Chapter 6: The Visual System

The visual system does not create an exactly accurate recreation of the external world for us to
interpret
The information projected onto the visual receptors is distorted, upside-down and twodimensional.
From this information the brain creates a representation of what is 'out there'
Two types of research inform us about our visual system:
Research that explores the visual system with neuroanatomical, neurochemical, and
neurophysiological techniques
Research that focuses on the assessment of what we see
6.1: Light Enters the Eye and Reaches the Retina
Some animals have adaptations that allow them to see in very dim light, but no visual system is
able to work in total darkness
Light: Often defined as waves of electromagnetic energy that are between 380 and 760
nanometers in length.
Light has some properties of waves and some properties of particles.
This range is the spectrum we are able to perceive with our eyes, however some animals are
capable of perceiving wavelengths we cannot.
Two properties of light are of particular interest:
Wavelength: Used interchangeably with colour.
Intensite: Used interchangeably with brightness.
The Pupil and the Lense:
How much light reaches the retinas is regulated by a donut-shaped band of contractile tissue
called the iris.
Light enters the eye through the pupil, the hole in the center of the iris.
Sensitivity: The ability to detect the presence of dimly lit objects
Acuity: The ability to see details of objects
Pupil size is adjusted in response to light in order to balance sensitivity and acuity.
High illumination (ie. Sensitivity is not important) leads to constricting of the pupils to
allows for the greatest acuity.
Low illumination (ie. Sensitivity is important) leads to dilation of the pupils to allow for
greater sensitivity at the expense of acuity.
Constriction of the pupils allows a greater depth of focus, meaning a greater range of depth
differences are able to be kept in focus simultaneously.
Behind each pupil is a lens which focuses incoming light onto the retina.
Accomodation: The process by which the configuration of the lenses of the eyes bring
images into focus.
Ciliary Muscles: Muscles which hold the lens of the eye in place and adjust its tension
to allow us to focus either nearby or far away.
When we observe something near, the lens assumes its natural cylindrical shape,
allowing the lens to refract the light and bring close objects into focus.
When we observe something far away, the lens is flattened which reduces refraction and
brings distant objects into focus.
Eye Position and Binocular Disparity
One reasons why vertebrates have two eyes is because they have two sides

 By having an eye on each side of the head the most common vertebrate occular
arrangement animals can see in almost every direction without moving their heads.
Having both eyes on the front side of the head sacrifices being able to see what is going on
behind us for being able to see what is going on in front of us much more clearly.
In general, predators tend to have front-facing eyes because it enables them to accurately
perceive how far away their prey are.
Prey animals tend to have side facing eyes because this arrangement gives them a larger
field of vision and the ability to perceive oncoming predators from any direction.
The movements of your eyes are coordinated so that each point in your visual world is
projected onto corresponding points on each of the two retinas.
Because both eyes must see corresponding images, your eyes must turn in slightly.
Because your eyes are in two different places, you cannot actually have the same image
on both eyes at the same time.
Binocular Disparity: Difference in the position of the same image on the two retinas.
Binocular disparity is larger for close objects than for faraway objects.
The brain uses binocular disparity to create a three dimensional representation of our
world based on two dimensional images captured by the retinas.
6.2: The Retina and Translation of Light into Neural Signals
The retina converts light to neural signals, conducts them towards the CNS, and participates in the
processing of the signals.
Receptors: Cells that are specialized to receive chemical, mechanical or rediant signals from the
environment; Also proteins that contain binding sites for particular neurotransmitters.
o Fifth and final layer of ocular cells
Horizontal Cells: Type of retinal neurons whose specialized function is lateral communication.
o Specialized for lateral communication (Communication across the major channels of sensory
input)
o Fourth layer of ocular cells
Bipolar Cells: Bipolar neurons that form he middle layer of the retina.
o Third layar of ocular cells
Amacrine Cells: A type of retinal neuron whose specialized function is lateral communication.
o Specialized for lateral communication (Communication across the major channels of sensory
input)
o Second layar of ocular cells
Retinal Ganglion Cells: Retinal neurons whose axons leave the eyeball and form the optic nerve.
o First layar of ocular cells
o Project across the inside of the retina before gathering together in a bundle and exiting the eyeball.
Each of these cell types comes in a variety of subtypes. (> 50 unique cell types have been identified
in the retina!)
Retinal neurons communicate both chemically via synapses and electrically via gap junctions.
Light reaches the receptor layer only after passing through the other four layers.
Once the receptors have been activated, the signal is transmitted back out through the retinal layers
to the retinal ganglion cells.
This arrangement of signal reception and transfer results in two problems:

