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2/9/2014 11:06:00 AM

Three points of view on perception

Objectivist view: our senses precisely, and accurately, reflect the physical world.
They provide us with a true, complete, and accurate representation.

Championed by JJ Gibson (Cornell)

People believe it because of the good perceptual skills humans havevery

successful at interacting with the physical world

Subjectivist View: There is no inherent organization to the world, but rather, our
brains organize our perceptionstherefore we believe the world is, itself, organized.

Gestalt psychologists (Wertheimer, Wallach etc)

But how? If we are so good at detecting and discriminating physical

stimuli, then we must be perceiving exactly what is there

Our perceptions often do not simply record a stimulus


Illusory contours: unconscious inferenceyou cant consciously

control or turn off the illusion

Much of what we perceive is an illusion: part played by our senses

is at least as important in our perceptions of the world as the part
played by the world

Part played by our minds is at least as important as the events that

actually transpire

Synthetic view: The world appears to us the way it does because: we perceive only
within the limits of our nervous system but, our nervous system has evolved to
reflect certain aspects of the world very accurately.

Combines both subjectivist and objectivist viewsagrees with subjectivist

that our senses themselves play a big role in determining how we
perceive the worldbut also agrees with objectivist that our senses
evolved over millions of years under constraints imposed by the world
our senses have evolved so that we can function successfully.

The sensory systems of your brain must make some kind of inference
because you do not have the opportunity to perceive the world directly.

The neurons in your brain perform computations to process and

transform the incoming sensory stimuli to make these inferences.
Given the limited/indirect information available, the sensory
systems of your brain do remarkably wellmost of the time
unconscious inferences are correct

Occasionally, these unconscious inferences fail, giving rise to

perceptual illusions

Brain and Neurophysiology

Bold, unsubstantiated claim: the human mind (perceptions, emotions,
memories, thoughts) is an interesting and complicated machine. All behaviour is a
reflection of brain functioneverything you feel/think is no more than the electrical
and chemical activity of a vast assembly of nerve cells.

Descartes believed that the mind is separate from the brain

Descartes description of a reflex

Filaments (come from innermost part of the brain and compose the
marrow of the nerves) are arranged in every organ of sense so that they
can very easily be moved by the objects of that sense and that

When they are moved (doesnt matter by how much force) they
simultaneously pull the parts of the brain which they come fromopen
the entrances to certain pores in the internal surface of the brain

The animal spirits in its cavities begin to immediately make their way
through these pores into the nervesmuscles give rise to movements

Includes some correct observations (nerves are involved, there are fluid-filled
ventricles in the brain) but details are very inaccurate
Charles Sherrington replaced Decartes mechanical and hydraulic model of the
reflex with a model based on electrical signals in neurons
Nearly all psychiatric and many neurological disorders are characterized by
dysfunction in the neural systems that mediate these neural processes
Structure: dendrite, cell body/soma, axon (electric signal goes down axon starting
from cell body)

Dendrites: input from other neurons arrives here, combined in various

ways (input signals might be added, subtracted, divided by one another)

Axon: result of input computation propagated along axon to serve as

inputs to other neurons, some axons very short (local connections
between neurons in one part of the brain) and some very long (>1 metre
down the spinal cord to innervate a muscle)

Soma: cell body between dendrites and axon

Golgi stain: stains a random handful of neurons so you can see them distinctly in
the midst of all the other cells packed in around them
Pyramidal cell: one of many different types of neurons found in the cortex
Axons make both local and far away connections

Action potentials
Neurons represent and transmit information electrically. If you place an electrode
close to a neurons soma then you can measure and record the signals.
Action potentials are often referred to as spikes or impulses

When not stimulated, typical neuron rests at a voltage of about -70mV

When stimulated, fires an action potential, voltage increases to +40mV

then comes back down again

Action potential starts at a neurons soma and travels down the axon

Neurons use firing rates to transmit information

You can measure a neurons response in terms of firing rate (number of action
potentials that occur per unit of time)

E.g. the response of a retinal ganglion cell depends on the contrast of the
test lightdim test light has fewer action potentials than bright test

When you increase brightness, action potentials dont get bigger, just
become more frequent

An analogous result is observed when you measure the firing rates of touchsensitive nerves and vary the strength of a stimulus that presses on the skin
Neurons respond selectively
When you excite the visual receptors in any way, the response in the nervous
system is that of a visual response, if you excite an auditory neuron, the resulting
sensation will be one of hearingknown as the Law of specific nerve energies

Formulated by Johannes Muller

Neurons in the different parts of the brain are selective for different

There are even neurons that respond selectively to faces

Neural responses are noisy

Recordings of action potentials are noisy, if you present the same stimulus
repeatedly, each time you hear a slightly different responseaction potentials
happen at different times on each trial, total number of action potentials varies
from one trial to the next.
The axon of one neuron comes very close but doesnt touch the dendrite of its
The narrow space between the two neurons is called the synapse
Electrical signal from the pre-synaptic neurons axon is not transmitted directly
across the synapse, when the action potential comes along, it causes the release of

certain chemicals called neurotransmittersthese diffuse across the synapse and

blind receptors in the membrane of the post-synaptic neurons dendritecauses a
change in the electrical properties of the postsynaptic neuron.
A given neuron may have thousands of
synapses distributed on its dendritic tree
if there is enough combined excitation from
all these synapses then the postsynaptic
neuron will fire an action potential
if there is even more excitation, it will fire
several action potentials (firing rate
increases with dendritic excitation)
Anatomy of the brain
Cerebral cortex is the outer layercovers the rest of the brain
Lobes: frontal, parietal, occipital, temporal
Sulcus: infolding
Gyrus: outfolding
Grey matter: folded sheet containing cell bodies, dendrites, local axon collaterals
White matter: axons, long range connections
Different brain areas have differences in layeringmotor cortex (involved in
controlling muscle movements) has an expanded layer 5 and a reduced layer 4.

Layer 5 is the output layer where neurons send their axons down the
spinal cord, layer 4 is the input layer.

This is the motor control area so it makes sense to have lots of output
and not much input

Visual cortex has expanded layer 4 because it has a lot of inputs

Neurons perform computations

A neurons response might depend on the sum of the responses from two of its
The function of the neurons in a particular brain area is best understood in terms of
the computations they perform
Human brain physiology
Neuropsychology: study patients with brain damage to specific brain areas

e.g. blind sight (residual ability to discriminate visual stimuli when forced
to guess, even though the subject has no conscious experience of the

e.g. face agnosia (inability to recognize faces)

Electrical microstimulation: sometimes during surgery for epilepsy patients, they

keep the patient away so that they can electrically stimulate the brain and make
sure they dont accidentally remove critical brain centres.

In the process of coming up with this method, Penfield localized specific

regions of the brain involved in vision, hearing, touch and motor control

By plotting movements and sensations that are evoked by electrical

stimulation along the central sulcus, he revealed a homunculus, a
representation of the body surface

Transcranial magnetic stimulation (TMS): is the use of powerful rapidly changing

magnetic fields to induce electric fields in the brain by electromagnetic induction
without the need for surgery or external electrodes
One reason TMS is important is that it can demonstrate causalityyou can tell that
the region of the brain that is activated is due to that task
Functional magnetic resonance imaging (fMRI):

Non-invasive method for measuring activity in the human brain, and for
investigating the relationship between brain activity and behaviour

Technique is similar to conventional MRI but can generate images of brain

activity in addition to brain anatomy

fMRI works by measuring blood flow, taking advantage of the coupling between
neuronal activity and hemodynamic (local control of blood flow and oxygenation) in
the brain to allow the non-invasive localization and measurement of brain activity.

Brain controls the flow of oxygenated blood to where it is needed

Iron in the blood is magneticoxygenated and deoxygenated blood have

different magnetic properties

MRI (1970s) based on the physics of magnetic resonance

MRI scanner can be reprogrammed to pick up differences in

magnetization that take place when the brain ships oxygenated blood to
where it is needed.

Ultimate success of fMRI as a measurement of brain function depends on the

relationship between fMRI signal and the underlying neuronal activity.
Vascular source of the fMRI signal places important limits on the usefulness of the
techniquewe know that fMRI signal is triggered by metabolic demands of
increased neural activity, but the details of the process arent fully understood
fMRI is an indirect measure of the underlying neural activity

but there are numerous demonstrations that fMRI measurement is tightly

linked with underlying neural firing rates

as contrast increases, firing rate increases

Cerebral control of blood flow

Pulsations in the brain increased in magnitude during periods of mental


When the brain is active, an oversupply of oxygenated blood is

deliveredthis has the effect of making the fMRI images slightly brighter
in the vicinity of active neurons

Positron Emission Tomography (PET)

A patient lies in the PET scanner after being injected with radioactive-labelled
substance (water/glucose)

More blood goes to the active brain regionsmore radioactivity in those

regions, PET scanner detects and localizes them

Scanner detects the unique radioactive decay of positrons (positively

charged particles) attracted to electrons in the blood, when they come
together theyre annihilated and energy is released in the form of two
photons (light)scanner detects where photons produced

Test condition: flickering visual stimulus

Control condition: blank screen
Often, the studies average results across many subjects which can be a problem
because individual brains can be different in shape
Ideally you would want to test multiple times on one person but too much exposure
to radioactivity
Electroencephalogram (EEG) and the event-related potential (ERP)

Also used to study brain function

At each electrode, the electrical activity is recorded at fixed intervals

following a stimulus presentation

Electrical values at each interval are taken from many trials and averaged
together so that electrical activity not caused by stimulus averages to 0
shows only activity produced by stimulus

Result is called an ERPusually 10-100 presentations of the stimulus

suffice to produce reliable measurement that reflects characteristics of
individual brain and particular stimulus

Magnentoencephalogram (MEG)

Measures tiny magnetic fields evoked by electrical currents in neurons (generated

when electrical currents travel along lots of nearby dendrites at the same time
Comparing fMRI, ERP, MEG and PET
fMRI and PET depend on regional control of blood flow in the brain but:

fMRI methods go way beyond conventional PET subtraction methodology.

fMRI is non-invasiveno radioactive stuff neededcan do repeated fMRI

on same subjectcan improve experiment and do it again on the same

spatial and temporal resolution of fMRI and PET are limited by blood
flowfMRI responses depend on average activity of neurons in tiny part
of the brain, averaged over time.

