The Retina

The Retina

• The goal of the processes of the iris, pupil, and lens is to focus an
image on the retina, the eye’s photosensitive surface (Schwartz &
Krantz, 2016).
Anatomy of the Retina

• The retina is the location in the eye where transduction takes

place
• Two types of photoreceptors are in the retina, rods and
cones.
• Rods prevail in the periphery whereas cones are clustered
in and near the fovea, wc is a small dimple on the center
of the retina.
• There are about 120 million rods and 7 million cones in
each eye.
Fovea

• It is the location on the retina with the highest density of

cones, and it has no rods.
• Other retinal cells are not present at the retinal surface,
allowing light to reach the surface of the fovea with
minimal light scatter.
• Both of these features evolved to enhance visual acuity, or
clarity of vision.
optic disc
• The part of the retina where the optic nerve leaves the eye
and heads to the brain.
• No receptors are at the optic disc, so this location is called
the blind spot for that eye.
• The optic nerve carries the neural signal into the brain
Retinal Physiology

• Rods and cones convert light into a neural signal by using

chemicals called photopigments, the molecules that absorb light
which releases an electric potential by altering the voltage in the
cell.
• When a photopigment absorbs a photon of light, it changes
shape, initiating a series of biochemical processes leading to a
neural signal.
• Photopigments are composed of two bound molecules, an opsin
in rods and (chromopsin in cones) and retinal (a form of vitamin
A).
• In rods, the photopigment is called rhodopsin.
• When a photon of light is absorbed by the
photopigment, the rhodopsin straightens out and the
photopigment breaks apart, causing the receptor to
have a hyperpolarization
• Hyperpolarization causes release of less of the
neurotransmitter.
• Thus, light is peculiarly inhibitory but only in the sense
that the neurotransmitter is also inhibitory.
 • This ultimately excites the visual system.
• In cones, there are three classes of photopigments, each
of which is maximally sensitive to a particular frequency of
light.
• The interaction of these three photopigments and
subsequent perceptual processing give us complex color
vision.
The Duplex Theory of Vision

• The duplex theory of vision suggests that there are two

distinct ways in which our eyes work.
• The first system is called the photopic system, which is
associated with cones and daytime vision.
• The second system is called the scotopic system, which is
associated with rods and nighttime vision.
• There is also a range of intermediate light intensity in which
both systems work, which results in mesopic vision.
Spatial Summation and Acuity

• Acuity refers to the ability to see or resolve fine details.

• The relative acuity of photopic and scotopic vision is different
because of how well each system carries out spatial
summation.
• Spatial summation refers to the ability to pool light across
different regions of space.
• Rods are more sensitive to dim light than cones, partly
because many rods connect to one retinal ganglion cell,
allowing the scotopic system to pool responses across
different rods in order to maximize sensitivity to light. Rods
cannot sense color or focus well.
• A drawback of pooling across rods, however, is that the
scotopic system loses some ability to converge light.
• Cones are for bright stns, and they focus well.
• Each cone connects to only one retinal ganglion cell,
providing photopic vision with greater visual acuity.
Dark and Light Adaptation