o Incoming light is distorted by the retinal tissue because it must pass through layars of tissue
before it reaches the receptors.
o In order for the bundle of retinal nerves to leave the eye, there must be a gap in the receptor
layar.
o Blind Spot: The gap in the receptor layar where the bundle of retinal nerves exits the eye.
Fovea: Indentation about 0.33cm in diameter and the center of the retina.
o Specialized for high-acuity vision (ie. the seeing of fine details)
o The thinning of the retinal ganglion cell layer at the fovea reduces the distortion of incoming
light.
Completion: The visual system;s automatic use of information obtained from receptors around the
blind spot, or scotoma, to create a perception of the missing portion of the retinal image.
o When the visual system detects a straight bar going into one side of the blind spot and another
straight bar leaving the other side, it fills in the missing bit for you, regardless of what is
actually there.
o Completion phenomenon is evidence that the visual system does more than create a faithful
copy of the external world.
o Completion is not a reaction to retinal blind spots, it occurs in all areas of vision.
o Our eyes detect key features which are transmitted to the brain (most importantly edges and
location) where a recreation of the expected object is generated based on that partial
information.
o Surface Interpolation: The process by which we perceive surfaces; The visual system extracts
information about edges and from it infers the appearance of large surfaces.
Cone and Rod Vision:
o Two different types of receptors in the the human retina
Cones: Cells in the human retina that are responsible for photopic vision.
Species active only during the day tend to have cone-only retinas.
Short and stocky appearing cells
In the center of the fovea there are exclusively cones
Cone density decreases as you look closer to the perimiter of the foveal indentation.
Rods: Cells in the human retina that are responsible for scotopic vision.
Species active only at night tend to have rod-only retinas
Long and 'fibrous' appearing cells
There are no rods in the fovea
Around the outside of the foveal indentation there are more rods and significantly less
cones
Highest concentration of rods is located at 20 degrees from the center of the fovea.
There are more rods in the nasal hemiretina (the half of the retina closer to the nose)
than in the temporal hemiretina( the half of the retina closer to the temples)
o Duplexity Theory: The theory that cones and rods mediate different kinds of vision.
o Photopic Vision: Predominates in good lighting and provides good acuity. Allows for detailed
and coloured perceptions of the world.
In a photopic vision system, the output of only one or two cones converge are on a single
retinal ganglion cell.
It takes more light to trigger a reaction from a cone, but when it does react there is less
ambiguity than in scotopic vision about where the stimulus which triggered the reaction.

o Scotopic Vision: Predominates in dim illumination. Lower acuity and lacks details and colour,
but allows sight in very low levels of light.
Vision is used when there is not enough light to excite cones.
In a scotopic system, the output of several hundred rods converge onto a single retinal
ganglion cell.
Because the information from many cells is summed to create a single image, all of the light
information is used in a situation of dim light. (Dim light would not be enough for a single
cone to cause the retinal ganglion cell it is attached to to fire)
High degree of sensitivity, low degree of acuity
When a retinal ganglion cell that receives input from hundreds of rods changes its firing, the
brain has o way of knowing which portion of the rods contributed to the change.
Spectral Sensitivity
o Generally more intense light appears brighter, but wavelength also has an effect on how bright
we perceive light to be.
o Our visual systems are not equally senstive to all wavelenths in the visible spectrum.
o Lights of the same intensity but of different wavelengths can appear to be very different
'brightnesses'
o A graph of relative brightness of lights of the same intensity (but at different wavelengths) is
called a spectral sensitivity curve.
o Animals with both cones and rods have two spectral sensitivity curves:
Photopic Spectral Sensitivity Curve: The graph of the sensitivity of cone-mediated vision
to different wavelengths of light.
Can be determined in humans by having subjects judge the relative brightness of
different wavelengths of light shone on the fovea.
Maximally sensitive to wavelengths of about 560 nm (ie. a light at 500 nm would have
to be much less intense than a light of 560 nm in order for them to appear as the same
brightness.)
Scotopic Spectral Sensitivity Curve: The graph of sensitivity of rod-mediated vision to
different wavelengths of light.
Can be determined in humans by asking subjects to determine the relative brightness of
different wavelengths of light shone on the periphery of the retina at an intensity too low
to activate te few peripheral cones that are located there.
o Purkinje Effect: In intense light, red and yellow wavelengths look brighter than blue or green
wavelengths of equal intensity, in dim light, blue and green wavelengths look brighter than red
and yellow wavelengths of equal intensity.
Eye Movement
o The eyes continually scan the visual field, and our visual perception at any instant in s a
summation of recent visual information.
o It is because of this temproal integration that the world does not vanish momentarily each time
we blink.
o Even when we fix our gaze, our eyes continuously move.
o Fixational Eye Movement: Involuntary eye motions that allow us to see during fixed
observation by keeping images moving on the retina.
Three kinds of fixational eye movement
Tremors
Drifts

