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Physiology of Vision: a swift overview
Parietal
visual
cortex
Dorsal
Stream Striate
cortex
(V1)
LGN
Thalamus
Extrastriate
cortex
Eye Optic
Ventral
nerve Temporal Stream
visual
cortex
Some figures from Steve Palmer
Pixels to Percepts
A. Efros, CMU, Spring 2011
Understanding the Brain
Anatomy versus Physiology
Anatomy: The biological study of the
physical structure of organisms.
Physiology: The biological study of the
functional structure of organisms.
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
The Brain is tricky business
• Aristotle thought it’s for cooling the blood
• Localized or distributed?
Phrenology
How bad Machine Learning got started…
Localized or Distributed?
Evidence from patients with partial brain
damage
• Lots of useful data from soldiers in the Russo-Japanese war
But also evidence for distributed nature of
processing
• E.g. [Lashley] showed “graceful degradation” of memory
performance in rats
The Visual System
Both eye and brain are required for
functional vision
Two kinds of blindness:
Normal blindness (eye dysfunction)
Cortical blindness (brain dysfunction)
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
The Visual System
Eyes register optical information
Pathways to occipital cortex
Two pathways from V1
“What” pathway to temporal cortex
“Where” pathway to parietal cortex
Convergence on frontal cortex
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Pathways to the Brain
Anatomy of Pathway to Visual Cortex
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Pathways to the Brain
The Lateral Geniculate Nucleus (LGN)
Waystation in Thalamus
Projections from both eyes
Six layers
Projects to cortical area V1
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Visual Cortex
Map of Visual Areas in Cortex
“Unfolded” view of
visual areas in the
macaque cortex
(sizes not to scale).
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Visual Cortex
What/Where Pathways
Dorsal Pathway
("Where" System)
Parietal
Frontal
Occipital Primary
Visual
Cortex
Temporal
A
Ventral Pathway
("What" System)
Evidence from lesions
of monkey cortex
Dorsal Lesion
Ventral Lesion
B
Object Discrimination
C
Landmark Discrimination
© Stephen E. Palmer, 2002
Visual Cortex
What/Where Pathways
Evidence from Neuropsychology
Visual agnosia:
Inability to identify objects and/or people
Caused by damage to inferior (lower) temporal lobe
Disruption of the “what” pathway
Visual neglect:
Inability to see objects in the left visual field
Caused by damage to right parietal lobe
Disruption of the “where” pathway
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Visual Cortex
Feature-based Pathways Hypothesis
Visual Features
Color
Shape
Depth
Motion
Color
Depth
Form
Color
Featural Pathways
Separate neural pathways
in which different features are
processed.
Motion
V2
Form
Depth
Motion
MT
V1
Color
Form
Depth
Motion
LGN
Color
Form
Depth
Motion
RETINA
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
The Gross Summary
The visual system is composed of
many interactive functional parts:
Eye (optics of image formation)
Retina (light transduction)
LGN (waystation?)
Area V1 (hypercolumns)
Higher cortical areas (features)
Cortical pathways (what/where)
© Stephen E. Palmer, 2002
© Stephen E. Palmer, 2002
Image Formation
Digital Camera
Film
The Eye
Monocular Visual Field: 160 deg (w) X 135 deg (h)
Binocular Visual Field: 200 deg (w) X 135 deg (h)
The Eye is a camera
The human eye is a camera!
• Iris - colored annulus with radial muscles
• Pupil - the hole (aperture) whose size is controlled by the iris
• What’s the “film”?
– photoreceptor cells (rods and cones) in the retina
The Retina
Cross-section of eye
Cross section of retina
Pigmented
epithelium
Ganglion axons
Ganglion cell layer
Bipolar cell layer
Receptor layer
Retina up-close
Light
Two types of light-sensitive receptors
Cones
cone-shaped
less sensitive
operate in high light
color vision
Rods
rod-shaped
highly sensitive
operate at night
gray-scale vision
© Stephen E. Palmer, 2002
Rod / Cone sensitivity
The famous sock-matching problem…
Distribution of Rods and Cones
# Receptors/mm2
.
Fovea
150,000
Rods
Blind
Spot
Rods
100,000
50,000
0
Cones
Cones
80 60 40 20 0
20 40 60 80
Visual Angle (degrees from fovea)
Night Sky: why are there more stars off-center?
© Stephen E. Palmer, 2002
Electromagnetic Spectrum
Human Luminance Sensitivity Function
http://www.yorku.ca/eye/photopik.htm
Visible Light
Why do we see light of these wavelengths?
…because that’s where the
Sun radiates EM energy
© Stephen E. Palmer, 2002
The Physics of Light
Any patch of light can be completely described
physically by its spectrum: the number of photons
(per time unit) at each wavelength 400 - 700 nm.
# Photons
(per ms.)
400 500
600
700
Wavelength (nm.)
