Senses I: Vision

In this lesson, we will discuss...

1. Eyes and Light

2. Color Perception
3. Lateral Inhibition
4. Retina

5. Primary Visual Cortex

6. “What” and “Where” Pathways

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1. Eyes and Light

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Sensation & Perception
• the processes of transforming
physical energy into
psychological experiences/feelings
• Our systems have evolved to process
biologically useful information.
• Some info lies beyond our sensory
or perceptual limits (e.g., infrared lights, ultrasound)
• Each of our senses has specialized
receptors that are sensitive to a
particular kind of energy,
e.g., vision: light, audition: sound
• Transduction: the process of converting physical energy (e.g.,
light) into neural signals
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 The Physics of Lights
• Lights are a form of electromagnetic (EM) waves,
which are oscillations of electric and magnetic energy.
• There are many types of EM waves, characterized by their
differences in wavelengths.
• Visible lights: from 400nm (violet) to 700nm (red)

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The Eye
• Light enters the eye through
the pupil.
• The iris controls the size of
the pupil.
• Afterwards, the lens focuses
lights onto the retina.
• Ciliary muscles control the
thickness of the lens for us to
focus at different distances.
• Vitreous body (a gel-like
substance) fills the eyeball.

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Retina: Overview
• the first stage of visual processing
• contains photoreceptors and other retinal neurons
• Images projected on the retina are inverted both vertically
and horizontally.
• Light from the left side of the world strikes the right side of
the retina, and vice versa.

left right

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Retina: Types of Cells
• Photoreceptors (rods & cones):
transduce lights
• Bipolar cells: gather inputs from
photoreceptors and send output to
ganglion cells
• Ganglion cells: gather inputs from
bipolar cells and takes signals out of
the eye to the brain
• Horizontal cells: send inhibitory
signals laterally to multiple
neighboring cells (see later slides)
• Amacrine cells: inhibitory neurons
that facilitate various processes
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Retina: Structure
• It has an inverted (or backward)
structure: Light passes through all
the neurons before reaching the
photoreceptors.
• Fovea: The central part of the
retina, which contains almost no
ganglion axons or blood vessels
Periphery: Non-central regions of
the retina
• Blind spot:
– where the optic nerve leaves
the eye
– does not contain
photoreceptors (hence “blind”)
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 Photoreceptors: Rods and Cones
• transduce light energy to neural signals
• send signals by varying glutamate release
Rods Cones
In periphery Abundant Very scarce
In fovea Absent Densely packed
Sensitivity in dim light High Low
Number (each eye) ~120 million ~6 million
Output convergence many rods -> 1 ganglion cell 1 cone -> 1 ganglion cell
Wavelength selectivity Only one type Three types of cones

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Wavelength Selectivity
Cones: 3 types are named based on their “preferred” wavelengths:
Short (S)-, Medium (M)-, and Long (L)-wavelength cones
Rods: all have the same wavelength selectivity (preferring mid-range).
S cones Rods M cones L cones

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2. Color Perception

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From Physics to Psychology, e.g., color
• Wavelengths are physical. Colors are psychological.
• This is a variant of the mind-body problem in the specific
domain of color perception!
• Trichromatic vs Opponent-process theories

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The Trichromatic Theory
The theory states that our perception of color is determined by
the ratio of the responses from the 3 cones (S:M:L),
e.g., Blue = 90:30:20, Cyan = 30:50:30, Orange = 0.1:15:70

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The Opponent-process Theory
• The brain has two independent opponent color continuums:
Red-Green and Yellow-Blue. Perceived color is determined based
on the strength of activation on each continuum.
e.g., Suppose {Yellow=-ve | Blue=+ve} {Red=-ve | Green=+ve}.
Then, orange = {-0.75} {-0.75}, Green = {0} {+1}, Violet = {+1} {-1}
• supported by the processing mechanisms in some retinal neurons
• This theory can explain perception of negative afterimages.
+ve

–ve +ve

–ve 15
Color Constancy
• Perceptual constancy: the ability to recognize the same
stimulus across different conditions (e.g., brightness, color,
face recognition, etc.)
• Color constancy: allow us to recognize colors under different
lighting conditions
The (in)famous dress
• Explanation: one stimulus, two possible interpretations
• No evidence
to support the idea
that you’re
smarter /
more sensitive /
more outgoing /
more romantic / etc.
if your brain
interprets it one way
but not the other!