• Dark adaptation is the process whereby the visual system’s sensitivity

to low light levels is increased. It takes nearly 30 minutes to complete.
• Light adaptation the process whereby the visual system’s sensitivity
to low light levels is decreased so that it can operate in higher light
levels. It takes 5 minutes or less to complete (Schwartz & Krantz,
2016).
• The difference in time to completion is due partly to light adaptation
being driven actively by light entering the eye whereas dark
adaptation is a passive response.
Retinal Ganglion Cells and Receptive
Fields
• Each photoreceptor is connected to one or more retinal
ganglion cells, which convey the signal into the optic nerve
toward the brain.
• There are fewer ganglion cells than there are
photoreceptors; therefore, most ganglion cells receive inputs
from many different photoreceptors through a process called
convergence.
• The receptive field is the area in the visual field that a
particular vision neuron responds to.
• Receptive fields of cones are quite small because one or a
few cones maps onto each retinal ganglion cell.
• Receptive fields of rods are much larger because many
rods (sometimes hundreds) map onto a single ganglion
cell.
• Edge detection involves determining the location at which
one object ends and another begins.
• The earliest stage of edge detection occurs in the retinal
ganglion cells and is known as center-surround receptive
fields.
• In on-center receptive fields, the cell’s center produces
activation, whereas the surround produces inhibition.
• In off-center receptive fields, the cell’s center produces
inhibition, whereas the surround produces activation.
• In either case, if you present light that covers the entire
receptive field of the cell, the response will be small,
because the excitation and inhibition will cancel each
other out.
• The physiological mechanism that creates center-surround
receptive fields is quite complex, involving horizontal cells,
bipolar cells, rods, cones, and retinal ganglion cells.
• Horizontal cells are specialized retinal cells that contact both
photoreceptor and bipolar cells (Wolfe, 2015).
• Bipolar cells are retinal cell that synapses with either rods or
cones and with horizontal cells, and then passes the signals
on to ganglion cells.
• Lateral inhibition facilitates edge detection by reducing a
response of the eye to light stimulating one receptor by
stimulation of nearby receptors, caused by inhibitory signals
in horizontal cells.
• Lateral inhibition enables the signals that reach retinal
ganglion cells to be based on differences in activation
between nearby photoreceptors.
Refractive Errors and Diseases of
the Eye
Myopia (Nearsightedness)
• Myopia is a common form of mild visual impairment in which
people can focus well on near objects but faraway objects
appear blurry.
• The eye tends to be too long from front to back for the lens,
projecting distant objects in front of the retina instead of onto
it, making them appear blurry.
• To correct the problem, the lens must be weakened by using a
diverging or negative artificial lens such as those in eyeglasses.
 Hyperopia (Farsightedness) and Presbyopia (Old-Sightedness)
• Hyperopia is a common form of mild visual impairment in
which people can focus well on far objects but near objects
appear blurry.
• The eye tends to be too short for the lens, projecting near
objects behind the retina instead of onto it, making them
blurry.
• To correct the problem, the lens must be strengthened.
• Presbyopia is a condition associated with older eyes.
• As we age, the lens hardens, and the ciliary muscles lose
power, making it difficult for older eyes to accommodate to
nearby objects.
• As with hyperopia, the lens projects near objects behind the
retina.
Astigmatism

• Astigmatism is a condition that develops from an irregular

shape of the cornea or lens, making it impossible for the lens
to accommodate a fully focused image.
• The cornea may be unsymmetrical, bending light more
strongly in one direction.
• Thus, an object in a particular location and orientation will
not be in focus.
• Some orientations are blurry but others are still relatively
clear.
Cataracts

• Cataracts result from the clouding of the lens due to water

buildup, eventually leading to blindness unless there is
surgical intervention.
• Cataracts may result from complications of diabetes,
exposure to ultraviolet light, or just natural aging.
Macular Degeneration

• Macular degeneration is a disease that destroys the fovea

and the area around it, causing a blind spot in central vision.
• It occurs in two forms, wet and dry.
• Wet macular degeneration has a very fast onset but is
partially treatable.
• It occurs because of abnormal growth of blood vessels,
leading to the leaking of blood below the retina,
causing scarring of the macula.
• Dry macular degeneration may take years to develop but
no treatment is available.
• It results from degeneration of the cells that produce
photopigments for the photoreceptors in the macula,
resulting in impaired function of the photoreceptors.
Retinitis Pigmentosa