Saccades: Small jerky movements or flicks

o We are normally unaware of our constant eye movements.
o If we were to actually fix our eyes, our vision would fade and disappear, since visual neurons
respond to change.
o If retinal images are artificially stabalized, parts of the image begin to disappear and reappear.
Visual Transduction: The Conversion of Light to Neural Signals
o Transduction: Conversion of one form of energy to another
Visual transduction is the conversion of light to neural signals by the visual receptors.
o Rhodopsin: Red pigment extracted from rods. Discovered in 1876.
When exposed to conintuous intense light, it became bleached (ie. lost its colour)
When returned to the darkness, it regained its redness and light absorbing capacity.
Rhodopsin's absorbtion of light is the first step in rod-mediated vision.
The rate at which rhodopsin absorbs lights of different wavelengths is related to the ability
of animals with rods to detect the presence of different wavelengths of light under scotopic
conditions.
Our sensitivity to light is a direct consequence of rhodopsin's ability to absorb varying
wavelengths of light.
G-protein-coupled receptor that responds to light rather than neurotransmitter molecules.
Like other G-protein-coupled receptors, they initiate a cascade of intracellular chemical
events when they are activated.
When rods are in darkness, sodium channels are partially open, thus keeping the rods
slightly depolarized and allowing a steady flow of excitatory glutamate neurotransmitter
molecules to emanate from them.
When rhodopsin receptors are bleached by light, the resulting cascade of intracellular
chemical events closes the sodium channels, hyperpolarizes the rods and reduces the release
of glutamate.
Signals are often transmitted through neural systems by inhibition.
o Absorbtion Spectrum: A graph of the ability of a substance to absorb light of different
wavelengths.

6.3: From Retina to Primary Visual Cortex

Many pathways in the brain carry visual information
Retina-Geniculate-Striate Pathways: Most studied and largest visual pathway; Pathway which
conducts signals from each retina to the visual cortex.
o Lateral Geniculate Nuclei:
Each lateral geniculate nucleus has six layars
Each layar receives input from all parts of the contralateral visual field for one eye
Each lateral geniculate nucleus receives visual input from the contralateral visual field: three
layars from one eye and three layars from the other
Most of the laterak geniculate neurins that project to the primary visual cortex terminate in
the lower part of cortical layar IV, producing a characteristic strike when viewed in a cross
section.
o Retina-geniculate-striate pathway conducts signals through the lateral geniculate nuclei.
o 90% of retinal ganglion cells become part of the retina-geniculate-striate pathway
o No other sensory system has such a predominant pair of pathways (right and left) to the cortx.

o All signals from the left side of the visual field are transmitted to the right hemisphere for
processing either
Ipsilaterally from the temporal hemiretina of the left eye
Contralaterally from the nasal hemiretina of the right eye via the optic chasm
o The reverse is true for the right side
Retinoscopic Organization
o Retinotopic: Mapped like the retina.
o Each area of the retina-geniculate-striate system is retinoscopic.
o Two stimuli presented to adjacent areas of the retina excite adjacent neurons at all levels of the
system.
o The retinotopic layout of the primary visual cortex has a disproportionately large area dedicated
to the fovea.
25% of the visual cortex is devoted to analyzing signals from the fovea
The M and P Channels
o There are at least two parallel pathways of information through the lateral geniculate nucleus
o Parvocellular Layars: Top 4 layars of l.g.n. First pathway of communication
Called parvocellular because it is composed of small cells (parvo means small)
Particularly responsive to colour, details and still images
Input is provided mainly by cones
o Magnocellular Layars: Bottom 2 layars of l.g.n.. Second pathway of communication.
Called magnocellular because it is composed of large cells (magno.p means big)
Input is provided mainly by rods.
o Parvocellular and magnocellular pathways project to different sites in the striate cortex
o In turn, these M and P portions project to different parts of the visual cortex.

6.4: Seeing Edges

Edges are the most important features of any visual display because they define the extent and
position of the various objects in it
In a sense a visual edge is nothing more than a point where two different areas of a visual image
meet.
The perception of edges is really the perception of a contrast between two adjacent areas of visual
field.
Lateral Inhibition and Contrast Enhancement
o Mach Bands run adjacent to the edges of brightness contrast in order to allow us to more easily
perceive an edge.
o Contrast Enhancement:
Every edge we look at is contrast enhanced
Experiments of the physiological basis of contrast enhancement were conducted on the eyes
of a horseshoe crab
Horseshoe crab eyes are made up of very large receptors, called ommatidia.
Each ommatidium has its own large axon.
Axons of the ommatidia are interconnected by a lateral neural network.
Ifa singe ommatidium is illuminated, it fires at a rate that is proportional to the intensity of
the light striking it. (ie. more intense light produces more rapid firing)
Lateral Inhibition: When a receptor fires, it inhibits its neighbours via the lateral neural
network.