Spatial resolution with fMRI is better than PETbecause of the

basic differences in technology and because can be repeated on the
same person (PET patients have different shaped brains)

EEG>fMRI because of temporal resolution

Huge disadvantage for EEG is localizationdont know where electrical signals are
coming from
Strength of EEG signal at each electrode depends on how far it is from sourcecan
infer roughly where signal is coming from, if there are large number of sources, no
way to tell
MEG (like EEG) has advantage of millisecond time resolution, but limited spatial
resolution and localization

Study of the ethical, legal and social questions arising when scientific findings about
the brain are carried into medical practice, legal interpretations, and health and
social policy

Applications to lie detectionusing fMRI, 88% accuracy

The anterior cingulate cortex has often been reported to be active when people lie,
but no causal relationship between activity here and lie detectioncant know for
Limits of neurophysiology: neuroscience methods (recording firing rates,
neuroimaging etc.) are useful but cant answer things about sensation and
Behavioural experiments: goal is to interpret brain activity in terms of
perception and guided behaviour
Starting point is to come up with a phenomenon (like an illusion of observationwe
can see where sound is coming from based on combining sounds from both ears)
Many of the basic phenomena in vision and hearing were discovered and described
by Helmholtz
Want to establish casual link demonstrating that a particular neural mechanism
causes a particular perceptual phenomenonneed to have quantitative
measurements and need to show they are consistent to establish strong link
Three basic experimental protocols that we use in perceptual psych experiments:
magnitude estimation, matching, and detection/discrimination
Magnitude estimation: subject rates some aspect of a stimulus (how bright e.g.)

Relationship between intensity of a stimulus and perception of magnitude

follows the same general equation in all senses: P=kSn

P= perceived magnitude

S= stimulus intensity

K= constant

Matching: in matching experiments, subject has to adjust one of two stimuli so

that they are the same as the other

Also sometimes done by showing a series of trials and asking subject to

pick closest match

Detection/discrimination: subjects task is to detect small differences in stimuli

Psychophysical procedures for detection/discrimination: Signal

detection theory informs us about the general principles regarding
sensitivity measurement but how do we do a detection experiment?

Method of adjustment: ask subject to adjust light intensity until

they think its barely detectablethis method is subjectivedifficult
for them to judge their own threshold, can be stressful

Yes-No/method of constant stimuli: observer presented with

series of tones of various intensitiesobserver asked to report
whether or not they heard it, calculate probability of detection (%
yes responses) and can be potted against tone intensity

These curves are called psychometric functions: plot signal

strength against %age yes (50% usually used as estimate of

Whats wrong with this? No noise-only trialsno false

alarmscant use data from yes-no to estimate sensitivity
separately from criterion (d)

Forced choice: most popular choice, signal presented sometimes,

sometimes not presentedsubject forced to say whether it was or
wasnt presentedcan count both hits and false alarms to get
estimate of d

Comparing methods: the different procedures lead to different

estimates of thresholdeven for experienced observers, adjustment
thresholds and forced-choice thresholds are usually different by about a
factor of 3

Forced choice and signal detection theory:

SDT used to analyse and think about detection and discrimination experiments
To summarize SDT:

Your ability to perform a detection/discrimination task is limited by

internal noise

Information (signal strength) and criterion are the 2 components that

affect your decisionthey both have different kinds of effects

Because there are 2 components, we need to make 2 measurements to

characterize the difficulty of the taskby measuring both hits and false
alarms, can estimate d (measure of task difficulty independent of

d increases as hit/false alarms increasewhen the light intensity is

increased, discrimination is easier, hit rate goes up, false alarm rate goes
down, d goes up

Choosing the threshold: Usually just pick a particular value of dusually 1 (75%
correct) so the intensity that yields d=1 is called the threshold
Discrimination: dont have to measure a threshold of signal against no signal, can
also find threshold when discriminating between two different levels of signals
(difference threshold) (signal v no signal is absolute threshold)
Webers Law: measuring difference thresholdhave been performed in many
different sensory modalitiesto be able to discriminate between light intensity,
sound, pressure, weight etc. Result is always the same

The difference threshold is proportional to the baseline/starting intensity

Fechners Interpretation: came up with the idea of internal response,

hypothesized that when two signals are just noticeably different, it is as though
they are separated by one unit of internal response

the threshold from the zero background

to the first discriminable intensity level
defines one unit of internal response
curve is logarithmic

Fechner didnt measure internal

response directlyinferred that the
internal response increases with the log
of stimulus intensity [s(x)=log(x)]

Can make predictions with thisthere

ought to be neurons that response in
proportion to the log of stimulus intensity

Weber-Fechner Summary
Webers Law: to perceive a difference between a background level x and the
background plus some stimulation x+dx, the size of the difference must be
proportional to the background dx=kx
Fechners Interpretation: the relationship between stimulation level and the
perceived sensation is s(x)= log(x)
Nowadays, people dont believe Fechners interpretationdata is the same but
interpretation changedWebers law can also be explained by assuming that
internal response is proportional to contrast (ratio of intensity at a point divided by
local average intensity)
Sound and The Ear

Acoustic stimuli are communicated through changes of air pressure

The tendency of the high pressure region to invade the region surrounding it means
that it will have to invade increasingly large regions of space
When you clap, air molecules at your ear are undisturbed but change in air
pressure is rushing outwards in all directions so when it reaches your each there is
a momentary change in the air pressure in your ear

This brief disturbance is the only information available to you about the
hand clap

Complex sound signal

Real sound signals can involve very complex changes in sound pressure level over
Pure tones: what you get from a tuning fork

Pure tone is characterized by frequency and amplitude

Frequency (Hz)= rate of air pressure modulation (pitch)


Hznumber of cycles of sound pressure change per second

Amplitude (dB)= sound pressure level (volume)


Decibels (dB) are a log scale

Hearing sensitivity to different frequencies: for very low/very high frequencies, pure
tone must have a very high amplitude to be heard

20-20,000 Hz is a rough range of frequencies to which humans are


More sensitive to 1000-5000 Hz

Frequency decomposition
Sound signals are often characterized in two
different ways, left graph shows sound pressure
level (SPL) against time, right graph shows Fourier
spectra of the soundsSPL against frequency
Any sound signal can be decomposed as a sum of
pure tones or frequency components

Fourier spectrum graph plots the

intensities (SPL) of the separate pure
tones than make up the entire sound

Controls of a stereo are designed around Fourier frequency

decompositionincreasing bass increases low frequency signals,

equalizers divide up the signal into many frequencies to give you finer
Swinging pendulum
When you swing the pendulum back and forth slowlylong string swingslow
frequency and vice versa
Fourier spectrum is essentially a plot showing the amplitudes of swing for each of
the pendula
Real sounds that are produced naturally typically have a fundamental frequency
and harmonics (multiples of the fundamental)
The Ear
Pinnalarge bit of skin on the side of
your head that focuses sound ways
into your auditory canal
Tympanic membrane (ear drum) is a
cone-shaped membrane
Sound waves propagates through the
auditory canal and introduces a
differential between the air pressure
levels on the two sides of the
tympanic membrane

Differential causes ear drum

to moveinternal pressure
on one side of the tympanic membrane acts as a baseline estimate of the
general, atmospheric pressure.

We must re-adjust this level occasionally when we are subjected to

changes in the ambient air pressure

Displacing the tympanic membrane causes tiny

bones (ossicular chain) to move.
The malleus/hammer is attached to the tympanic
membrane. The incus/anvil acts as fulcrum. The
stapes/stirrup is attached to the inner ear in the
oval window.
The ossicular chain acts as an amplifierthe
pressure per unit area applied by the stapedial
footplate to the oval window is about 14 times as
great as the pressure per unit area applied to the ear drum by the external changes

in air pressurethis increase is important because the opposite side of the oval
window is filled with liquid that is harder to compress or move than air.
Amplification is accomplished by two factors:

The area of the tympanic membrane is about 10x greater than that of the
oval window

The fulcrum-like action of the ossicular chain.

Ossicular chain is suspended in the middle ear
by several tendonswhen tympanic membrane
moves, the bones pivot about the anvil
because of the position of the pivot point, a
small motion of the membrane leads to a larger
motion of the oval window

Cochlea (inner ear)

Oval window: see above
Round window: another elastic membrane, whose
role is simply to dissipate any excess
energy/pressure in the cochlear fluid
Semi-circular canals: primary organs of your
vestibular system that is responsible
the cochlea is divided into three regions: scala
vestibule, scala media, scala tympani
When the ossicular chain presses against the oval
window, it gives the fluid in the cochlea a good push.
This causes a local change in fluid density which
propagates through the scala vestibuli, around the
outside of the scala media to the end of the cochlea.
At the end of the scala vestibuli, there is a small
opening that connects up the two outer scalacalled
the helicotrema.

The energy in the fluid is largely dissipated by the time it reaches the

Whatever's left is dissipated by elasticity

of the round window, the crucial region is
the scala media, that moves up and down
as the fluid pressure waves propagate by.

The arches of corti are a complicated set of structures sitting on the basilar
membrane. The main parts were interested in are:

Inner and outer hair cells: transduce/convert mechanical motion of basilar

membrane into neural signalsmain cell bodies span distance from
basilar membrane to the tectorial
membrane, with cilia on top

Tectorial membrane sits just above

stereocilia of the hair cells, forming a

When a sound pressure waveform arrives at
your ear, air pushes the ear drum, which in turn
pushes the ossicular chain, which in turn
presses the oval window.
That introduces pressure difference between the fluid in the scala vestibuli and the
scala tympani with the elastic basilar in between. The pressure difference pushes
the basilar membrane down. When the air pressure subsides, the tympanic
membrane moves out pulling th`e oval window with it. Now the pressure difference
is in the opposite direction and the basilar membrane moves up.
Motion of the arches
The arches are connected on one side to the
edge of the basilar membrane, and on the
other side they are connected to the centre
of the basilar membrane.
Side connected to the centre moves up and
down with the basilar membrane, but the
other side doesnt movearch tilts over a

As the arch tilts over, the hair cells

are bent back and forth

This is the key motion that converts the mechanical motions into an
electrical signal used by the rest of the nervous system.

Conversion from mechanical (motion of basilar membrane) energy into neural

(electrical) response in hair cells is called transduction
As the stereocilia are bent, positively charged potassium ions flow into the neuron,
this is the electrical signal that initiates neural conduction.

