The visual and oculomotor systems

Peter H. Schiller, year 2006

Review,
the visual and oculomotor systems

Basic wiring of the visual system

Primates
Horopter (Vieth-Muller circle)

fix

left hemifield

right hemifield

temporal

nasal

m he ht rig

le ft he m

is ph er

e

er ph is

FRONTAL LOBE TEMPORAL LOBE PARIETAL LOBE

optic chiasm

e

NOT
p
terminal nuclei

LGN MT V

m

4

superior colliculi
V
3

V

2

V

1

Retina and LGN

pigment epithelium

rods

cones photoreceptors

OPL
cone horizontal

H

ON

OFF

bipolars

ON

IPL
AII ON OFF
amacrine ganglion cells

incoming light

to CNS CNS

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 Parasol

Left Eye

Right Eye

Radical Model

Sub-cortical

Left Eye

Right Eye

Swirl Model

Figure by MIT OCW.

1m

m

Striate Cortex Output Cell

Intracortical
Midget ON Midget OFF Midget ON Midget OFF

LEFT EYE INPUT

Parasol ON Parasol OFF

Parasol ON Parasol OFF

RIGHT EYE INPUT

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
V1
Midget Mixed Parasol

V2

w

LGN

P
M

?
Parasol

MT

V4

Midget

PARIETAL LOBE

TEMPORAL LOBE

Spatial Frequency
H

L

Processing Capacity

L

H

Midget System Parasol System

Temporal Frequency
H

L 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

saturation

Yellow

Green Red

Blue

hue

black

Response to Different Wavelength Compositions in LGN Blue ON cell Yellow ON cell
90 135 45 135
Spikes per Second

90
45

180

10 20 30 40 50 60

0

180

20 40 60 80 100

0

225 270

315

225 270

315

Green OFF cell
90 135 45 135

Red ON cell
90 45

180

10

20

30

40

0

180

10 20 30 40 50

0

225 270

315

225 270

315

maintained discharge rate

Depth perception

Cues used for coding depth in the brain
Oculomotor cues
accommodation vergence

Visual cues
Binocular
stereopsis

Monocular
motion parallax shading interposition size perspective

stereo camera

MOTION PARALLAX, the eye tracks

1

a

2 b a

eye movement

object motion

1

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.
2 b

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 FEF ce LIP sts ls BS

SC

V2 V1

medial eye fields
frontal eye fields parietal cortex

superior colliculus ablated

visual cortex
MEF

Anterior system

p

FEF ce

SC sts

LIP

V2 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
muscimol bicuculline
INTERFERENCE

Visual discrimination
muscimol bicuculline
DEFICIT

V1 FEF

INTERFERENCE

V1 FEF

DEFICIT

INTERFERENCE

FACILITATION

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
B

V1, V2, V4, IT, LIP, etc.
what?

A C B

1. 2. 3. 4.

What are the objects in the scene? Which object to look at? Which object not to look at? Where are the objects in space?

2
A

5. When to initiate the saccade?

V1, V2, LIP, FEF, MEF

Br a

which?

C B

3
A

in a

V1, V2, LIP

re a

si nv olv ed

which not?

C B

4
A
C B

V1, V2, FEF, SC
where?

5
A C

LIP
when?

V1

Midget Mixed Parasol

V2
?

Auditory

?

system

w

LGN

P M

Somatosensory

Midget Parasol

MT Posterior system
PARIETAL LOBE

V4

system

?

Olfactory
system

TEMPORAL LOBE

w

SC

?

BG

rate code

BS

SN
vector code vector code

BS

FRONTAL LOBE FEF MEF

Smooth pursuit
system

place code

Anterior system

Vergence system

Accessory optic system

Vestibular
system

Motion perception

Summary of cell types in V1

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

D

L D

s2
L D

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

D

L

.1

.2

.3

.4

.5

.6

.7

.8

.9

1.0

1.1 deg

s3
.1 .2

L
.3 .4 .5 .6 .7 .8 .9

L

D

deg

D

s7
L
.1 .2 .3 .4 .5

D L
.6 .7 .8 .9 1.0 deg

s4
L
.1 .2 .3

L

D

L

D
.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

2 3

Cortex
1 Ant

Cerebellum

climbing fibers Prime axes of retinal direction-selective neurons
NOT

Inferior Olive
D
1

Semicircular canals

M
2,3

L

rate code

Vestibular Nucleus

2,3

BS

BS

Terminal Nuclei

vestibulo-ocular reflex

Effects of lesions on vision

Summary of lesion deficit magnitudes
VISUAL CAPACITY
color vision texture perception pattern perception
BASIC VISUAL FUNCTIONS

PLGN
severe severe

MLGN
none none none none none none none none none none none

V4
mild mild mild mild
none none none

MT
none none none none none none none

fine fine coarse

severe severe mild
none none

shape perception

brightness perception coarse scotopic vision contrast sensitivity fine coarse stereopsis fine coarse motion perception flicker perception

severe mild severe pronounced
none none

mild
none none none none none

mild mild
none none

moderate severe
none
not tested not tested

moderate pronounced
none none
not tested

INTERMEDIATE

choice of "lesser" stimuli visual learning object transformation

severe severe
not tested not tested

severe severe pronounced

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
5

90
4 3 2 1

5

90

4 3 2 1

135

45

135

45

180

0

180

0

225

315

225

315

270
2 1 3 4
270 270

270
4 3 2 2 1 3 4
270 270

4

3

1

2

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
5

256 points 45 135

5 5 4 3

90 45

135

4 3 2 1

2 1

180

0

180

0

225

315

225

315

270
2 1 3 4
270 270

270
4 3 2 2 1 3 4
270 270

4

3

1

2

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

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