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The visual and oculomotor systems

Peter H. Schiller, year 2006

Review,

the visual and oculomotor systems

Basic wiring of the visual system

fix

Primates left right


hemifield hemifield

Horopter (Vieth-Muller circle)

temporal nasal

rig
e
er

ht
ph

he
is

m
m

is
he

ph
optic
ft

er
FRONTAL LOBE
le

chiasm

e
TEMPORAL LOBE

PARIETAL LOBE
NOT

p
LGN terminal
m nuclei

MT

V superior
4
colliculi
V
3
V
2
V
1
Retina and LGN

pigment epithelium

rods cones

photo-
receptors

OPL

cone horizontal H ON OFF bipolars ON

IPL
AII
amacrine
ON OFF
ganglion cells

incoming light to CN
CNSS
Visual cortex

Transforms in V1

Orientation

Direction

Spatial Frequency

Binocularity
ON/OFF Convergence
Midget/Parasol Convergence
Three models of columnar organisation in V1

Original Hubel-Wiesel "Ice-C


ube" Model

Cortical
Midget Left Eye Right Eye
Parasol
Sub-cortical
Radical Model

m
1m
Left Eye Right Eye

Swirl Model

Figure by MIT OCW.


Striate Cortex Output Cell

Intracortical

Midget ON Midget ON
Midget OFF Midget OFF
LEFT EYE 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

ON OFF ON/OFF

inhibition inhibition 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

neuronal response profile

ON OFF ON OFF

time

Projections of the midget and parasol systems

Midget
V1
Mixed V2

Parasol

LGN P
M ? MT V4

Midget
Parasol

PARIETAL LOBE TEMPORAL LOBE


Spatial Frequency
H

Processing Capacity L

L H

Midget System
Parasol System

Temporal Frequency
H

L H

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

white

white

Yellow

saturation

Green

Red

Blue hue

black

Response to Different Wavelength Compositions in LGN


Blue ON cell Yellow ON cell
90 90

135 45 135 45
Spikes per Second

180
10 20 30 40 50 60
0 180 20 40 60 80 100
0

225 315 225 315

270 270

Green OFF cell Red ON cell


90 90

135 45 135 45

180
0 180 10 20 30 40 50
0
10 20 30 40

225 315 225 315


maintained discharge rate
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

1 a

b
eye
object motion
a movement

The eye tracks the circle, which


therefore remains stationary on the fovea

Objects nearer than the one tracked move 2 b


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

medial eye fields


frontal eye fields
parietal cortex

visual cortex

MEF

p LIP
FEF SC V2

ce sts V1
ls

BS
medial eye fields

frontal eye fields parietal cortex

superior colliculus
ablated

visual cortex

MEF

p LIP
Anterior system FEF SC V2

ce sts V1
ls

BS
Posterior system
Summary of the effects of electrical stimulation:

FACILITATION INTERFERENCE FIX INCREASE NO EFFECT

V1 & V2, upper

V1 & V2, lower

V4

LIP

FEF

MEF

Summary of the effects of the GABA agonist

muscimol and the GABA antagonist bicuculline

Target selection Visual discrimination

muscimol bicuculline muscimol bicuculline

V1 INTERFERENCE INTERFERENCE V1 DEFICIT DEFICIT

FEF INTERFERENCE FACILITATION FEF MILD DEFICIT NO EFFECT

LIP NO EFFECT NO EFFECT LIP NO EFFECT NO EFFECT

SC INTERFERENCE FACILITATION Hikosaka and Wurtz


Saccade to new location

A
1. What are the objects in the scene?
1 2. Which object to look at?
B 3. Which object not to look at?
V1, V2, V4, 4. Where are the objects in space?
A
IT, LIP, etc. 5. When to initiate the saccade?
what?
C 2
B
V1, V2, LIP,
A
FEF, MEF
which?
C 3
Br B
a in V1, V2, LIP A
a re
a which not?
C 4
si
nv B
olv V1, V2, A
ed FEF, SC
where?
C
5
B

A
LIP
C
when?
Midget
V1
V2 Auditory

Mixed
? system

Parasol
?

LGN
P Somatosensory

M MT V4 system

Posterior
Midget system
Parasol

PARIETAL LOBE TEMPORAL LOBE Olfactory

system

w ?
SC BG

rate code
FRONTAL LOBE
SN Smooth pursuit

BS vector code FEF MEF


BS
system

vector code
place code
Anterior
system

Vergence Accessory Vestibular

system optic system system

Motion perception

Summary of cell types in V1

s1 s5
D
D
L
.1 .2 .3 .4 .5 .1 .3 .4 .5 .6 .7 deg
.2
DEGREES OF VISUAL ANGLE
L
D

s2
L D
s6
.1 .2 .3 .4 .5 .6 .7 .8
D L
DEGREES OF VISUAL ANGLE

.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 1.1 deg

s3 L
D
L

.1 .2 .3 .4 .5 .6 .7 .8 .9 deg

s7 D
D
L
L
.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 deg

L
D
s4 L
L D

.1 .2 .3 .4 .5 .6 deg

CX
D
D L

.1 .2 .3 .4 .5 .6 .7 .8 .9 1.0 1.1 1.2 deg

Figure by MIT OCW.


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

Cortex
2 1 Cerebellum
Ant
3

climbing fibers
Prime axes of retinal NOT
direction-selective neurons

Inferior Olive
D Semicircular
1 canals

M
2,3
Vestibular
L Nucleus
rate code 2,3

BS Terminal
BS Nuclei

vestibulo-ocular
reflex
Effects of lesions on vision

Summary of lesion deficit magnitudes


VISUAL CAPACITY PLGN MLGN V4 MT
color vision severe none mild none

texture perception severe none mild none

pattern perception fine severe none mild none

shape perception fine severe none mild none


BASIC VISUAL FUNCTIONS

coarse mild none none none

brightness perception none none none none

coarse scotopic vision none none none none

contrast sensitivity fine severe none mild mild

coarse mild none none mild

stereopsis fine severe none none none

coarse pronounced none none none

motion perception none moderate none moderate

flicker perception none severe none pronounced


INTERMEDIATE

choice of "lesser" stimuli severe


severe none severe none

visual learning not tested not tested severe none

object transformation not tested not tested pronounced not tested


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

5 5

135 4 45 135 4 45
3 3
2 2

1 1

180 0 180 0

225 315 225 315

270 270

3 4 4 3 3 4 4 3
2 270 270 2 2 270 270 2
1 1 1 1

315 225 315 225

0 180 0 180

45 135 45 135

90 90 90 90
The size and location of the regions activated in the monkey V1 by the dots presented in the visual field
90 256 points 90
5 55

135 4 45 135 4 45
3 3
2 2

1 1

180 0 180 0

225 315 225 315

270 270

3 4 4 3 3 4 4 3
2 270 270 2 2 270 270 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