This voltage is graded, an analog signal

Hair cells dont fire action potentials

The hair cells themselves are attached to a bundle of nerve fibres, if the voltage
signal in the hair cell is large enough, it causes an auditory nerve to fire an action
Hudspeth managed to stimulate hair cells directly using a micromanipulator
simultaneously displaced stereocilia and measured the electrical voltage in the hair
cell; measured size of the change in electrical signal as a function of the size of
displacementover a large fraction of its range, response of the hair cell is linear
double displacement=double the response
The direction of bending also makes a differencesystem designed so that under
normal circumstances, sound signal will cause stereocilia to bend in a specific

Hudspeth measured what happens when its bent in another direction

bending in normal direction excites neurons the most

Even though there are more outer than inner hair cells, most of the ascending
nerves fro the 8th cranial nerve get input from inner hair cellsresponsible for
capturing what we hear and communicating to our brain.

Outer hair cells change shape and rigidity to control motion of the basilar
membraneamplifying weak sounds and attenuating loud sounds

Linear Systems
Systems, Inputs, and Responses
One possible way to characterize the response of the ear to sound might be to build
a look-up table: a table that shows the exact neural response for every possible
auditory stimuluswould take forever
Linear Systems
To see whether a system is linear, need to test whether they satisfy:

Homogeneity: Sometimes called scalar rule. If you double input, output

should double.

Additivity: S1+S2=R1+R2

Superposition: systems that satisfy both homogeneity and additivity are

considered linear systems. These two systems together, often referred to
as the principle of superposition

Shift-invariance: If you present ear with S1 and it creates a response, if

you present it with S1 again but at a later time, response will be the same
but shifted in time Not all linear systems are shift-invariance

Why impulses are special: homogeneity, additivity, and shift invariance suggest
that the systems response to an impulse can be the key measurement to take.
Trick is to conceive of the complex stimuli we encounter as a combination of
We can approximate any complex stimulus as if it were simply the sum of a number
of impulses that are scaled copies of one another and shifted in time
For shift-invariant linear systems, can measure the systems response to an
impulse and well know how to predict the response to any stimulus (combinations
of impulses) through the principle of superposition.
To characterize shift-invariant linear systems, we only need to measure one thing:

The way the system responds to an impulse of a particular intensity

impulse response function of a system

Once weve measured this function, we can predict how the system will respond to
any other possible stimulus
Sinusoidal Stimuli
Have a special relationship to shift-invariant linear systems
Distance from one peak to the next is called the wavelength/period of the
sinusoidgenerally indicated by lambda
Inverse of the wavelength=frequency: number of peaks in the stimulus that arrive
per second at the ear

Longer the wavelength, lower the frequency

Mathematical expression of the sine wave= Asin(2pift)

The response of shift-invariant systems to sine waves:
Can express any periodic stimulus as the sum of a series of sinusoids at different

Called Fourier Series expansion of the stimulus

s(t) = A0 + A1 sin(2 f1 t + p1) + A2 sin(2 f2 t + p2) + A3 sin(2 f3 t + p3)

+ ...

Even the abrupt onsets and offsets of a sound can be decomposed as a sum of
smoothly modulating pure tones
Decomposition is important because if we know the response of the system to
sinusoids at many different frequencies, we can use the same thing we did for
impulses to predict the response via impulse response function

First, measure systems response to sinusoids of all different frequencies

Then, take our input stimulus and use the Fourier Transform to compute
the values of the coefficients in the Fourier Series expansion

Has been broken down as the sum of its component sinusoids

Can predict the systems response by adding the responses for all the
component sinusoids

Why use sinusoids?

We use sinusoids as input to a shift-invariant system because the

systems responses is always a copy of the input at the same frequency
as the input

Measuring the response to a sinusoid for shift-invariant entails measuring

only two numbers: shift and the scale

Representation of how the shift-invariant

linear system behaves (plot of values of the
shift and scale for each possible input
frequency) is called modulation transfer
functionsame as providing you with
impulse response function

Can use these numbers to compute

response to any stimulus

Frequency Tuning and Pitch Perception

Frequency Tuning
Found that basilar membrane acts as a shift invariant linear system, then used
sinusoidal stimuli to measure the frequency response at different points along
basilar membraneable to predict response (motion of membrane) for any sound
Cochlea varies in thickness and elasticity as it curls from the oval window out to
helicotremaeffect is that different parts of the basilar membrane respond more
strongly to some sounds than others
For pure tones, each point on the basilar membrane oscillates up and down at the
same frequency as the soundonly size of oscillation differs
Entire motion that occurs on the basilar membrane in response to a sound stimulus
is called a traveling wave
Wave begins at the oval window, rises to a crescendo somewhere along the basilar
membrane, and falls off with the energy being absorbed around the helicotrema
Envelope for several frequencies

Each point along the basilar membrane oscillates a different amount, depending on
the frequency of the sound.
Points near oval window oscillate largest amount in response to high frequency
Points near helicotrema oscillate largest amount in response to low frequency tones

Happens because the stimulus begins with a push at the oval window
forces the part of the cochlea nearest to the oval window to begin
oscillating, takes time to propagate down the cochlea

The motion of the basilar membrane in response to a complex sound is the sum of
the responses to the pure tone components of that sound
Each auditory nerve fibre is connected to a small number of hair cells, near one
another, on the basilar membranenerve fibres response is governed by motion of
a small region on basilar membrane

Basilar membrane in any small region undergoes its largest motion only
for a small range of frequencies

Most sensitive frequency for an auditory nerve fibre is called the neurons
characteristic frequency
Representation of information on basilar membrane and 8th nerve is very different
from representation at tympanic membrane
Place and Temporal Code Theories of Pitch Perception
Pitch is a perceptual attribute, not a property of physical stimulus
Place Code Theory: Helmholtzs theory of pitch is based on observations of the
anatomy of the ear, sensation of a low frequency pitch derives exclusively from the
motion of a particular group of hair cells, and same for high pitch.
Each sensation is perfectly identified with an action of an anatomical location along
the basilar membrane.
Place code theory given its name because it identifies each pitch with a particular
place along the basilar membrane
Temporal Code Theory: The location of activity along the basilar membrane is
irrelevant, pitch is coded by the firing rates of nerve cells in the auditory nerve
Low frequency tone causes slow waves of motion in basilar membrane and that
might give rise to low firing rates in auditory nerve, high frequency causes high
firing rates
There is a problem with temporal codeear is sensitive to frequencies up to
20,000Hz and nerve cell cannot fire that fastdoesnt make sense
Cochlear Microphonic: places doubt on place code, supports temporal code.

Is a small electrical signal that can be measured by an electrode placed

near the hair cells of the cochlea

We know that the cochlear microphonic arises from the sum of electrical potentials
in the hair cells of the cochleamimics the form of the sound pressure waves that
arrive at the ear
Low frequency tones result in low frequency modulations of the cochlear
microphonic electrical signal
Cochlear microphonic is a shift-invariant linear system that obeys the scalar,
additivity, and shift-invariant rules
Volley principle: reconciles the fact that the cochlear microphonic mimics the
sound pressure waves with the implausibility of the temporal code.

Suggested that while one neuron alone cant fire for 20,000Hz tone, 20
neurons together cancollectively can fire together

Phase Locking: empirical observation that supports volley principlewhen

auditory nerve neurons fire action potentials, tend to respond at times
corresponding to a peak, result is that there are a bunch of neuron firings near the
peak of every cycle of a pure toneone neuron cant fire at every peak but different
neurons can fire successively

What you need for temporal code theory is for neural activity to look like
the sound pressure waveformresponse must follow rise and fall of sound
pressure level

Wevers temporal code theory rejected Place Code Theory and was backed up.
Whites Cochlear Implants:
He tested place code theory by measuring the dependence of pitch on which
electrode was being stimulated
He tested temporal code theory by measuring the dependence of pitch on the
frequency of electrical stimulation

Results: both mechanisms play a role in pitch perception

Virtual Pitch:
Both the place theory and the temporal code/volley theory play roles in pitch
perception, however neither theory provides a complete explanation for pitch
Loudness Perception and Critical

Like pitch, its perceptual

Equal Loudness Curves
Two sounds that have the same amplitude but different frequency are often
perceived to have different volumes
Firing Rate Hypothesis: rate of firing in the auditory nerve might determine
perceived loudness

For weak tones, basilar membrane displaced little, hair cells not pushed
very farfew spikes in the auditory nerve fibres

The rising phase of each cycle of the sound signal evokes bursts of spikes
in a collection of auditory nerve fibresamplitude of the sound
determines the number of spikes per burst

If this was true, we could just control perceived loudness by injecting patterns of
current into the neuron that causes it to respond at a more rapid firing rate
Number of Neurons hypothesis:
Most neurons fire to a louder sound, as a traveling wave passes down the basilar
membrane, each point of the membrane oscillates at the frequency of the tone
When the sound is weak, displacements are generally small and only a small region
of the membrane moves sufficiently to evoke any responses
In the auditory nerve, this means that additional nearby auditory nerve fibres will
be recruited when the sound intensity is increased
The physics, physiology, and anatomy do not define the perceptual code:
Both the firing rate hypothesis and the number of neurons hypothesis could
plausibly serve as the mechanism for our perception of loudness
Experiments show that both play a role in perception of loudness
Critical Bands
Band-limited noise: any sound can be decomposed as a sum of pure tone
sinusoids, can represent pure tone components of a sound by drawing a graph of its
Fourier spectrum
Band-limited noise stimuli have equal energy at all frequencies within some region,
and no energy outside of that region
Can describe this stimulus with three values:

Centre frequency: frequency marking centre of the region where there is


Bandwidth: width (in Hz) of the region of frequencies where there is


Total energy: summed energy of all pure tone components, which is the
area under the Fourier spectrum curve

Band-limited noise on the basilar membrane: if we change only centre

frequency, shift along the basilar membrane
If we change only bandwidth, change how much of the basilar membrane we are
Allows us to test the number of neurons hypothesisif we change total energy,
change the amount of displacement at each excited positiontest firing rate
Zwickers loudness matching experiment:
Showed that there is a range of bandwidths for which the perceived loudnesses are
There is a range of bandwidths for which the perceived loudness increases with the
increase in the number of neurons
Critical bands:
Zwicker determined that there was a region over which the cochlea adds up the
energy that it is receivingwithin this critical region sounds of equal total energy
have equal loudness
As soon as the sounds that we present are spread over a large frequency range,
sound with larger bandwidth sounds larger
Critical band corresponds to a pooling along the basilar membrane: width in terms
of frequency corresponds to an estimate of the physical length along the membrane
over which the auditory nerve signals are pooled
Summary: a neural code for both pitch and
Pitch depends on both a place code (which
position exhibits the larger firing rate) and a
temporal code (firing in bursts that phase lock
to the stimulus frequency)
Loudness depends on firing rates (more spikes
per burst for louder sounds) and number of
neurons firing (at high intensity, you get some
spikes from both positions)
Auditory Pathways and Sound Localization
Auditory Pathways

Primary auditory pathway begins with auditory receptors in the cochlea

These synapse on spiking neurons in the spiral ganglia, the axons of which from the
auditory nerve
These lead to the cochlear nucleus then to the superior olive then inferior colliculus
then medial geniculate nucleus then auditory cortex
Crossing of the Fibres: significant number of nerve fibres cross the brain and
make connections with neurons on the side opposite from where they started
happens very early on in the auditory system
Inter-aural comparisons are an important source of information for the auditory
systemwhere a sound came from
Tonotpic organization: spatial layout of frequencies in the cochlea along the
basilar membrane is repeated in other auditory areas in the bran

Cochlear nucleus is also organized tonotopically

Analogous tonotopic organization is also found in the auditory cortex

Auditory Scene Analysis

How does the brain separate sounds that are overlapping/intertwined?

