidget System
Figure by MIT OCW.
Color vision and adaptation
Basic facts and rules of color vision
1. There are three qualities of color: hue, brightness, saturation
2. There is a clear distinction between the physical and psychological attributes of color: wavelength vs. color, luminance vs. brightness.
3. Peak sensitivity of human photoreceptors (in nanometers): S = 420, M = 530, L = 560, Rods = 500
4. Grassman's laws: 1. Every color has a complimentary which when mixed propery yields gray. 2. Mixture of non-complimentary colors yields intermediates.
5. Abney's law: The luminance of a mixture of differently colored lights is equal to the sum of the luminances of the components.
6. Metamers: stimuli producing different distributions of light energy that yield the same color sensations.
Basic facts about light adaptation
1. Range of illumination is 10 log units. But reflected light yields only a 20 fold change (expressed as percent contrast).
2. The amount of light the pupil admits into the eye varies over a range of 16 to 1. Therefore the pupil makes only a limited contribution to adaptation.
3. Most of light adaptation takes place in the photoreceptors.
4. Any increase in the rate at which quanta are delivered to the eye results in a proportional decrease in the number of pigment molecules available to absorb those quanta .
5. Retinal ganglion cells are sensitive to local contrast differences, not absolute levels of illumination.
The color circle
whitewhite
Yellow
Green
Red
hueBlue
saturation
black
Response to Different Wavelength Compositions in LGN Blue ON cell Yellow ON cell
90 90
30 40 0
45135
225 315
270
50 40 60 80
45135
180
225 315
270
Spikes per Second
20 1006010 20
20 0
10
45
90
135
225 315
90
135
20 30 40
45
180
225 315 maintained discharge rate
10
Green OFF cell Red ON cell
5030 40
0180
0180
270 270
Depth perception
Cues used for coding depth in the brain
Oculomotor cues Visual cues
accommodation Binocular vergence
stereopsis
Monocular motion parallax
shading interposition
size perspective
stereo camera
MOTION PARALLAX, the eye tracks
object motion
1
2
a
b
1
2
a
b eye movement
The eye tracks the circle, which therefore remains stationary on the fovea
Objects nearer than the one tracked move at greater velocities on the retinal surface than objects further; the further objects actually move in the opposite direction on the retina.
Form perception
Three general theories of form perception:
1. Form perception is accomplished by neurons that respond selectively to line segmens of different orientations..
2. Form perception is accomplished by spatial mapping of
the visual scene onto visual cortex.
3. Form perception is accomplished by virtue of Fourier analysis.
Eye-movement control
superior colliculus
sts ls
ce
p SC
visual cortex
FEF
MEF
frontal eye fields
medial eye fields
parietal cortex
V1
V2 LIP
BS
medial eye fields
ls ce
p FEF
MEF
frontal eye fields
BS
Anterior system
Posterior system
visual cortex
parietal cortex
V1
V2 LIP
SC
superior colliculusablated
sts
Summary of the effects of electrical stimulation:
V1 & V2, upper
V1 & V2, lower
V4
LIP
FEF
MEF
FACILITATION INTERFERENCE NO EFFECT FIX INCREASE
Summary of the effects of the GABA agonist
muscimol and the GABA antagonist bicuculline
Target selection Visual discrimination
muscimol bicuculline muscimol bicuculline
V1 V1
FEFFEF
LIPLIP
INTERFERENCE INTERFERENCE
INTERFERENCE FACILITATION
NO EFFECT NO EFFECT
DEFICIT DEFICIT
MILD DEFICIT NO EFFECT
NO EFFECT NO EFFECT
SC FACILITATION INTERFERENCE Hikosaka and Wurtz
Saccade to new location
A
what?
B
C
A
1 1. What are the objects in the scene?
2
A
B
Cwhich?
2. Which object to look at?
A
B
C
5
when?
5. When to initiate the saccade? 4. Where are the objects in space?
where?
A
B
C
4
3. Which object not to look at?
which not?
3
A
B
C
Brain areas involved
V1, V2, V4, IT, LIP, etc.
V1, V2, LIP, FEF, MEF
LIP
V1, V2, FEF, SC
V1, V2, LIP
BS
ras
?