Amount of lateral inhibition produced by a receptor is greatest when the receptor is most
intensely illuminated
Lateral inhibition has the greatest effect on its immediate neighbours
Receptor adjacent to the edge on the more intense side fires more than other neurons fired
by the light of the same intensity.
Other receptors on the side of the intense light receive the same rate of illumination and
the same rate of lateral inhibition.
Receptors directly adjacent to those that are receiving dim light receive the same rate of
illumination but a lowered rate of lateral inhibition since it is not inhibitted by the
receptor receiving a lower rate of illumination.
The reverse is true for the receptors on the side with dim illumination.
Receptor Fields of Visual Neurons
o David Hubel and Torsten Wiesel (2004) invented a very influential method for studying the
neural mechanisms of vision.
o Their technique analyzes th firing pattern of a single neuron in the occular systme of laboratory
animals.
o The tip of a microelectrode is positioned near a single neuron in the part of the visual system
that is under investigation.
o During testing eye movements are inhibited by paralyzing the eye muscles.
o Images on a screen are focuesed sharply onto the retina by an adjustable lens
o Receptive Field: The area of the visual field within which it is possible for a visual stimulus to
influence the firing of that neuron.
The next step in the procedure is to determine the receptive field of the neuron
o Final step is to record the responses to the neuron to various stimuli within its receptive field in
order to determine what kinds of stimulation influence that neuron's activity.
o The electrode is then advanced to the next neuron and the process is repeated.
o Strategy is to work from the neurons nearest to receptors and work up through progressively
"higher" levels of the system, with the goal of understanding the increasingly complex neural
responses at each level of the system.
Receptive Fields: Neurons of the Retina-Geniculate-Striate System:
o When Hubel and Wiesel compared the receptive fields of the retinal ganglion cells, lateral
geniculate nuclei, and lower layar IV neurons, they found four things in common:
At each level, the receptive fields in the foveal area of the retina were smaller than those at
the periphery.
All neurons had receptive fields that were circular.
All neurons were monocular - each had a receptive field in one eye but not the other.
Many neurons at each level had receptive fields that comprised an excitatory area and an
inhibitory area separated by a circular boundary.
There is an area of the receptive field that caused the neuron to fire when a light was
present and illuminated
There is also an area of the receptive field that caused the neuron to inhibit firing when
the light was present and illuminated, but displayed a burst of firing when the light was
removed or turned off.
For most neurons, the pattern of "on" firing and "off" firing was quite predictable
depending on whether they were on-center cells or off-center cells.

On-Center Cells: Respond to light shone in the central region of their receptive fields
with "on" firing and to lights shone in the periphery of their receptive fields with
inhibition followed by "off" firing when the light is turned off.
Off-Center Cells: Respond with inhibition followed by "off" firing to lights on the
center of the field, and "on" firing to lights the periphery of the field.
On-center and off-center cells respond best to contrast. The optimal way to increase
firing of these neurons is to maximize the contrast between the center and the peripery
of the receptive field by illuminating either region while leaving the other region
completely dark.
Diffusely illuminating the entire receptive field has little effect on firing.
Hubel and Wiesel concluded that one function of many of the neurons in the retinageniculate-striate system is to respond to the degree of brightness contrast between te
two areas of their receptive fields.
o Most visual system neurons are continually active, even when there is no visual input.
o Spontaneous activity is a characteristic of most cerebral neurons, and responses to external
stimuli consume only a small portion of the constant energy required by ongoing brain activity.
(Raichle, 2006)
o The level of activity of visual cortical neurons at the time that a visual stimulus is presented
influences how the cells respond to the stimulus - one way cognition may influence perception.
(Arieli et al., 1996)
Receptive Fields: Simple Cortical Cells
o The neurons of lower layar IV are exceptions. Their receptive fields are unlike any other neuron
in the visual system.
o Most other neurons fall into one of two categories:
Simple cells
Complex cells
o Simple Cells: Cells with receptive fields that can be divided into antagonistic "on" and "off"
regions; Do not respond well to diffuse light; Simple cells are all monocular; Borders between
"on" and "off" regions are straight instead of circular.
Simple cells respond best to bars of light in a dark field, dark bars in a light field or single
straight edges between dark and light areas.
Each simple cell responds maximally when the straight-edge stimulus is in a particular
position and in a particular orientation.
Receptive fields of simple cells are rectangular rather than circular.
Receptive Fields: Complex Cortical Cells
o Complex Cortical Cells:
Complex cells are more numerous than simple cells
Complex cells share some properties with simple cells.
Rectangular receptive fields
Respond best to straight-line stimuli in specific orientations
Unresponsive to diffuse light.
Complex cells differ from simple cells in three key ways:
Larger receptive fields
It is not possible to divide the receptive fields into static "on" and "off" regions
o Complex cells respond to a particular straight-edge stimulus of a particular
orientation regardless of its position within the receptive field of that cell