Called the problem of auditory scene analysis

A lot of things contribute to perceptual organization (grouping/segregating)

Dont always notice echoes
Your brain suppresses the naturally occurring echoes
Play the sound of a hammer backwards and it sounds completely differentthe
natural echoes in the environment are suppressed by your auditory system and
never reach your consciousness
When you clap your handsthe stimulus that reaches your ears is a combination of
the sound wave that comes directly from your hands plus the sound waves that
bounce around the room for a while before reaching your ears
We dont hear a large portion of the physical stimulus coming to our ears
Our auditory system makes an unconscious inference from the sound wave to the
physical event that caused it
Spatial Localization
Sounds can be localized using three coordinates:

Azimuth: angle left or right of straight ahead

Elevation: angle above or below horizontal plane


Two cues for localizing a sound source are:

Inter-aural intensity difference (IID)

Inter-aural timing difference (ITD)

When you hear something from a cornersound arriving at the ear further away is
delayed (time difference) and lower in amplitude (intensity difference)
Intensity difference: sound intensity decreases by the time it reaches the further
ear (by 1/d^2)
The head interferes with the sound-wave, casting the auditory equivalent of a
shadow on the far ear

Sound shadow is more effective for sounds with higher frequency

IID depends on both azimuth of the sound source and the frequency of the sound
Timing difference: this is easy to see with abrupt soundscompare onset of the
sounds in each ear
For continuous pure tones, more difficult as it involves a phase difference between
the sound in two ears

This phase difference is ambiguous, can only be clearly resolved for lower

Abrupt onset sounds and low frequency sounds give rise to ITD
For normal person, sound delay is very short
Measurements of timing differences
Two cues work togetherpeople can adjust the intensity of one tone and
compensate for the time delay so that they hear a single tone in the middle of their
headboth cues are combined to localize sound sources
The intensity difference needed to accomplish this is quite largevery small
differences in time between the two ears requires large differences in intensity to
compensate for the perceived displacement of sound
Neurophysiological mechanisms of sound localization
Studied the inputs to the medial superior olive (MSO) and lateral superior olive
(LSO)they discovered neural circuits and neurons that appear to be well-suited to
representing these two types of information, both timing and intensity differences
LSO neurons measure intensity differencescontains neurons that are selective to
intensity differences
MSO neurons measure timing differences

If sound is coming from right, firing rate in right cochlear nucleus is

higher than leftLSO neuron computes difference between firing rates
and responds strongly


Responds strongly when the sound in right ear leads left ear by some
short time

Many MSO neurons work together to infer and represent the timing


MSO neurons act as coincidence detectorscoincident spikes

arriving from two ears evoke response in MSO neurontiming of
individual spikes must be very precise at least for low frequencies
phase locking

Anatomy of MSO:

The neurons are connected in a way that looks like Jeffries wiring

inputs from the two ears are combined to generate neuons whose
preference fro inter-aural time differences varies systematically

Cone of confusion
Sound sources located at any position in the cone generate exactly the same IID
and ITD cues for the listener and thus cannot be distinguished using IIDs and ITDs
Two ways to distinguish direction

Azimuth and elevation

You can rotate your head so that one ear is directly in front of the source

The outer ears are asymmetricfilters sound differently depending on

where the source is and what frequency

if we measure the intensity of sounds at the eat drum as a function

of their azimuth, elevation and frequency, result is called Headrelated transfer functiondescribes IID as a function of frequency
by attenuation characteristics and ITD as function of frequency in
phase delay

Cues to distance
Intensity can be a cue to sound source distancebecomes less intense as it reaches
your ear
But we also have some degree of loudness constancy (sounds the same volume
even though intensity decreases)
You normally dont perceive echoes as separate sounds, but the timing of these
reflections and the number and intensity of them is a cue to the kind of space you
are in and the distance to the sound source

Cochlear Implants and Speech Perception

Speech analysis
Phonemes: the smallest unit that, if changed, can change the meaning of the word
Phonemes correspond to produced speech sounds
Vowels: voiced
Consonants: constrict/stop air flow
Graph of frequency v time
Higher amplitudethicker line
Higher frequencyhigher y coordinate
Computed using Fourier Transform
Vowels/consonants that are voiced are visible in spectrograms as a series of thin,
horizontal bands corresponding to the separate harmonics

These peaks are called formants

Difference between ba da and ga is in the formant transition

Length of the formant transition and the time at which voicing begins
following the stop are indications of whether the stop consonant is voiced
or unvoiced

Cues in Spectrograms
Prosody: extra-verbal indication shown by change in pitch from high to low or low
to high shows whether it is a statement or question etc.
Auditory Cortex, Wernickes Area and Brocas Area
Primary auditory cortex is located laterally near top of temporal lobe
Auditory cortex critical for speech perception and language comprehension

Aphasia: collective deficits in language comprehension and production

that accompany brain damage

Damage to Wernickes area causes disorder of language comprehension

Damage to Brocas area causes disorder of speech production

Language learning impairment:

Kids have difficulty with speech because they have deficits in the fast temporal
processing needed to distinguish brief formant transitions
Cochlear Implants
Several electrodes mounted on a carefully designed support that is matched to the
shape of the cochlea

Design is important because it places electrodes very close to nerve cells

Computer decomposes signal into frequency components via Fourier

Transform and sends the components to the corresponding electrodes

2/9/2014 11:06:00 AM
The Eye and Image Formation
The Eye
Vision begins when light enters the eye, focused by the cornea and lens onto the
retina, a thin layer of neural tissue at the back of the eye.
Photoreceptors, specialized neurons that transduce light into neural signals,
respond with graded potentials, pass the signal on to bipolar cells and then to
retinal ganglion cells.
Retinal ganglion cells are the only neurons in the retina that fire action potentials.
Axons of ganglion cells make up the optic nerve
Blind Spot:

Photoreceptors are at the back of the retina, ganglion cells are at the
front, and ganglion cell axons make up the optic nerve that goes through
a hole at the back

Cant be any photoreceptors where the hole is

Photoreceptors transform light energy into a neural signalno

photoreceptorsno light

Eye Anatomy, Cornea, Glaucoma, and Corneal Transplants

Sclera: white part

Pupil: hole

Iris: Coloured part, has muscles that change its shapechanging pupil

Cornea: thin, transparent covering of the eyeball, serves as chief

refracting/focusing element of the eye, about 2/3 of the optical power of
your eye is in the cornea and 1/3 in the lens

Lens: adjustable focus for near/far

Fovea: part of the retina corresponding to the central part of the visual
fieldpit shaped because the bipolar and ganglion cells are pushed off to
the side so that the photoreceptors have better access to incoming light

Optic disc: part of the retina corresponding to the blindspot

Optic nerve: made up of ganglion cell axons that exit through the optic

Anterior chamber/aqueous humor: fluid filled region in front of the lens

Posterior chamber/vitreous humor: fluid filled region behind the lens

Cornea is living tissue but must be transparentno blood vessels running through it

Has two practical consequences:

Glaucoma: without blood vessels, nutrients are passed to the cornea and
lens through a continual flow of liquids through the eye

Glaucoma is a disease caused by a blockage of the flow

Normally, nutrients flow around the lens to the anterior chamber of

the eye then out through a pore called Shlemms canal.

Flow can either be blocked at the pupil (fixed by making a hole in

the iris) or blocked at Shlemms canalfixed with drugs

End result of glaucoma is damage at the back of the eye, leading to

blindnessincrease in the pressure of the eye causing blood
vessels in the eye to compress, sometimes destroying them. Can
also crush optic nerve.

Loss of blood vessels can lead to retinal detachments (can also be

caused by eye injury). Can be fixed by surgery if caught
immediately by laser.

Glaucoma can be diagnosed is through tonometry (using Shiotz

Tenometer). Eye was anaesthesized and weights were used to
displace the eyeballamount of weight was measured which
showed something about pressure in eye. Nowadays just blow at
your eye

Corneal Transplants: because there is no blood, cornea can be

transplanted with very little fear of rejectionno antibodies

New research in stem cell technology enabled procedures for

repairing corneas for people who rejected cornea transplants

Image Formation
Each point in the retina receives light from a single point in the scenetwo objects
of different sizes and at different distances can project through the pin hole to
make precisely the same image

Has two important consequences:


There is an inherent ambiguity in the way that images from in the

eye, makes the visual systems job of estimating size and distance

Second consequence of ambiguity between an objects size and its

distance from the observer is that we need some way of specifying
the size and position of a visual stimulus independently of its

Visual angle= angle subtended by an object

If two objects have the same visual

angle, they project onto the same
retinal area

If pinhole is too bigimage is blurry

If pinhole is too smallonly small fraction of

the light can make it through the opening,
cant operate well in medium/low light, diffraction if hole too small

To make sure pinhole is the right size, open it a bit to let enough light in
and avoid diffraction, add a lens to avoid blurring.

Cornea bends light rays that are incident on your eyeacts like a lens
Behind cornea is the lensbends light rays even further.