BS
rate code
SC
SN
BG
vector code
P
Midget
1V
w
w
Parasol
LGN M
PARIETAL LOBE TEMPORAL LOBE
2
V4
V2
Midget
?
??
Mixed
Posterior system
Pa ol
MT
FRONTAL LOBE FEF MEF
vector code
place code Anterior system
V Auditory
system
Somatosensory
system
Olfactory
system
Smooth pursuitsystem
Vergence Accessory Vestibularsystem optic system system
Motion perception
Summary of cell types in V1
CXDL
L
LL
L
L
L
L
L
L
L
D
D
D
D
D
D
D
D
D
.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 1.1 1.2 deg
.1 .2 .3 .4 .5
.1 .2 .3 .4 .5 .6 .7 .8
.1 .2 .3 .4 .5 .6 .7 .8 .9
.4.1 .2 .3 .5 .6
.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 deg
.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 deg
deg
deg
1.1
.1 .2 .3 .4 .5 .6 .7 deg
L
D
D
s6
s7
s5s1
s2
s3
s4
DEGREES OF VISUAL ANGLE
DEGREES OF VISUAL ANGLE
Figure by MIT OCW.
BS
Major Pathways of the Accessory Optic System (AOS) Velocity response of AOS neurons = 0.1-1.0 deg/sec Number of AOS RGCs in rabbit = 7K out of 350K
BS
rate code
D
M
L
Terminal Nuclei
Inferior Olive
Cerebellum
climbing fibers
Ant
NOT
Vestibular Nucleus
Semicircular canals
12
3
1
2,3
2,3
Cortex
Prime axes of retinal direction-selective neurons
vestibulo-ocular reflex
Effects of lesions on vision
severe
Summary of lesion deficit magnitudes
VISUAL CAPACITY PLGN MLGN V 4 MT color vision
coarse scotopic vision
brightness perception
contrast sensitivity
motion perception
flicker perception
fine
coarse
stereopsis fine
coarse
pattern perception
shape perception
texture perception
fine coarse
severe
severe
severe
severe
severe
severe
severe
mild
mild
mild
mild
mild
mild
mild
mild
none
none
none none none
none none
none none none
none none
none none
none none
none none
none none
none none none
none none none
none
none
none
none
none
none
moderate moderate
pronounced
mild
pronounced
fine
choice of "lesser" stimuli
not testedvisual learning
not tested
not tested
not tested
none none
none
not tested
severe
severe
severe
pronounced object transformation
INTE
RM
EDIA
TE
BA
SIC
VIS
UA
L FU
NC
TIO
NS
Prosthetics
Figure by MIT OCW.
The size and location of the regions activated in the monkey V1 by the dotted circle presented in the visual field 90 90
3
2
1
4
5
3
2
1
4
5
1 2 3 4
1 234
0
45
315
270
90 90
180
225
135
270
270
0180
135 45
315225
1 2 3 4
1 234
0
45
315
270
90 90
180
225
135
270
270
0180
135 45
315225
The size and location of the regions activated in the monkey V1 by the dots presented in the visual field
590 256 points 55
90
135 4 45 135 4 45
3 3
2 2
1 1
180 0 180 0
225 315 225 315
270 270
2 3 4 270 270 4 3 2 2 3 4 270 270 4 3 2 1 1 1 1
315 225 315 225
0 180 0 180
45 135 45 135
90 90 90 90
Illusions
The Hermann grid illusion
The most widely cited theory purported to explain the illusion:
ON ON
larger response smaller response
Due to antagonistic center/surround organization, the activity of ON-center retinal ganglion cells whose receptive fields fall into the intersections of the grid produces a smaller response than those neurons whose receptive fields fall elsewhere.
Differently oriented vertical and horizontal lines reduce illusion
Retinal ganglion cell receptive field layout at an eccentricity of 5 degrees
At the eccentricity of 5 degrees the 0.5 by 0.5 degree visual angle area outlined impinges on 365 midget cells and 50 parasol cells. Half of these are ON and half OFF cells. The layout of the ON cells is shown in B and C.
5mm 5 deg of visual angle
Retinal midget cells Retinal parasol cells
0.5 deg of visual angle