• Retinitis pigmentosa is an inherited progressive

degenerative disease of the retina that may lead to blindness
due to degeneration of the photoreceptors, particularly rods
at the periphery.
• This results in tunnel vision at first but may result in total
blindness as it spreads to the cones.
• Presently, no treatment is available
Colour perception
• Natural sunlight varies in the distribution of
wavelengths throughout the day.
• Artificial light sources tend to be mixes of
wavelengths and can thus be classified as white light.
Key definitions
• Heterochromatic light consists of many wavelengths (white
lights), whereas monochromatic light is light of only one
wavelength or a narrow band of wavelengths.
• When monochromatic light reflects off a white surface,
that surface is seen as the colour associated with that
wavelength.
• Surfaces that reflect all light equally are achromatic (without
colour).
• These surfaces are said to be white to gray to black.
• What matters is the proportion of ambient light they
reflect.
• An important property of the visual system is that we
respond to the proportion reflected rather than the total
amount reflected.
• Hue refers to the colour quality of the light and corresponds to
the colour names that we use.
• Hue is a quality, meaning it is a value that changes but the
change does not make the value larger or smaller.
• colours associated with particular wavelengths are called
monochromatic colours, which include basic or spectral
colours (i.e., red, green, orange, yellow, blue).
• colours made of combinations of more than one
monochromatic colours are known as (e.gnonspectral .,
purple, brown, silver, gold).
• Saturation refers to the purity of light.
• The more saturated the stimulus, the stronger the colour
experience. The less saturated the stimulus, the more it
appears achromatic.
• Brightness refers to the amount of light present.
• The brighter an object is, the easier it is to see and the more
evident its colours are.
• Brightness is distinguished from lightness, which refers to the
amount of light that gets reflected by a surface.
• Additive colour mixing is the creation of a new colour by a process
that adds one set of wavelengths to another set of wavelengths.
• This occurs when we mix lights.

• Subtractive colour mixing is the creation of a new colour by the

removal of wavelengths from a light with a broad spectrum of
wavelengths.
• This occurs when we mix paints.
• It occurs when we mix substances with different absorption
spectra.
The Retina and colour

• There are three classes of cones in the fovea of the retinae.

• Each cone type contains a different photopigment and is thus
sensitive to a different band of wavelengths.
• The S-cone has a maximum response to light at 420nm (short
wavelength).
• The M-cone has a maximum response to light at 535nm (medium
wavelength).
• The L-cone has a maximum response to light at 565nm (long wavelength).
• When these three cone systems are combined, we can see colour over a
range of approximately 400 to 700nm.
• When reflected light hits the retina, there will be a pattern of responses
in the three cone systems that induce the experience of a particular
colour.
Univariance
• Univariance is the principle whereby any single cone system
is colourblind, in the sense that different combinations of
wavelength and intensity can result in the same response
from the cone system.
• That is, colour vision requires the comparative inputs of the
different cone systems.
The Trichromatic Theory of Colour
Vision
• The trichromatic theory of colour vision states that the
colour of any light is determined by the output of the three
cone systems in our retinae.
• Evidence supporting the trichromatic theory:
• colour-matching experiments show that it takes a
minimum of three primary colours to make a metameric
match to a single monochromatic light.
• There is overwhelming evidence of the existence of three
classes of cones in the human retinae.
• The trichromatic theory predicts what happens when a
person loses a cone class.
• The person still sees in colour, but cannot distinguish
between hues normal three-coned individuals can.
• In the classic form, most colour-deficient people cannot
distinguish between reds and greens due to the loss of
either M- or L-cones.
The Opponent Theory of colour
Perception
• The opponent theory of colour perception is the traditional
rival theory to the trichromatic theory.
• The opponent theory of colour perception states that colour
perception arises from three opponent mechanisms, for red-
green, blue-yellow, and black-white.
Findings That Support Opponent
Theory
• Ewald Hering proposed that colour vision was organized with
four primaries, or unique hues
• The opponent theory was advanced for the following
reasons:
• Nonprimary colours can look like combinations of two
primary colours.
• Our perception of colours supports the idea that red
and green do not combine and that blue and yellow do
not combine.
• In colour-sorting experiments, people tend to sort colours
into four basic groups (red, green, yellow, and blue) rather
than three colours.
• Colour afterimages are visual images seen after an actual
visual stimulus has been removed.
• Simultaneous colour contrast occurs when perception of
one colour is affected by a colour that surrounds it.
• This effect occurs when a colour is surrounded by its
opponent colour and not by other colours or achromatic
backgrounds.
Hue Cancellation