Combined effect is that they focus the light rays onto the surface of your


When you tae a picture, you need to adjust the focus depending on the
distance of your objectyou have an analogous mechanism in your eye to
do this

Focusing power of cornea is fixed, but focusing power of lens can be adjusted
Lens has muscles attached to it that change its shape and focusing power

When muscles are relaxed, lens has little curvature, serves for viewing
distant objects

When you need sharp focus on a close object, adjust shape by pulling
muscles tautcalled accommodation

If you cant focus properly for distant objectsneed glasses

Failures of focusing:

Myopia: (near-sightedness): eye is too long/lens is too fateven with

relaxed accommodation, far objects are focussed in front of the retina
corrected with concave glasses

Hyperopia: (far-sightedness): eye is too short/lens too thin, with relaxed

accommodation, distant objects are focussed behind the retinasome
degree of accommodation required for focussing on far objects and less
required for near objectsfixed with convex glasses

Astigmatism: cornea is football-shaped (more curvature in one direction

than the other)corrected by adding a cylindrical correction, specified in

eyeglass prescription as cylindrical lens power plus angle at which that

correction should be oriented
Lens of your eye tends to get more rigid with agetil where the muscles can no
longer bend itpeople who have 20/20 all their lives end up needing glasses
around 50presbyopia. Have difficulty seeing under low light levelseye tries to
compensate for the failure of lens accommodation by closing down pupilbetter
focus even without a lens, but suffers from not having enough light
Another condition associated with agingcataractclouding of the lens. May be
present at birth or result of injury/infection, most common cause is old age
If serious enough, can require surgerybreak up and remove old lens and install a
new plastic one with fixed focus (cant accommodate)
Eye Movements
Eye has 3 pairs of muscles that move left/right, up/down, rotate about line of sight
(torsional movement)

First two used to direct gaze to objects

Last used to ensure that the torsional state of the eye is the same no
matter what sequence of eye movements resulted in that choice of

Types of eye movements:

Saccadesquick, ballistic, to change fixation to new object or regard

Smooth pursuitsmooth movements that keep gaze on chosen moving


Nystagmussequence of short smooth pursuits interspersed with

saccades in opposite direction.

Opto-kinetic nystagmuswhen you look at an extended moving


Vestibulo-ocular reflexreflex smooth movement in opposite direction of

head motionkeep fixation on fixed object in the world

Vergenceabove eye movements are conjunctiveboth eyes move in

same direction, vergence is disjunctiveeyes move in opposite directions
to view objects at difference distances from observerconverging to view
nearby object, diverging to view distant object

Tremor, microsaccadesvery small movements that serve to keep images

from being perfectly stable on the retina and from fading (stabilized

images lead to adaptation effects and can fade away completely

The Retina
Purkinje Tree
Light comes in through the pupil, focused by cornea and lens onto retina, passes
through aqueous and vitreous humours and several layers of neural tissue
(ganglion cells, bipolar cells) before reaching photoreceptors.
Apart from the neural tissue between the lens and the light receptors of your eye,
there are also a bunch of blood vessels called the Purkinje treebranches out from
the optic disc (blind spot) where blood supply comes in and optic nerve goes out
Ophthalmoscopebright light with silvered mirrors to stop your head from casting a
Stabilized vision: reason why you cant see blood vessels in your eye shows
important phenomenon about how your visual system worksthey are fixed to the
retina so move with you eye when it movesis a stabilized imagepattern doesnt
change with respect to your retina, your eye is insensitive to stabilized images

Special devices been built that stabilize imagesused for vision

experiments that require careful control over eye movements. Every time
you move your eye, moves the visual stimulus by the same amount.

Under these conditions (stabilized viewing conditions) image of the world

that is projected onto your retina does not make it to your conscious
awarenessafter a few seconds the scene fades, just like the shadows of
your blood vessels.

Diseases of the Retina

Diabetic retinopathy: swelling and abnormal growths of capillaries

insufficient oxygen supply to retina causes bleeding into vitreoustreated
by laser to seal off blood vessels

Detached Retina: retina separates from pigment epithelium, scotoma

(blind spot) in the area that detaches. Caused by glaucoma/injury. Use
laser to cause scar tissue to form and hold retina in place

Macular degeneration: degeneration of part of the retina called the

maculaincludes fovea and surrounding region. Leads to loss of central
visionage related

Retinitis pigmentosa: hereditary disease, initially affects rods and

peripheral then starts to affect conescomplete blindness

Retinal Implants:

No good treatments for retinal diseasesinterest in developing

technology that would help alleviate diseases.

E.g. computer and camera glasses to direct light onto healthy part of

Conceptual design for a retinal implant

Another ideaimplanting microchip with digital camera on one side and

stimulating electrodes on the other sidelike cochlear implants

Right now, only one solution on the marketLVES (Low Vision

Enhancement System)provides variable focus and contrast rather than
neuronal stimulation

Retinal Circuitry (Parallel Pathways)

Retina is a highly organized, layered structureinfo begins at receptors and flows
through a second layer of cellsbipolar cells, and continues through third layer
At the level of the receptor to bipolar connections, there are horizontal cells
connect from receptors to other receptors.
At level of interconnections between bipolar and ganglion cellsamacrine cells
take as their input signals from bipolar cells and feed output back onto other bipolar
In living eye, neurons in retina are transparent so dont interfere with light to
From top to bottom:

Clear ganglion cell axons (optic nerve), stained ganglion cell bodies,
bipolar cells, photoreceptors, black layer of pigment epithelium (absorb
stray light that help in process of regenerating bleached photopigment)

None of the peripheral cells in the retina (receptors, horizontal, bipolars, amacrine)
generate action potentials.

Horizontal/amacrine done even have proper axons

They respond with graded potentialsif the resulting current is large

enough at their synapses, release a neurotransmitter

Ganglion cells (with axons that reach the brain) create action potentials

Big theme in sensory neuroscienceinformation split and processed by separate

subparts of the visual system
Typical properties of parallel pathways:

Physiologically/functionally distinct: rods give you high sensitivity

see at night. But if you didnt have coneswould be blind in high light
levelsboth rods and cones needed

Anatomically distinct: rods and cones have different shapesfor many

parallel pathways, neurons in the separate streams are separated from
one another (except for rods and cones)

Complete coverage: of the visual field by each streamno rods in

central fovea but apart from that they cover entire visual fieldcone
mosaic covers entire visual field

Recombine: parallel streams often remerge eventually. Rod-cone

recombine at ganglion cells. Need different sensitivity of rods and cones
at front end of visual system but later when light is neural signalno
need to keep separate processing.

Midget, Parasol, and bistratified retinal ganglion cells:

Example of parallel pathways

Anatomically distinct:

Parasol dendtritic trees are big, combine inputs from many cones

Midget trees are small

In fovea, midget ganglion cells receive synapses from only one bipolar cell
each which in turn receive synapses from only one cone

In periphery, dendritic fields of both ganglion cell types get bigger

To distinguish between parasol and midgetpay attention to where you

are in the retina and how big the tree is

Complete coverage:

covers large area with no overlap


Midget and parasol streams stay segregated in lateral geniculate nucleus

(LGN), recombine to large extent in visual cortex

Physiologically distinct:

Parasol Ganglion cells (magnocellular LGN cells):


Receive inputs from many photoreceptors

Respond faster and respond to brief transient or moving stimuli

believed to be involved in fast visual processing like visual motion
perception, control of eye movements, and maybe reading

Have poor spatial resolutioncombine inputs from many

photoreceptors (blurring the retinal image)

Combine inputs without regard to cone classcolour blind

Midget ganglion cells:


Receive inputs from fewer cones

Have high (unblurred) spatial resolution

Slower than magno cells

Magnocellular pathway and dyslexia:

Hypothesis that dyslexia is related with a deficit in magnocellular


Used fMRI to testfast-moving, dim light levels stimulus used to isolate

magno pathway to visual cortex.

Brain activity in visual cortex and reading rate measured

Results: weaker magno responses in dyslexic subjects, one can predict

reading rate f rom the strength of the magnocellular pathway responses

Conclusion: dyslexia is related (doesnt necessarily cause dyslexia) to

deficit in magno pathwaypathway that is critical for fast visual

Other parallel pathways initiated in the retina:

About 40 ganglion cell types, 90% of them send axons to LGN then visual cortex,
10% to superior colliculus in midbrain (involved in control of eye movements),
pretectum in midbrain (pupil size, focus), superchiasmatic nucleus above optic
chiasm (circadium rythms)
Information Processing in the Retina
Retina performs 5 important jobs:


Data compression

Light adaption

Spatial filtering

Wavelength encoding

Photoreceptors and Transduction

Two types of photoreceptorsrods and cones (different shapes)

Rods specialized for low light levels (night vision)

Cones specialized for high light levels and colour vision

Rod outer segmentcellular membrane folds in and out

Internal membranes are highly structured and filled with photopigment


In the rods, photopigment is called rhodopsincomplex molecule that is

made up of two parts: opsin, and a derivative of vitamin A

When a quantum of light is absorbed by a molecule of rhodopsin, changes the

chemical state of the photopigmenttwo parts of the molecule splitcalled the
isomerization of the photopigment

Isomerization sets off biochemical chain reaction that leads to an

electrical current flowing in the rodif its big enough, neural signal is
transmitted to the bipolar cells in the retina because neurotransmitter is
released at the rod-bipolar synapse

Absorbing a single photon of light is enough to evoke regular photocurrent

Shown in experiment, rhodopsin pigment colour bleaches when exposed

to light

Tapetum: white reflective surface at the back of alligators and cats eyes, can see
Human cone mosaic:

Rods all have the same photopigment but there are three different types
of cones in the human retinaeach with slightly different photopigment

The 3 photopigments absorb and reflect different wavelengths of light

Retinal Inhomogeneity (Data compression)

Roughly hundred mil light receptors in each eye, only half a million ganglion cells

In order to make original image on the photoreceptors and send it to the

more central portions of the brain, compression takes place

Large part of compression accomplished by having blurry vision in

peripherynot optically blurby analysing that retinal image using fewer
receptors, bipolars, ganglions etc.