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3. Lateral Inhibition

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 [for your interest ☺]

Lateral Inhibition
• The excitation of one neuron inhibits the activities of laterally
neighboring neurons.
• to highlight contrasts and emphasize the borders of objects
(see the illusions below)
• caused by inhibition from the horizontal cells

Mach bands

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 [for your interest ☺]

another Illusion from lateral inhibition

• lighter = more light, stronger stimulus; darker = less light, weaker stimulus
• Q: Why do I see the illusory dark spots at the intersections?
• A: This is the result of lateral inhibition: The lighter lines activates some
neurons, and those neurons send laterally inhibitory signals to neighboring
neurons. Neurons that pick up signals at the intersections are, in fact,
neighbors of all those neurons that pick up signals from the lines.
Therefore, they receive a lot of inhibition → you see “dark spots”
Hermann grid a variant by Lingelbach (1994)

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4. Retina

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Bipolar cells
• receive inputs from either rods or
cones, but not both
• send outputs to ganglion cells
using graded potentials
• In fovea, a bipolar cell receives
input from only one cone
(hence the low convergence)
• In periphery, a bipolar cell
receives input from many rods
(hence the high convergence)

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Foveal vs Peripheral vision
Foveal Peripheral
Receptors cones many more rods than cones
Convergence of input one-to-one many-to-one
Sensitivity in dim light low high
Spatial resolution high low
Color sensitivity high low

"Human photoreceptor distribution" by Cmglee - Own work. Licensed under

CC BY-SA 3.0 via Commons -
https://commons.wikimedia.org/wiki/File:Human_photoreceptor_distributio
n.svg#/media/File:Human_photoreceptor_distribution.svg 23
Receptive Field
• the area in the visual world to which a
neuron responds (could be excitatory or
inhibitory)
• E.g., photoreceptors have the smallest
receptive field in the visual system
(i.e., a point)
• If a ganglion cell receives inputs from
many photoreceptors, it should have a
large receptive field; and vice versa.

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Retinal Ganglion Cells (RGC)

Types of RGCs Midget Parasol Bistratified

a.k.a. P cells M cells K cells
in Kalat as… “parvocellular” “magnocellular” “koniocellular”
abundance ~80% ~10% 8-10%
receive inputs from mostly cones mostly rods possibly blue cones?
receptive field size small large Mostly small, but variable

conduction speed slow fast moderate

• Some color opponent processes are found in P cells RGCs.
• some remaining RGCs are even photosensitive!
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Lateral Geniculate Nuclei (LGN)
• the visual relay station located in the thalamus
• Signals from the retina are sent to LGN before reaching the cerebral cortex.
• Optic chiasm: a point at which the left and right optic nerves cross before
reaching LGN
• Signals from the left and right
visual fields are sent to the
contralateral LGN:
Right visual field -> Left LGN
Left visual field -> Right LGN
• Parvocellular (“small cell”) layer:
from midget RGCs (P cells)
• Magnocellular (“big cell”) layer:
from parasol RGCs (M cells)
• Koniocellular layer:
from bistratified RGCs (K cells)

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 [for your interest
☺]
Center-Surround Receptive Fields
• Receptive fields of RGCs and LGN cells tend to have a
center-surround configuration.
• 2 types (“On” means excitatory, “Off” means inhibitory):
– On-center, Off-surround (center=excite, surround=inhibit)
– Off-center, On-surround (center=inhibit, surround=excite)
• ideal for detecting dots or small blobs in visual scenes

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5. Primary Visual Cortex

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Cortical Visual Processing
• can be divided into multiple stages
(or levels) of processing, e.g.,
– Precortical: RGC, LGN
– Early stages: V1-V2
– Later stages: V2-V4,
middle temporal (MT),
interior temporal (IT), etc….
• Visual neurons can be thought of as
feature detectors: neurons respond to
the presence of
particular visual
feature/ stimuli