• Hurvich and Jameson (1957) conducted a series of important

experiments describing hue cancellation, which supports the
opponent theory.
• It is a procedure in which a subject is shown a monochromatic
reference light and is asked to remove, or “cancel,” the one of the
colors in the reference light by adding a second wavelength
(Goldstein & Brockmole, 2016).
• Some colours can be described in terms of other colours (e.g.,
orange is a mix of yellow and red) whereas unique colours can only
be described in terms of themselves (red, green, blue, yellow).
Colour Deficiency

• Colour deficiency refers to the condition of individuals who

are missing one or more of their cone systems.
• Colour deficiencies are usually the result of genetic defects
and are more common in men because the genes leading to
colour deficiency are on the X chromosome.
• Colour deficiency is most often determined using Ishihara
plates, in which different coloured, but isoluminant dots are
presented.
Rod Monochromacy

• Rod monochromacy is a very rare form of colour deficiency,

affecting 1 in every 30,000 people.
• No functioning cones of any kind are present, resulting in
true colourblindness.
• Individuals with rod monochromacy:
• See the world in shades of gray.
• Are very sensitive to bright light because they always use
scotopic vision.
• Have very poor visual acuity
Cone Monochromacy

• Cone monochromacy is an extremely rare form of colour

deficiency, affecting 1 in every 100,000 people.
• Only one functioning cone system is present.
• S-cone monochromats have been observed.
Dichromacy

• Dichromats have two working cone systems.

• They see in colour, but cannot make some of the
discriminations that are easy for trichromats.
• There are three major forms of dichromacy: protanopia,
deuteranopia, and tritanopia.
Protanopia
• A protanope perceives short-wavelength light as blue, and as
wavelength is increased, the blue becomes less and less saturated
until the protanope perceives gray eg when one enters the disco
room.
• The wavelength at which the protanope perceives gray is called
neutral point.
• A deuteranope perceives blue at short wavelengths, sees yellow at
long wavelengths and has a neutral point about 498nm (Boynton,
1979).
• Tritanope sees blue at shortwavelength, red at long wavelenghths
and a neutral point at 570 nm (Alpern et al, 1983).
• A very few people have unilateral dichromacy, in which
there is dichromacy in one eye but normal trichromatic
vision in the other eye.
• Unilateral dichromacy allows researchers to understand
the true nature of dichromacy.
Cortical Achromatopsia

• In cortical achromatopsia, loss of colour vision occurs

because of damage to the occipital lobe, usually from
damage to V4.
• The primary symptom is seeing in black, white, and gray or
having the impression that colours are washed out.
• Sometimes, patients can still discriminate by wavelength.
• In other cases, patients lose the ability to remember colour
(Schwartz & Schwartz).
Colour Constancy
• Colour constancy refers to the ability to perceive the colour
of an object despite changes in illumination.
• A system that sees an object as a constant colour across such
changes leads to accurate perception.
• There are several situations in which colour constancy fails.
• Monochromatic light allows an object to only reflect that
wavelength and appear the colour of that light.
• When an object is viewed in front of a pure black
background, it makes it difficult for the visual system to
get the context to see the object as the right colour.
Depth perception
• Our visual system must re-create the 3D world using an
essentially flat, 2D retina.
• The solution that our visual system uses is known as the cue
approach to depth perception.
• We must infer the third dimension from other cues our visual
system provides.
• A cue is a factor that aids you in making a nonconscious and
automatic decision.
• Cues tell us about which objects are closer and which are
farther away.
Cue Approach to Depth Perception
• These cues include oculomotor cues such as
accommodation and convergence, monocular cues such
as occlusion, relative height, and relative size, and
binocular cues from comparing images from each retina.
Oculomotor Cues

Cues based on our ability to sense the position of our eyes and
the tension in our eye muscles (Goldstein & Brockmole, 2016).
Accommodation