Rods and cones are not distributed uniformly across retina

Most cones in/near foveacentral part of the visual field

Most rods are in peripheral part of the visual field

Cones very tightly packed in fovea

Peripheral has a lot more rods than cones, but cones are big to absorb a
lot of light

Peripheral ganglion cells pool inputs from many receptors, have bigger dendritic
trees, have bigger receptive fields, and there are fewer of themhigh spatial
resolution for foveal viewing but still have some vision in periphery and a small
optic nerve

Light/Dark Adaption
Most important thing retina does is transduction by photoreceptors
Second is light and dark adaption
Main challenge that is common to signals carried by all visual neurons is that they
must remain sensitive as ambient light intensity varies over magnitudes
If the light level changes by a relatively small
amount, visual system compensates almost
immediately, if it changes by a lot, eye takes
long time to re-adjust
Adjusting from light to dark is dark adaptation
Threshold intensity decreases over time in the

Data naturally falls into two parts

double branching is due to transition
from cones to rods

The recovery is a fairly slow processsubjects sensitivity to the light

flash continued to increase over a very long timesubject did not fully
recover to the best value for nearly 40 minutes in the dark

Light adaptation in the retina

Part of the mechanism responsible for controlling light/dark adaptation is switchover from rods to cones
But even with a pure rod/cone regime there is still significant adaptation

Only a small part of adaptation is due to pupil size

A lot of the adaptation occurs in the photoreceptors themselvesdue to

photopigment bleaching/regenerationless photopigment available at
high light levels results in weaker responses to light increments at those
high light levels

Adaptation within photoreceptors also helped by feedback from horizontal

cells into the photoreceptors to control the responsiveness of
photoreceptorsrespond strongly then they tell photoreceptors to turn it
down a bit

Differences between rod and cone vision

Daylight vision: photopic, using cones
Low-light vision: scotopic, using rods

At the beginning of the dark adaptation graph, use photopic/cone system.

Cones most numerous in foveasensitivity is best there for photopic

Cones come in three typesresponsible for your ability to

discriminate coloursviolet colour in the graph above was evidence
of cone portion of the curve

Yellow flash of light is easiest to detect and will appear brightest in

photopic vision

Cones are fast respondersability to see fast flicker in photopic

range is high

Cones are more sensitive to stimuli passing through the centre of

the pupil than the edgeStiles-Crawford effectcones are
positioned and oriented toward centre of pupil

Rods (second branch of curve) sensitivity is highest in parafovea (around


Spectral sensitivitygreen appears brightest

Rods are slowcan only detect very slow flicker)

Show no Stiles-Crawford effect

At twilight light levels (mesopic) both cones and rods are available for vision
After being exposed to bright lightexperience afterimage (sometimes bluish) spot
in visual field
Afterimage moves with you when you move your eyesafter exposing a bit of your
retina to bright light, retina becomes light adapted ONLY where the light fell.
Light adaption is local to the region of the retina that was stimulated
If you look at a bright uniform field (white wall) after adapting to small bright light,
you see negative (darker) afterimage.
If you instead look at a very dark uniform field (black wall), afterimage will appear
lighter than backgroundpositive afterimage

Theory is that the adapted retina is signalling a weak stimulus when no

stimulus is present

Consequences for studying the rest of the visual system

Light/dark adaptation happens at the front end of the visual system
Important consequence is that you need to avoid light/dark adaptation when
studying later stages of visual processing
Can ignore light adaptation by restricting choice of visual stimuli to intensity
distributions that modulate about a fixed background intensity

The percent change in the light intensity in an image relative to the average light
Contrast= amplitude of modulation divided by the mean
Contrast of a sinusoidal grating stimulus= difference between the highest and
lowest intensities, divided by the sum of the two.
Maximum possible contrast=1 (100%)

Contrast cant go higher than this because that would mean having the
intensity of the darkest portions of the stimulus go negative

Contrast of a stimulus is changed by scaling the image intensities above

and below the fixed mean background intensity

Neural responses in most of the visual system increase with contrast

Retinal Ganglion Cells
Receptive Fields
Ganglion cells are the first neurons in the retina that respond with action potentials
Response in ganglion cell depends on responses of cells that feed to ganglion
(photoreceptors, biopolars, horizontal, amacrine)
Interested in relationship between ganglion cells activity (firing rate) and visual
stimulus image
People analysed ganglion cells activity:

Small spot of light was flashed on surface of retina for a brief duration,
position of spot was systematically varied across retinal surface,
monitoring cell activity

For most positions on retinal surface, flashing a spot of light has no effect

Within particular region (receptive field), flashing spot affects ganglion

cells response

Receptive field:

Piece of the retina that connects (indirectly) to ganglion cell, or

Region of the visual field in which light stimuli evoke responses in the
ganglion cell

Fixed relationship between the two holds only for a fixed eye position
Receptive field subregions: area within the receptive field is subdivided into two
regions, centre and surround. Two primary types of ganglion cell receptive fields:

ON centre/OFF surround cell: flashing small bright spot in the centre

subregion increases cells response, flashing in surround subregion

inhibits cells response. Little/no response to large spot that covers both
regionsexcitation in centre cancels inhibition from surroundlateral

OFF centre/ON surround cell: oppositeinhibition from small spot in the


Combination of ON-centre/ OFF-centre is another example of parallel pathway

Dendrites of the a-type (OFF centre) and b-type (ON centre) ganglion cells extend
to different layers of the retina where they receive synapses from different kids of
bipolar cells
Linearity of Ganglion Cell Receptive Fields
GCs act like shift-invariant linear systemsinput= stimulus contrast, output= firing
Experiment shows that response of centre and surround cells added together
separately is the same as them responding togetheradditivity rule
GC response is a weighted sum of stimulus intensitieswith positive weights in ON
subregions and negative weights in OFF subregions

Add light intensities at each point (including weights)

Result is a pattern of activity across spatial array of GCs

Neural image (John Robson) refers to the distribution across the retina of GC

Since response is linearcan predict

Neural image (graph of GC responses against retinal position)

Retinal GCs respond to edgescentre surround receptive fields emphasize edges


Intensity: amount of light (physical properties)

Lightness: perceived shade of an object, ranging from black to white


Brightness: perceived intensity of light, ranging from bright to dark


Reflectance: amount of light reflected by an object, giving rise to

brightness/lightness percepts

Retina does not simply record light intensities, retinal responses depend on the
surrounding context (centre-surround receptive field)

Photoreceptor sensitivity depends on the average/ambient light intensity,

due to light adaptation

Retinal GC responses depend on the difference between light intensity in

the centre and that in the immediate surround

Light adaptation and Brightness Perception

Image of a black paper outdoors is more intense than the image of white paper
indoorsbrightness constancy
Perceived brightness depends on the surface reflectance, independent of the
illumination conditions.
The visual stimulus that reach the eyes depends both on the illumination level
(outdoor/indoor light) and the reflectance of the surfaces.
Not interested in the visual stimulus, interested in physical properties of the
surfaces in the world (reflectance of black and white paper) that give rise to the
visual stimulus
Visual system factors out illumination and we perceive the relative reflectance
Light/dark adaptation mechanisms in the retina effectively re-normalize or divide by
the average intensity in the image to
compensate for the level of illumination

Helps visual system achieve

perceptual constancycolours look
the same regardless of level of

Simultaneous brightness contrast:

Having a brightness percept that depends on the context also predicts some other
Illusion showing one side darker than otherretinal responses depend on the local
average image intensityblack side has smaller average intensity
Koffka ring illusion
The visual system is designed to try to achieve a perceptual constancy. The
putative mechanism for brightness constancy is light adaptionretinal responses
depend on average intensity. Most of the time this works but sometimes it screws
upyielding the simultaneous brightness contrast illusion
Spatial Filtering and Brightness Perception
Hermann gridintersections of the white are surrounded with more whiteresults
in more inhibition from the surround in on-centre receptive fields.

Lots of inhibition by surround+ small amount excitation by centre= low RGC

responseperceived brightness is darker resulting in percept of grayish circles in
But why do the gray spots disappear when you look at them?

Receptive fields sizes depend on retinal positioncentral fovea (where

you fixate) are the smallest

May be so small that they fit in the intersectionsno difference in GC

response between intersections and the streets in central fovea

Brightness percept doesnt match how intensity of image actually changes

Mach Bands:

Spatial filtering in GCs.

Shows that retina emphasizes edges

Perceive bright and dark bands at edges of smoth brightness gradient

Dark band is a result of more lateral inhibition

Light band is a result of less lateral inhibition

Surfaces and Brightness Perception

Figure shows different types of edges:

Shading: change in surface orientation from A to B

Reflectance: change in the surface material from B to C

Illumination: shadow has less illumination than the unshadowed portion of

the ground

Adelson Corrugated Plaid Illusion

Brightness judgements can be influenced by high level perceptual factors

(3D interpretation)

Brightness percept depends on the perceived reflectance of the surfaces

Because light appears to come from above, visual system infers that the
upper square is receiving more incident light than lower square.

Because they both have the same physical intensity, upper one must be
less reflective than lower

Perception of transparency also affects perceived brightness

3 processes in vision affect perceived brightness:

light adaptation (divide by the mean, convert to contrast)

spatial filtering/lateral inhibition in retinal GCs. Since these neurons dont

respond much to uniform fields, edges are critical

3D interpretation (shading, reflectance, illumination, transparency)

Physics of Colour/Wavelength
Spectral Power Distribution (SPD) is a plot of energy vs wavelength.