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Primary Visual Cortex (V1)
• receives signals from the LGN
• a.k.a. striate cortex (anatomical term)
• first stage of visual processing
in the cerebral cortex
• contains neurons that are
highly specialized in analyzing small images features
(see later slides)
• Damage to V1 could result in blindsight.
(a.k.a cortical blindness):
– the ability to respond to or act on visual stimuli without
the conscious experience of “seeing”

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Organization of neurons in V1
V1 neurons
• are grouped together in columns: Neurons within a given column are
selective to similar information (e.g., same orientation).
• form a retinotopic map: Specific neurons in V1 can be mapped to specific
locations on the retina (and then to the visual world).

columnar
organization

retinotopic
mapping

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Selectivity of V1 neurons
• Neurons in V1 respond selectively to certain
primitive image features, e.g.,
– orientations
– spatial frequencies (SFs)
– location
• Preferred stimuli → High firing rate
• Hubel and Wiesel’s experiment (YouTube)
(turn sound ON before watching!)

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Hubel and Wiesel’s experiment

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Orientation-selective Neurons in V1
Simple cells Complex cells
Found in… V1 V1 & V2
Receptive field size smaller larger
Distinctive feature of Fixed excitatory or No fixed excitatory or
receptive field inhibitory zones inhibitory zones
• Both simple and complex cells
are orientation-selective:
each simple/complex cell
has a specific
“preferred orientation”,
to which it produces
the highest firing rate.

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After V1, what’s next?
• The secondary visual cortex (V2)
receives information from area V1,
processes information further (e.g.,
comparisons), and sends it to other
areas (including feedback to V1).
• Afterwards, it is believed that
there are two visual pathways
(Goodale and Milner, 1992):
– The “what” pathway
(ventral stream)
– The “where” pathway
(dorsal stream)

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6. “What” and “Where” Pathways

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“What” and “Where” Pathways
Ventral stream (the “what” pathway)
• takes inputs from the parvocellular cells
• goes ventral (down) through the temporal cortex
• specialized in identifying and recognizing objects

Dorsal stream (the “where” pathway)

• takes inputs from the magnocellular cells
• goes dorsal (up) towards the parietal cortex
• specialized in processing
motion and spatial information,
and vision-guide motor responses

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Ventral (“what”) Stream: example areas
V4
• receives inputs from V2
• selectively responsive to simple shapes
• a color-processing center in the cortex

Inferior temporal cortex (IT)

• a high-level processing center in the
ventral stream
• neurons have large receptive field
• selectively responsive to complex
shapes, identifiable objects, or faces
• insensitive to changes in small features
or size/orientation

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Dorsal (“where”) Stream: example areas
Middle Temporal (MT) area (a.k.a V5)
• a motion-processing center
• neurons have large receptive fields
• neurons selectively respond to
particular motion direction or speed,
but insensitive to color

Medial Superior Temporal cortex (MST)

• like MT, but more complex
• neurons have even larger receptive
fields (larger than MT)
• selectively respond to complex motion
patterns (e.g., rotation, expansion, etc.)
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Visual Processing is Hierarchical
• Low- to mid-level processing
– retina (points) → LGN (dots/blobs) →
V1 (lines) → V2 (combining V1 outputs)
→ V4 (color comparison) → …
• High-level processing
– IT (shape/object/face),
MT (motion direction/speed),
MST (complex motion patterns), etc.
• Take-home message:
The higher the processing level…
– the larger the receptive fields
– the more complex the preferred stimuli

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Caveat on the “Two-stream” Theory
• The two streams are not that separate/independent;
they communicate.
• While many experimental results support the theory,
many other results show that there are significant
overlap and interactions between the two streams.
• Demo on biological motion

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 [for your interest ☺]

Visual Impairments
• Damage to specific areas could lead to
specific visual impairments.
• Visual agnosia: the inability to recognize
objects despite satisfactory vision (the
object/shape area in IT)
• Prosopagnosia:
the inability to recognize faces (the
fusiform face area (FFA) in IT)
• Motion blindness: the inability to
perceive motion (MT/MST)

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End of this lesson

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