• Accommodation is the process of adjusting the lens of the

eye so that one can see both near and far objects easily.
• To focus on more distant objects, the ciliary muscles that
contract the lens are relaxed.
• To focus on more near objects, the ciliary muscles are
contracted.
• These contractions and relaxations take place automatically,
but they can be sensed.
• It is the sensing of these movements that gives information
about depth.
Vergence (or Convergence)

• Vergence (also known as convergence) happens when the

eyes bend inward to see a near object and bend outward
(diverge) when looking at a more distant object.
• Like accommodation, this process is automatic but the
movements are sensed, giving us information about relative
depth.
• Vergence is probably a more useful cue than
accommodation, providing the visual system with reliable
depth information to about 2m in length.
Monocular Depth Cues

• Monocular depth cues are cues that provide information

about depth and distance but can be inferred from just a
single retina.
• Monocular cues include pictorial cues, from which we can
judge depth from static or nonmoving pictures, and
movement-based cues, in which moving objects let us make
inferences about depth.
• Visual distance and depth cues that require the sense of only
one eye are called monocular cues.
Occlusion (or Interposition)

• Occlusion happens when one object partially hides or

obstructs the view of a second object.
• Occlusion provides information about relative, not absolute,
distance.
• We infer that the hidden object is farther away from us than
the object that obstructs it.
Relative Height

• Relative height means that objects closer to the horizon are

seen as more distant.
• Objects below the horizon are seen as more near the viewer
if they are closer to the bottom of the visual scene but
objects above the horizon are seen as more near if they are
closer to the top of the visual scene (Schwartz & Kranz,
2016).
Relative Size

• Relative size means the more distant an object, the smaller

the image will be on the retina.
• Thus, if there are two identical objects, the one that is closer
will have the larger image on the retina.
• However, the object that is further away does not look
abnormally small due to the mechanism called size
constancy.
Familiar Size

• Familiar size means we judge distance based on existing

knowledge of the sizes of objects.
• If we know that a particular object is smaller than another
object, but it is taking up more space on the retina, we
assume the smaller object is closer and the larger object is
farther away.
Linear Perspective

• Linear perspective is the pictorial depth cue that arises from

the fact that parallel lines appear to converge as they recede
into the distance.
• Though parallel lines never meet in actuality, perceptually
they do at the edge of the horizon.
• A visual neuron that is most responsible to a line moving in a
particular direction is called complex cells
• A neuron in visual cortex that responds best to lining of a
particular width, orientation and position is called simple
cells
• Simple cells within visual cortex respond to lines of particular
qualities
• Parallel lines seem farther apart close up, because they take
up more space in the image on the retina.
• Linear perspective is a monocular distance cue, but is based
on the apparent merging lines with increased distance.
• Eg, the walls of a very long, narrow corridor appear to come
together as they recede from the viewer, and this distance
cue is called linear perspective.
• Linear perspective is an important technique when painting a
3D scene.
• Complex cells are similar to simple cells in responsiveness, but they
are not selective to the position of a stimulus in the receptive field.
• Hypercomplex cells differ from simple cells because they respond to
lines of specific lengths
Texture Gradients

• Texture gradients occur because textures become finer and

smoother as they recede in distance
• Common elements that are evenly spaced in an image
appear more close together in the distance than they do in
the foreground.
Atmospheric Perspective

• Atmospheric perspective is a pictorial depth cue that arises

from the fact that objects in the distance appear blurred and
tinged with blue.
• Because the atmosphere scatters light, more distant objects
appear blue.
Shadows and Shading

• Shadows are decreases in light intensity caused by the

blockage of light (Goldstein & Brockmole, 2016).
• Shadows are a depth cue arising because an object is in front
of its shadow, and the angle of the shadow can provide some
information about how far the object is in front of the
background.
• Objects in shadow must be farther from the light than
objects that are not in shadow.
Motion Cues