Can be measured using spectro-radiometer

Light that contains only one wavelength is called monochromatic light

Any light can be characterized as the sum of a bunch of monochromatic

lightsplotted in SPD graph

Colour Matching and Trichromacy

Experiment (Helmholtz/Young) to infer that there must be 3 types of
photoreceptors in our eyes

Subject can see test light next to 3 primary lights and asked to adjust 3
intensities to make it match test light


Task is possible, can match any test light as a sum of 3 primary

lights by varying intensities

Lights that are physically different can look identicalsuch pairs are
called metamers or metameric lights. Test light SPD is usually
different from combination SPD, but look identical to eye

3 primaries are always enough to match any test light, 2 not

enough, 4 too many

people behave like linear systemsobeys scalar rule: If you double

intensity, they double settings. If you add two test lights, subject
will set knobs to sum of settings

Application: Colour TV

3 types of phosphors painted on the CRT screen (green, red, blue)

Physiological Basis of Trichromacy

Explanation for colour matching experiment is that there are three types of cone

All that matters is the response of the 3 types of cones

With 3 primaries, can get any combination of responses in the 3 cone


Rods are most sensitive to 500nm monochromatic light

Most cones are sensitive to longer wavelengths than this

Because of this, the brightness of a blue object compared to a red object

increases during dark adaptationPurkinje shift

All rods have the same photopigment (rhodopsin)have the same spectral
Principle of univariance: the response of a photoreceptor is a function of just one
variablethe number of photons absorbed

The response can be identical for:


A weak light at the wavelength of peak sensitivity (few incident

photons, large fraction absorbed)

A strong light at a wavelength of lower sensitivity (many incident

photons, small fraction absorbed)

Cone Spectral Sensitivities

S conesmost sensitive to short wavelengths
L conesmost sensitive to long wavelengths
M cones peak in the middle
Changing the wavelength of a monochromatic light changes the relative responses
of the tree cone typesbasis of your ability to discriminate colours of the rainbow,
each wavelength evokes a unique ratio of cone responses
Cone responses to any test light can be computed by multiplying the test light SPD
by the spectral sensitivities of each cone, then summing over wavelength
Surface reflectance
SPD of light reaching eye = SPD of light source multiplied by the surface reflectance
Response of each photoreceptor depends on the SPD of the light reaching the eye
multiplied by the spectral sensitivity of the photopigment
Wavelength Encoding
Each point in a scene is illuminated by various light sources each with its own SPD.
Surfaces are characterized by the proportion of the light landing on them that is
reflectedspectral reflectance function (relative energy against wavelength)

Product of the illuminant and reflectance yields the colour signalthe SPD of the
light heading toward your eye from the surface.
This signal is analysed by your three cone receptors, respond differentially due to
their individual sensitivities
Only information your brain works with to characterize the colour percept is the set
of 3 responses by the different cone types.
Light source SPD*spectral reflectance function=colour signal (SPD to eye)cone
spectral sensitivitiescolour absorption
Colour Mixture
Lights mix additivelySPD of the sum of 2 lights= sum of two separate lights SPDs

Mixing more of one of the primaries gives more light

Subtractive colour mixturefor pigments

Absorption of the pigments is being combined

Mixing more of one of the pigments gives less light reflected

SPD of a light reflecting off a pigmented surface depends on the spectral

power of the incident light*reflectance of the surface.

Mixing more pigment reduces reflectancereduces SPD of the reflected

light at one or more wavelengths

Summary of Trichromacy theory:

3 types of cones differ in photopigments

each selective for different range of wavelengths

if two lights evoke the same responses in the three cone types, lights will
look the same.

Each cone outputs only a single number (satisfies univariance)tells us how many
photons were absorbed
Colour blindness: there are two basic forms of colour blindnesseither only have
one type of receptor (monochromat) or has 2 types of receptors (dichromat)

A dichromat only requires 2 primary lights to complete colour matching

will accept a trichromats match but trichromat wont accept theirs

Monochromats require 1 primary light to make match

A rod monochromat is missing all 3 cone typesonly have rods. They

dont see colour at all, only different shades of gray. Have to wear dark
glasses during dayotherwise their photoreceptors would be fully
bleached and they would basically be blind.

About 7% of men cant discriminate between red-green colours.

Colour Opponency

Red and green are opponent colours, so are blue and yellow
Established with hue cancellation experimentsubjects were instructed to adjust a
mixture of red and green lights until it appeared neither green nor red
Trichromacy falls our from the fact that you have 3 cone types with different
sensitivities. In the retina, cone signals get recombined into opponent mechanisms

White/black: adds signals from all three cone types L+M+S

Red/green: L-M

Yellow/blue: L+M-S

A colour appears reddish when the red/green mechanism gives a positive response,
greenish when it gives a negative response
Colour opponency in the retina:
Requires very specific wiring in the retinablue/yellow mechanism must receive
complementary inputs from specific cone typesinhibition from S, excitation from
M and L
Bistratified cells do this.

B/Y bistratified ganglion cell receives complementary inputs from the two
bipolar cell classes (S-cone bipolar cells and another) providing excitation
from the S cones and inhibition from the L and M cones

Most of the cones are M and L, only few Ss

Blue-yellow pathway has poor spatial resoltuion

S cones are easy to pigmentanatomists have been able to identify the

retinal circuitry for blue-yellow

Dendritic tree of bistratified GC branches into two separate layers of the retina

Inner tree avoids S cone bipolar cells

Outer tree has synapses with every S cone bipolar cell

Colour Constancy and Chromatic Adaptation

If you take a picture of a room in daylight and fluorescent lightscolours look
different. But not when you see the room
Called colour constancy:

Same as brightness constancy

Eye adapts to compensate for the colour SPD of the light source

Change in percept following adaptation is due to chromatic adaptation

Adapts to whatever colour the ambient illumination is

Each cone type adapts independently

Retinal image adjusts to compensate not only for the overall intensity of the light
source, but also for the colour of the light source

Chromatic adaptation can also lead to dramatic aftereffects

Normally when you look at a white field, L and M cones give about the
same response so the red/green opponent colours mechanism does not
respond at all

If you adapt to green, M cone sensitivity reduced, then when you look at
whiteL:M cones are out of balanceL is more sensitive so you see red
instead of white.

Visual system designed to try achieve perceptual constancy

But like brightness illusions, colour adaptations also result in


Simultaneous colour contrast (similar to simultaneous brightness contrast)

Colour perception, like brightness perception, depends on contrast/surrounding

2/9/2014 11:06:00 AM
Retinal Ganglion Cells
Receptive Fields
Ganglion cells are the first neurons in the retina that respond with action potentials
Response of the ganglion cell will depend on the responses of the cells that feed
into the GC including photoreceptors, bipolar cells, and various lateral
interconnections via horizontal cells and amacrine cells.
We are interested in the relationship between this GCs activity (firing rate) and the
visual stimulus image.

Scientist analysed ganglion cells sensitivity to light using this technique:

small spot of light flashed on surface of retina for brief duration, position
was systematically varied across retinal surfacemonitored GC response

For most positions on the surface of the retina, flashing a spot of light has no effect
on cells response (continues firing at spontaneous rate)
Within particular region (receptive field) flashing spot affects the GCs response:
Receptive field:

Piece of the retina/photoreceptors that connects (indirectly) to the GC

Region of the visual field in which light stimuli evoke responses in the GC

Fixed relationship between these two holds only for fixed eye position

Receptive field subregions: are within the receptive field is subdivided into two
regionscentre and surround. Two main types of GC receptive fields:

On centre/Off surround: Flashing light on centre increases cells

responseflashing bright annulus inhibits cells response. Little/no
response to large light covering both sectionsinhibition cancels
excitationlateral inhibition

Off centre/on surround: opposite.

ON and OFF centre GCs with superimposed receptive

fields give complementary responses, combination of
both ON and OFF centre is a parallel pathway

Dendrites of the a-type (OFF centre) and b-type (ON centre)

GCs extend to different layers of the retina where they receive
synapses from different kinds of bipolar cells
Linearity of Ganglion Cell Receptive Fields
GCs act like shift-invariant linear systemsinput=stimulus contrast, output=firing

Experiment shows that response to centre stimulus alone plus response of surround
stimulus alone equals response to simultaneous stimulation of both (additivity rule)
GC response is a weighted sum of stimulus intensities, with positive weights in ON
subregions and negative weights in OFF subregions

Result is a patter of activity across the spatial array of GCs

Neural imagedistribution across the retina of GC (or other neural)

responses. Since GCs are linear, can predict their responses.

GC response to light intensity in the neural image has spikes (batman head)
because of how it responds to edges

Check previous set of notes for explanation on edges response.

LGN and V1
Retino-Geniculate Visual Pathway
Optic nerve leads from the eye to the optic chiasm where some of the fibres cross.
Optic tract proceeds from the optic chiasm to the lateral geniculate nucleus (LGN)
Optic radiation leads from the LGN to the primary visual cortex (V1)
Visual Fields
Fixation point is the centre of the visual field
corresponds to the fovea
Vertical meridian splits the visual fields into left
and right hemi-fields, horizontal meridian splits
it into upper and lower hemi-fields
Blind spot is the region that corresponds to the
optic disc
Principle of lateralization:

Right half of the brain receives

sensory information from and sends motor
commands to the left half of the bodyright half of
the brain receives information about the left half of
the visual field (left half of both eyes)fibres cross
over at the optic chiasm

Visual deficits due to lesions at different points in the pathway:

If someone has vision problems, type of problem depends on location of problem

If deficit is only one eyes visual fieldophthalmologist

If deficit is in corresponding parts of both eyescentral problem


Lateral Geniculate Nucleus (LGN)

Has 6 layers

Cells have monocular input

Layers alternate inputs from each of the two eyes

Top four are parvocellular layers, two layers from each eye. Parvo (small)
LGN cells receive inputs from midget ganglion cells

Bottom two are magnocellular layers (one from each eye). Magno (large)
LGN cells receive inputs from parasol GCs

Parallel pathways

Retinal (retinotopic) map is preservedaxons from the retina preserve

their order

There is an entire map of the visual hemi-field in each layer of the LGN

Maps are in register in each layer

LGN Physiology
In the LGN, cells have centre-surround receptive fields like GCs
Little or no information processing beyond that done in the retina
Hypotheses of LGNs function:

Brings retinotopic maps from both eyes into register to make it easy for
cortex to combine inputs from the two eyes

Only 10% of inputs to LGN comes from retina, 90% are modulatory inputs
from cortex and the brainstembrainstem modulates information flow
from the eye to the visual cortex e.g. according to the sleep-cycle.
Cortical (feedback) inputs to LGN might have to do with attention. LGN is
a convenient bottleneck for these modulatory inputs from the brainstem
and cortex. If you try to send these projections from brainstem and
cortex back to the retinablind spot would be 10x larger

Primary Visual Cortex (V1)

Visual cortex is subdivided into separate and distinct regionsvisual cortical areas
V1 is located in the calcarine sulcus in the medial occipital love of the brain (back
middle left/right)

Primary because LGN sends most of its axons therefirst visual

processing area in the cortex

Processes information coming from the LGN and then passes its outputs
to the other visual cortical areas (V2, V3 etc.)