o A stimulus which produces "on" firing that is swept across the cell's receptive field
will create a continuous response as it moves.
o Many complex cells respond more robustly to the movement of a straight line across
their receptive filds in a particular direction.
Most complex cells are binocular, responding to stimulation from both eyes.
o Receptive fields in each eye are almost identical to each other
o If the appropriate stimulation is applied through both eyes simultaneously a
binocular cell usually fires more robustly than if only one eye is stimulated.
o Most binocular cells show some degree of ocular dominance, favouring one eye or
the other by firing more robustly to stimlation of the favoured eye.
o Some binocular cells fire best when the preferred stimulus is presented to both eyes
at the same time, but in slightly different positions on the two retinas.
These cells respond best to retinal disparity and likely play a role in depth
perception.
Columnar Organization of Primary Visual Cortex
o The characteristics of the receptive fields of visual cortex neurons are attributable to the flow of
signals from neurons with simpler receptive fields to those with more complex fields.
o It seems that signals flow from on-center and off-center cells in lower layer IV to simple cells
and from simple cells to complex cells.
o Primary visual cortex neurons are grouped into functional vertical columns.
o If an electrode is advanced vertically through the layers of the visual cortex the results show
that each cell in the column has a receptive field in the same area of the visual field.
o All the cells in a column respond best to straight lines in the same orientation.
o Neurons in a column that are either monocular or binocular with occular dominance are all most
sensitive to light in the same eye.
o If an electrode is advanced horizontally through the tissue of the primary visual cortex, each
successive cell is likely to have a receptive field in a slightly different location and to be
maximally responsive to straight lines of a slightly different orientation.
o During a horizontal electrode pass, the tip passes alternately through areas of left-eye and righteye dominance. These are commonly referred to as occular dominance columns.
o All of the functional columns in the primary visual cortex that analyze input from one area of
the retina are clustered together.
Half of the cluster is thought to receive visual information from the left eye and the other
half from the right.
Each cluster is thought to include neurons with preferences for straight-line stimuli of
various orientations.
Plasticity of Receptive Fields of Neurons in the Visual Cortex
o Most neuroscientific research is based on two assumptions:
Mechanisms of visual processing can be identified by studies using simplified, artificial
stimuli.
The receptive field properties of each neuron are static, unchanging properties
o Studies of the responses of visual cortex to natural scenes suggest that neither of these
assumptions are particularly true.
The response of a visual cortex neuron depends not only on the stiumli in its receptive field,
but on the larger scene in which these stimuli are embedded.

The ability to adapt to changing visual stimuli has been largely ignored, but appears to be a
fundamental property of the visual cortex.
Research based solely on the study of reaction to simple stimuli cannot provide a complete
explanation of how the visual system works - receptive field properties depend on the scene
in which the stimuli to its field are embedded.

6.5: Seeing Colour

Black is experienced when there is an absense of light.
The perception of white is produced by an intense mixture of a wide range of wavelengths in
roughly equal proportions.
The perception of gray is produced by the same mixture with a lower intensity.
Correct term for colour is hue.
To a larg degree, an object's colour depends on the wavelengths of light that it reflects into the eye.
Most objects absorb the different wavelengths of light that strike them to varying degrees and
reflect the rest.
The mixture of wavelengths that objects can reflect influences our perception of their colour.
Component and Opponent Processing
o Component Theory: The theory that there are three different kinds of colour receptors, each
with a different spectral sensitivity and that the colour of a particular stimulus is presumed to be
encoded by the ratio of activity in the three kinds of receptors.
Also called trichromatic theory;
Proposed by Thomas Young in 1802 and refined by Hermann von Helmholtz in 1852
Theory is based on the observation that any colour in the visible spectrum can be recreated
by mixing together three different wavelengths of light in different proportions.
This can be accomplished by any three wavelengths in which the colour of any one of them
cannot be matched by a mising of the other two.
o Opponent-Process Theory: Theory suggesting that there are two different classes of cells in
the visual system for encoding colour and another class for encoding brightness.
Proposed by Ewald Herring in 1878.
Hypothesized that each of the three classes of cells encoded two complementary colour
perceptions.
One class of colour coding cells signaled red by changing its activity in one direction and
signaled red's complementary colour green by canging its activity in the other direction (ie.
hyperpolarization)
Another class of colour-coding cells was hypothesized to signal blue and its compliment,
yellow.
Class of brightness encoding cells was hypothesized to signal both black and white.
Complementary Colours: Pairs of colours that prduce white or gray when combined in
equal measure.
Theory is based off of several observations
Complementary colours cannot exist together - there is no such thing as bluish yellow or
reddish green
The afterimage produced by staring at red is green and the afterimage produced by
staring at yellow is blue (The reverse is true for both examples)
o Subsequent research showed that portions of both opponent process theory and component
theory coexist in our visual systems.