• Objects moving at different speeds can reveal information about

relative distance.
Motion parallax
• Motion parallax is a monocular depth cue arising from the
motion of a person in the environment.
• Faster moving objects are closer to us; slower moving
objects are farther away.
Deletion and accretion
• Deletion is the gradual occlusion of a moving object as it
passes behind another whereas accretion is the gradual
reappearance of a moving object as it emerges from behind
another object.
• The object that is deleted and later accreted is the object
that is farther away than the object we can see
continuously.
Optic flow
• Optic flow is the motion depth cue that refers to the relative
motions of objects as a person moves forward or backward.
• Like with motion parallax, closer objects appear to move
more quickly and farther away objects appear to move
more slowly.
• Optic flow is often used to convey depth in movies
Binocular Cues to Depth

• Because humans have two eyes that see the world from
slightly different angles, we have an important cue to depth.
• In the area where both eyes see the same part of the world,
we have binocular vision.
• Stereopsis is the sense of depth that we perceive from the
visual system’s processing of the comparison of the two
different images from each retina.
Demonstration
• Hold one hand out at arm’s length and look at your own
thumb. First look at that thumb with your left eye and then
with your right eye.
Results
• You get an illusion that your hand has shifted somewhat as you go
back and forth from one eye to the other. This quick test allows us to
see the slightly different perspective on the world that each eye gives
us. If you have an object to look at just beyond your thumb, you can
see that the object may appear closer to your thumb through one eye
than through the other. This is binocular disparity, which helps
provide the basis for the determination of depth.
Binocular Disparity

• Disparity occurs because our two eyes are in different locations in our
head and thus have slightly different views of the world.
Size Perception

• Size-distance invariance refers to the relation between

perceived size and perceived distance.
• The perceived size of an object depends on its perceived
distance, and the perceived distance of an object may
depend on its perceived size.
• Visual angles, the angles of objects relative to one’s eyes, are
important to studying size-distance invariance.
 • If lines were drawn from the top and bottom of an object
to the eyes, we would have the visual angle of the object.
• Smaller objects close up can have the same visual angles
as larger objects farther away, like the moon and the sun.
• Depth provides information that allows us to appropriately
scale the size of an object.
 Size Constancy

• Constancy is the tendency to experience a stable view of the world,

even in a changing sensory environment(eg shape, color r
unchanged even during the night)
• Size constancy is the perception of an object as having a fixed size,
despite the change in the size of the visual angle that accompanies
the change in distance.
• We tend to see an object as the same size regardless of the size of
its image on the retinae.
• Size constancy normally helps us maintain a stable perception of
objects by taking distance into account
Visual Illusions of Size and Depth

• By manipulating depth cues, it is possible to see depth relations that

are not really present in an image.
The Müller-Lyer Illusion

• In this illusion, although both lines are exactly the same

length, one of the lines seen as longer than the other one
due to the smaller lines that split off from the main line.
• One viewpoint for why we see this illusion is that it is a result
of misapplied size constancy.
• The visual system wants to keep objects at a fixed size.
• The very mechanisms that help us maintain stable
perceptions in the three-dimensional world sometimes
create illusions when applied to objects drawn on a two-
dimensional surface (Goldstein & Brockmole, 2016).
• Another viewpoint is that the overall image of the line that
has outward oriented fins is longer and, thus, its components
are also perceived as longer.
The Ponzo Illusion

• In this illusion, the cow further away from the observer looks
bigger than the cow in the foreground.
• There are several cues in the photograph that give normal
indications of depth, such as the parallel lines of the side of
the road, the texture gradient of flowers and grasses, and
familiar size cues of the trees.
• The Ponzo illusion is a strong example of misapplied size
constancy as well as the influence of linear perspective on
size perception.
The Ames Room Illusion

• Two people of normal size are put into corners of a room

with trapezoidal walls and windows.
• When viewed through a peephole, the trapezoids are
converted into squares by our visual systems so that we
perceive the room as normal.
• Given that we perceive a normal room, we therefore infer
that the people are abnormally short or tall.
The Moon Illusion

• In this illusion found in natural phenomena, a full moon on

the horizon is perceived as being larger than when it is
higher up in the sky, though its size and distance from earth
remain constant.
• Because the horizon is perceived as being farther away than
the zenith overhead, an object that takes up the same
amount of space on our retina is perceived to be larger.