© Stephen E. Palmer, 2002
The Physics of Light
Some examples of the spectra of light sources
.
B. Gallium Phosphide Crystal
# Photons
# Photons
A. Ruby Laser
400 500
600
700
400 500
Wavelength (nm.)
700
Wavelength (nm.)
D. Normal Daylight
# Photons
C. Tungsten Lightbulb
# Photons
600
400 500
600
700
400 500
600
700
© Stephen E. Palmer, 2002
The Physics of Light
% Photons Reflected
Some examples of the reflectance spectra of surfaces
Red
400
Yellow
700 400
Blue
700 400
Wavelength (nm)
Purple
700 400
700
© Stephen E. Palmer, 2002
The Psychophysical Correspondence
There is no simple functional description for the perceived
color of all lights under all viewing conditions, but …...
A helpful constraint:
Consider only physical spectra with normal distributions
mean
area
# Photons
400
500
variance
600
700
Wavelength (nm.)
© Stephen E. Palmer, 2002
The Psychophysical Correspondence
# Photons
Mean
blue
Hue
green yellow
Wavelength
© Stephen E. Palmer, 2002
The Psychophysical Correspondence
# Photons
Variance
Saturation
hi. high
med. medium
low
low
Wavelength
© Stephen E. Palmer, 2002
The Psychophysical Correspondence
Area
Brightness
# Photons
B. Area
Lightness
bright
dark
Wavelength
© Stephen E. Palmer, 2002
Physiology of Color Vision
Three kinds of cones:
440
RELATIVE ABSORBANCE (%)
.
530 560 nm.
100
S
M
L
50
400
450
500
550
600 650
WAVELENGTH (nm.)
• Why are M and L cones so close?
© Stephen E. Palmer, 2002
Retinal Processing
© Stephen E. Palmer, 2002
Single Cell Recording
Microelectrode
Amplifier
Electrical response
(action potentials)
mV
Time
© Stephen E. Palmer, 2002
Single Cell Recording
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in ganglion cells:
On-center Off-surround
Response
Time
Stimulus condition
Electrical response
© Stephen E. Palmer, 2002
Retinal Receptive Fields
RF of On-center Off-surround cells
Neural Response
Response Profile
Receptive Field
Center
Firing
Rate
on-center
Surround
off-surround
On
Off
Horizontal Position
© Stephen E. Palmer, 2002
Retinal Receptive Fields
RF of Off-center On-surround cells
Neural Response
Surround
Center
Receptive Field
Response Profile
Firing
Rate
on-surround
Surround
Center
off-center
On
Off
Horizontal Position
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Retinal Receptive Fields
Receptive field structure in bipolar cells
Light
© Stephen E. Palmer, 2002
Retinal Receptive Fields
Receptive field structure in bipolar cells
LIGHT
Receptors
Horizontal
Cells
Direct Path
Indirect Path
Direct excitatory
component (D)
Indirect
inhibitory
component (I)
D+I
Bipolar Cell
A. WIRING DIAGRAM
B. RECEPTIVE FIELD PROFILES
© Stephen E. Palmer, 2002
Visual Cortex
Cortical Area V1
Parietal
visual
cortex
aka:
Primary visual cortex
Striate cortex
Brodman’s area 17
Dorsal
Stream Striate
cortex
(V1)
LGN
Thalamus
Extrastriate
cortex
Eye Optic
Ventral
nerve Temporal Stream
visual
cortex
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Single-cell recording from visual cortex
David Hubel & Thorston Wiesel
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Single-cell recording from visual cortex
Time
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Three classes of cells in V1
Simple cells
Complex cells
Hypercomplex cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Simple Cells: “Line Detectors”
B. Dark Line Detector
Firing
Rate
Horizontal Position
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Simple Cells: “Edge Detectors”
C. Dark-to-light Edge Detector
Firing
Rate
D. Light-to-dark Edge Detector
Firing
Rate
Horizontal Position
Horizontal Position
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Constructing a line detector
Retina
Receptive Fields
LGN
CenterSurround
Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Complex Cells
STIMULUS
NEURAL RESPONSE
00o
Time
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Complex Cells
STIMULUS
NEURAL RESPONSE
o
60
0
Time
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Complex Cells
STIMULUS
NEURAL RESPONSE
o
90
0
Time
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Complex Cells
STIMULUS
NEURAL RESPONSE
o
120
0
Time
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Constructing a Complex Cell
Retina
Receptive Fields
Cortical Area V1
Simple Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Hypercomplex Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Hypercomplex Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Hypercomplex Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Hypercomplex Cells
“End-stopped” Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
“End-stopped” Simple Cells
© Stephen E. Palmer, 2002
Cortical Receptive Fields
Constructing a Hypercomplex Cell
RETINA
Receptive Fields
CORTICAL AREA V1
Complex Cell
End-stopped Cell
© Stephen E. Palmer, 2002
Mapping from Retina to V1
Why edges?
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