o The development of a technique for measuring the absorption spectrum of te photopigment

contained in a single cone allowed researchers to scientifically test both theories.
There are indeed three different types of cones in the retinas of vertibrates with strong
colour vision.
Each of the three has a different photopigment with its own characteristic absorption
spectrum.
Some cones are most sensitive to short wavelengths, some to medium wavelengths and
others to long wavelengths
Colour coding by cones appears to happen by the rules of component theory, but subsequent
processing of colour information in the retina-geniculate-striate uses both component and
opponent-processing theory.
In all subsequent levels, there are cells that respond in one direction (ie. increased firing) to
one colour and in the opposite direction (ie. decreased firing) for its complementary colour.
o Most primates are trichromats, which means that we possess three colour vision photopigments.
o Most other mammals are dichromats - posessing only two colour vision photopigments.
These animals lack sensitivity to long wavelengths and thus have difficulty seeing light at
the red end of the visible spectrum.
o Some birds, fish and reptiles have four photopigments.
These animals are able to detect ultraviolet light, which is invisible to humans.
o In an amazing study, mice were given the gene for long-wavelength photopigment
development, effectively converting them from dichromat to trichromat.
Behavioural tests indicated that the transgenic mice had acquired the ability to perceive long
wavelengths and make colour discrimination involving light at that end of the spectrum.
Colour Constancy and the Retinex Theory
o Colour Constancy: The tendency of an object to appear the same colour ven when the
wavelengths of light it reflects change.
Single most important characteristic of colour vision
Colour constancy improves our ability to tell objects apart in a memorable way so that we
can respond appropriately to them;
Our ability to recognize objects is heightened by colour constancy.
We are normally completely unaware of colour constancy.
Colour constancy occurs as long as the object is illuminated with light that contains some
short, medium and long wavelengths (such as daylight, firelight and most artificial lighting)
and as long as the object is viewed as part of a scene and not in isolation.
o Experiment to test colour constancy involved three adjustable projectors, each emitting only
one wavelength of light: short, medium or long.
o Long shone the three projectors onto a display of blocky colours called a Mondrian
o Adjusting the amount of light emitted from each projector (and thus, how much light each
coloured rectangle relfected) made no difference in subjects' perception of colour.
o Retinex Theory: Theory which states that the colour of an object is determined by its
reflectance - the proportion of light of different wavelenths that a surface reflects.
The wavelengths of light reflected by a surface can change dramatically with changes in
illumination
The efficiency with which a surface absorbs each wavelength and reflects the unabsorbed
portion, however, remains constant

According to the retinex theory, the visual system calculates the reflectance of surfaces and
thus perceives their colours by comparing the light reflected by adjacent surfaces in the
three different wavelength bands (long, medium, short)
o Dual-Opponent Colour Cells: Neurons which are responsive to colour contrast.
Respond with vigorous "on" firing when the center of the circular receptive field is
illuminated with one wavelmength, ad the peripheral field is illuminated with another
wavelength.
Respond with "off" firing when the pattern of illumination of colours is reversed.
dual-opponent colour cells are not distributed uniformly throughout the visual cortex, with
the exception of lower layar IV.
These neurons are concentrated in the primary visual cortex in peglike columns that
penetrate the layers of the monkey primary visual cortex.
Most of the neurons in these peglike columns are particularly rich in cytochrome oxidase, so
distribution of the pegs can be visualized by staining slics of tissue with stains that have an
affinity for this enzyme.
Cytochrome Oxidase:
Blobs: Peglike, cytochrome oxidaserich, dual-opponent colour columns..
Located in the midst of ocular dominace columns.
fMRI studies have provided evidence of dual-opponent colour cells in the human visual
cortex.

6.6: Cortical Mechanisms of Vision and Conscious Awarenes

Visual Cortex is considered to be in three types:
o Primary visual cortex: The area of the sensory cortex that receives direct input from the lateral
geniculate nuclei (Also called the striate cortex)
Located in the posterior region of the occipital lobes
Mostly hidden by the longitudinal fissure
o Secondary visual cortex: Areas of the crebral cortex that receive most their input from primary
visual cortex
Located in the prestriate cortex and the inferotemporal cortex
Prestriate Cortex: Band of tissue in the occipital lobe that surrounds the primary visual
cortex.
Inferotemporal Cortex: Cortex of the inferior temporal lobe.
o Visual association cortex: Areas of the cortex that receive input from areas of the secondary
visual cortex as well as from secondary cortex of other sensory systems.
Located in several parts of the cerebral cortex, but the largest area is in the posterior parietal
cortex.
Posterior Parietal Cortex:
Major flow of visual information in the cortex is from the primary visual cortex to the various areas
of secondary visual cortex to the areas of association cortex.
As you move up the visual hierarchy, neurons have larger receptive fields and the stimuli to which
the neurons respond to are more specific and more complex.
Damage to Primary Visual Cortex: Scotomas and Completion
o Scotoma: Area of blindness in the visual field
Damage to an area of the visual cortex produces a scotoma in the area of the contralateral
visual field of both eyes.

Many patients with extensive scotomas are not even aware of their deficits due to
completion and other factors
A patient with a scotoma who looks at a complex figure will often report seeing the
complete figure
o Patients suspected of neurological damage to the primary visual cortex are usually given a
perimetry test.
o Perimetry Test: A patient's head is held motionless on a chin rest and the patient stares with
one eye at a fixation point on a screen. A small dot of light is then flashed on various parts of
the screen and the patient presses a button to record when the dot is seen. The entire process is
repeated for the other eye.
Result of a perimetry test is a map of the visual field of each eye, which indicates any areas
of blindness.
o Completion: The visual system's automatic use of information obtained from receptrs around
the blindspot, or scotoma, to create a perception of the missing portion of the retinal image.
completion sometimes depends on residual visual capacities in the scotoma.
Completion also occurs in cases which this explanation can be ruled out, such as patients
who are hemianopsic.
o Hemianopsic: Having scotoma covering half of the visual field.
Patients who are hemianopsic may see an entire face when they focus on a person's nose,
even when the side of the face in the scotoma has been covered by a blank card.
Damage to Primary Visual Cortex: Scotomas, Blindsight, and Conscious Awareness
o We generally assume that if a person sees something, he or she will be consciously aware or
seeing it.
o In reality, this assumption is not really true.
o Blindsight: The ability of patients to respond to visual stimuli in their scotomas even though
they have no conscciouus awareness of the stimuli.
Of all visual abilities, perception of motion is the most likely to survive damage to primary
vvvisual cortex.
Two neurological interpretations of blindsight have been proposed
The striate cortex is not completely destroyed and remaining islands of functional cells
are capable of mediating some visual abilities in the absense of conscious awareness
Those visual pathways that ascend directly to the secondary visual cortex frm
subcortical visual structures without passing through the primary visual cortex are
capable of maintaining some visual abilites in the absense of cognitive awareness.
Some support has been found for both theories, but nothing conclusive for either
It is possible that both mechanisms contribute to the phenomenon
Functional Areas of the Secondary and Association Visual Cortex
o Secondary cortex and portions of association cortex that are involved in visual analysis are both
composed of different areas, each specialized for a particular type of visual analysis.
o In the macaque monkey, more than 30 different functional areas of visual cortex have been
identified - 24 areas of the secondary cortex and 7 areas of the association cortex
o Various functional areas of secondary and association cortex are extensively interconnected.
Anterograde and retrograde studies have revealed over 300 connecting pathways
Connections between pathways generally move information both ways
Generally, however, the flow of information is from simpler cells to more complex ones

Various areas of the human secondary and association cortex have been identified through
the use of PET, fMRI and evoked potentials
Researchers have identified about a dozen functional areas of the human visual cortex.
Most are similar in terms of location, anatomical characteristics, and function to areas
already identified in the macaque.
Dorsal and Ventral Streams
o The information from the two lateral geniculate nuclei is received in the primary visual cortex,
combined, and then segregated into multiple pathways that project separately to the various
functional areas of secondary, and subsequently association visual cortex.
o Many pathways that conduct information from the primary visual cortex to other specialized
visual processing areas of the cortex follow one of two pathways
Dorsal Stream: Flows from the primary visual cortex to the dorsal prestriate cortex to the
posterior parietal cortex.
Most neurons in the dorsal stream respond most robustly to spatial stimuli, such as
stimuli that indicate where an object is spatially or indicate directionality of movement
Dorsal stream neurons are involved in the perception of "where" objects are.
Ventral Stream: Flows from the primary visual cortex to the ventral prestriate cortex to the
inferotemporal cortex.
Most neurons in the ventral stream respond most robustly to characteristics of objects
such as colour and shape.
Some evidence suggests that there are clusters of neurons in the ventral stream that
respond specifically to particular classes of objects such as faces, bodies, letters,
animals, tools, etc...
Ventral stream neurons are involved in the perception of "what" objects are.
o Where versus What theory:
Suggests that damage to some areas of cortex may abolish certain aspects of vision while
leaving others unaffected.
Most support for the where versus what theory comes from analysis of what occurs to
people's vision when they have selective damage to either the dorsal or ventral visual
stream.
Patients with damage to the posterior parietal cortex often have difficulty accurately
grasping an object they can easily describe orally.
o Control of Behaviour versus Conscious Behaviour Theory: Suggested that the difference
between the dorsal and ventral streams is not the kinds of information they carry, but the use to
which that information is put.
Suggested that the function of the dorsal stream is to direct behavioural interactions with
objects, whereas the function of the ventral stream is to mediate the conscious perception of
objects.
Suggested that conscious awareness medated by the ventral stream is what separates our
visual capacity from that of our closely related primate ancestors.
This theory also explains the observations of individuals with selective brain damage to a
single stream.
Major support from this theory comes from two related observations
Some patients with bilateral lesions to the dorsal stream can consciously see objects but
cannot interact with them.

Some patients with bilaterals lesions to the ventral stream have no conscious experience
of seeing and yet are able to interact with objects under visual guidance.
o The two streams do not function in isolation from one another, but little attention has been paid
to how they interact
Prosopagnosia: Visual agnosia for faces.
o Agnosia: Failure of recognition that is not attributable to a sensory deficit or to verbal or
intellectual impairment.
o Visual Agnosia: Specific agnosia for visual stimuli.
Visual agnosics can see visual stimuli but they don't know what they are.
Visual agnosias are often specific to a particular aspect of visual input and are named
accordingly.
Movement agnosia: Difficulty recognizing movement
Object agnosia: Difficulty recognizing objects
Colour agnosia: Difficulty recognizing colours
Assumed that each specific visual agnosia results from damage to an area of secondary
visual cortex that mediates the recognition of that particular attribute.
o Prosopagnosics can usually recognize a face as a face, but they have problems recognizing
whose face it is.
o Often report seeing a jumble of individual facial parts that are never fused, or bound, into and
easy-to-recognize whole
o In extreme cases, prosopagnosics cannot recognize themselves
o It was once believed that prosopagnosia applied only to faces because people were diagnosed
with prosopagnosia for being able to distinguish a pencil from a chair from a door, but not a
face from another face.
o This logic is flawed because the question of comparison should be whether or not the patient
can distinguish a chair from another chair.
o Evidence that one farmer lost the ability to distingish one cow from another.
o Evidence that some individuals experience no symptoms other than difficulty recognizing faces
in people.
o The diagnosis of prosopagnosia is often associated with damage to the ventral stream in the area
of the boundary between the occipital and temporal lobes. (Known as the fusiform face area)
Portions of this area are selectively activated by human faces
Other portions of the fusiform face area respond selectively to classes of visual stimuli other
than faces.
Similar face-specific areas have been found in the ventral streams of macaque monkeys
The extent to which the development of the fusiform face area depends on a person's early
experience with faces is still unclear.
o It makes sense that facial recognition would be designated such a specialized region of the brain
since human social behaviour patterns are highly dependent on facial cues.
o Patients who are shown photographs of individuals they know mixed in with photographs of
individuals they do not know will respond that they do not recognize any of them. Skin
conductance tests, however, demonstrate that the body reacts to photographs of familiar
individuals, even if the patient cannot consciously piece together the face.
o Patients recognize the faces in the photographs with undamaged portions of their brain.
Akinetopsia: A deficiency in the ability to see movement progress in a normal smooth fashion.
o Can be triggered by high doses of certain antidepressants

o Often associated with the middle temporal (MT) area of the cortex
The middle temporal (MT) is sometimes reverred to as V5 or MT/V5
Function of the middle temporal (MT) seems to be the perception of movement.
Some neurons at lower levels of the visual cortex respond to movement, but they provide
little information about the direction of movement becase their receptive fields are too
small.
95% of the neurons of middle temporal (MT) respond to specific directions of movement
and little else.
Each middle temporal (MT) neuron has a large binocular receptive field, allowing it to track
movement over a wide range.
o Four lines of evidence suggest that middle temporal (MT) is responsible for the perception of
motion and that damage to that area causes akinetopsia.
Patients with akinetopsia tend to have unilateral or bilateral damage to middle temporal
(MT)
As measured by fMRI, activity in middle temporal (MT) increases when humans view
movement.
Blocking activity in middle temporal (MT) with transcranail magnetic stimulation produces
motion blindness.
Electrical stimulation of middle temporal (MT) in human patients induces the visual
perception of motion.