All 6 layers of LGN project to area V1 in the cortexmagno and parvo layers
project separately in the input layers of V1 but then merge later (parallel pathways)

V1 retinotopic map
The retinotopic map is laid out across the folded cortical surface in the grey matter
of the calcarine sulcus.
Central (foveal) part of the visual field is represented at the very back and more
peripheral regions of the visual field are represented further forward (anterior).
The retinotopic map is lateralized so that the left hemisphere V1 represents the
right half of the visual field vice versa.
Retinotopic map in V1 is distorted so that the central 10 degrees of the visual field
occupies roughly half of V1makes sense because of the poor acuity in the
peripherythis distortion is called cortical magnification
V1 Physiology
Hubel discovered three different types of neurons in V1 that can be distinguished
based on how they respond to visual stimuli:

Simple cells

Complex cells

Hypercomplex cells

V1 neurons transform information (unlike LGN) so that they are orientation

selective and direction selective
Orientation Selectivity: Most V1 neurons are orientation selectiverespond strongly
to lines/bars/edges of a particular orientation but not to the orthogonal orientation
Direction selectivity: Some V1 cells are also direction selective meaning that they
respond strongly to oriented lines/bars/edges moving in a preferred direction and
not in the opposite direction
Simple cells

Respond best to elongated bars or edges

Orientation selective

Can be monocular or binocular

Have separate ON/OFF subregions

Perform length summation (bigger responses with increasing bar length

up to some limitthen plateau)

Each simple cell sums inputs from LGN neurons with neighbouring/aligned receptive
fields to build an elongated receptive field that is most responsive to elongated
Complex cells

Orientation selective

Have spatially homogeneous receptive fields (no separate ON/OFF


Nearly all binocular

Perform length summation

Hypercomplex cells
Like complex cells except there are inhibitory flanks
on the ends of the receptive fieldresponse
increases with increasing bar length up to some
limit, but then as the bar is made longer the
response is inhibited (no response) (called endstopping)
V1 functional architecture
Hubel found that neurons in V1 with similar response
properties (e.g. same orientation preference) lie nearby one another
Columnar architecture: as one moves an electrode vertically through the thickness
of cortex, most have the same selectivity (same orientation preference and eye
Ocular dominance columns: as you move the electrode tangentially through the
cortex, first find cells that respond to the left eye, then binocular (respond to
both/either eye), then right eye, then binocular, then left again etc.
Orientation columns: as you move the electrode tangentially in the orthogonal
direction, first find cells selective for vertical, then diagonal, then horizontal etc.
Hypercolumn: chunk of the cortex that contains neurons, all with approximately the
same receptive field location, but with all different orientation selectivities, direction
selectivities, and both eye dominances.
Amblyopia (cortical blindness)
Variety of visual disorders when there is no problem with the eye (optics and retina
are fine) but one eye has better vision than the other

Can be caused by strabismus (wandering eye) if not corrected at infancy

During eye development, if the signals from one eye are weak or out of
register with input from the other eye, brain develops in a way that
ignores the signals from the weak/misdirected eye

There is a critical period during developmentmonocular deprivation

during this period messes up the ocular dominance columns, deprivation
after critical period has no effect

To fix in baby, cover good eye for a few hours everyday until end of
critical period

Spatial Frequency Channels

Sine wave gratings
Linear systems theory for vision is more complicated

If talking about an image(picture, retinal image, neural image)which

is a function of two spatial dimensions x and y

If interested in temporal sensitivity (to understand visual motion), signal

is a function of three variables x, y, and t

Analogous stimulus for vision is the sine wave gratingcan vary in spatial
frequency (measured in cycles/degree, for a retinal image), orientation,
phase, and contrast

Contrast for sine wave gratings is defined as Michelson contrast= (ImaxImin)/(Imax+Imin) or (Imax-Imin)/(2 Imean).

Ranges from 0 (bright and dark bars have the same intensity as the midgraygrating is invisible) to 1 (bright bars twice the intensity of the mean
and dark bars are black)

Same definition as Weber contrast or (change in I)/(I)

Characterization of a system in the linear systems theory is the Modulation Transfer

Function (MTF)degree to which different frequencies are amplified or attenuated
by the system.
Behavioural analogy to the MTF is the contrast sensitivity function
(CSF) which describes how sensitive an observer is to sine wave
gratings as a function of their spatial frequencymeasured using
a contrast detection experiment where one determines the
minimum contrast required to detect sine wave gratings of
various spatial frequenciessensitivity is defined as 1/threshold
contrastif threshold is low, sensitivity is high
By tracing out the boundary between visible and invisible
you can make out the curved shape of your CSF
Contrast Sensitivity Function

You are most sensitive for an intermediate

range of spatial frequencies (4-6
cycles/degree) and less sensitive to spatial
frequencies both lower and higher than this

Highest spatial frequency you can see (high frequency cutoff) determines
your spatial acuityfinest spatial patterns you can see. Acuity limit
decreases with age

Multiple Spatial Frequency Channels

CSF is not thought of as the MTF of a single kind of neuron,
but an envelope of sensitivity over several underlying
mechanisms, each corresponding to neurons with differing
preferred spatial frequencieswith different sizes of
receptive field, larger= lower spatial frequency preference)

Graph shows four spatial frequency channels,

notion is that the CSF represents the sensitivity
pooled over those underlying channelssensitivity
is primarily determined by whatever channel (set of
neurons) is most sensitive to the stimulus

Low spatial frequency filters encode coarse luminance variations in the

world (large objects, overall shape)

High spatial frequency filters respond to the fine spatial structure of the
world (small objects, detail)

Evidence for the existence of multiple spatial frequency channels:

Physiological: in V1 and beyond, for each

location in the visual field there are neurons
varying in preferred spatial frequency,
orientation, direction of motion and so on.

Behavioural: consider effects od adapting to a

particular spatial frequencybegin by
measuring CSF then observer stares at
particular sine wave grating for extended
period. Visual system adapts to the pattern
any neurons or mechanisms that were sensitive
to that pattern become desensitized
temporarilyre-measure CSF

Dotted curvepost adaption CSFsensitivity is reduced only for gratings

with spatial frequencies near that of adapting gratingthose with spatial
frequency preferences distant from the adapter remain unaffected

Spatial frequency adaptation affects threshold and the appearance of the

supra-threshold gratings

Adaptation to the higher spatial frequency desensitizes

the higher spatial frequency channels
When you adapt to a lower spatial frequency grating, you
see the opposite shift in the response
Similar shifts happen with orientation and direction of
motion, indicating that there are channels tuned for
various orientations and directions of motion
Two other common methods of demonstrating the
existence of multiple spatial frequency channels

Summation: if you ask an observer to detect a combination of two

gratingsensitivity is much higher if the two gratings are close in spatial
frequency (detected by the same channel)

Masking: observer is asked to detect on TEST grating (frequency f1) in

the presence of another masking grating (frequency f2)masking grating
is always present, question is to what degree does it mask the test
grating. When the masking grating is similar in spatial frequency to test,
masking is strong, when dissimilarmasking is weak. One grating masks
another only to the extent that both are detected by the same channel

Visual Motion Perception

Motion is a perceptual attribute: visual system infers motion from the changing
patter of light in the retinal imageoften incorrect.
After viewing continuous motion in the same direction for a long time, if you look at
a stationary object it appears to move in the opposite direction

Shows that this adaption is local in the retina (to the right of where you
were looking, you were adapting to rightward motion, to the left you
adapted to leftward)take this as evidence for the existence of neurons
that are sensitive to motion and selective for the direction of motion,
which adapt to the stimulus

Role of motion perception: has lots of helpful functions

Detecting that something is movingdraws your attention to it

Segmentation of foreground from background

Computing 3D shape

Computing distance to various objects in a scene and estimating direction

in which you are heading in a scene

When there is strong motion on your retina, especially in peripheral

regions, you can misattribute that motion and perceive yourself as
moving (vection)

Recognising actionsmovements

Optic flow: physical motion in an image while an observer moves through the

Focus of expansion (point that all the arrows come out of) where observer
is heading

First step in motion perception is for visual system to estimate optical flow
from the changing patter of light in the retinal image

3D motions of observer and objects can be inferred from optical flow

which then provides info about the observers heading and relative
distance to each surface in the world

CAVEAT: small close object when youre moving slow looks same as far
big object when youre moving fast

Object motion:

When an observer moves through a stationary environmententire

retinal image changes over time

When object moves and observer is stationarysmall region of retinal

image changes over time

Motion provides info about shape even in absence of other shape cues

Biological motion: provides information to allow recognition of human movements

Visual Motion in the Brain
Functional Specialization hypothesis: there are specific brain areas that are involved
in visual motion perception
Current opinion is that the optical flow field is computed and represented by
neurons in area MT

Neurons in MT are selective for motion direction

Neural responses in MT are correlated with the perception of motion

Damage to MT or temporary inactivation causes deficits in visual motion


Electrical stimulation in MT causes changes in visual motion perception

Computational theory quantitatively explains both responses of MT

neurons and the perception of visual motion

Well-defined pathway of brain areas underlying motion specialization in


MT neurons receive inputs from direction-selective neurons in V1they are velocity

selective, independent of stimulus patternresponds to almost any patter with
almost any contrast as long as it moves with the right velocity
Human cortical area MT can be identified with fMRI
There is columnar architecture in MT for stimulus motionneurons with similar
motion preferences lie nearby one another
MT and motion perception: activity of MT neurons is correlated with motion
perception (experimenters used electrical microstimulation to change the responses
of a small number of MT neurons and showed that this affected motion perception

Trained monkeys to perform difficult motion discrimination taskviewed

moving dots and decided which way they went, then characterized how
MT neurons responded during the taskfor a neurons preferred direction
of motion the neurons respond more as coherence increased.

Inserted stimulating electrodes into a particular column of MT neurons

and injected pulses of current while the monkeys performed the motion
discrimination task

Results showed that stimulating MT neurons directly influences behaviour,

presumably because it influences the monkeys conscious percepts.

Visual area MST and optic flow

Cortical area MST is right next to MTMST neurons have very large receptive fields,
respond selectively to complex optic flow fieldsexpansion, contraction, rotation.
It is believed that MST is involved in 3D motion perception, inferring 3D motion of
objects/observer from optical flow.
Neurons in the area STS respond selectively to biological motion, like the point-light
Eye Movements and Motion Perception
When observer is moving, visual system uses the changing retinal image to infer
the observers trajectory and 3D structure.
Visual images are combined with other information to inform you about the motion
of your eyes, head, and body
Vestibular system provides information about the motion of your head and body.
Vision is combined with vestibular and eye movement signals in Area MST.

Brain is divided into motor areas and sensory areas. Corollary discharge is a copy of
the motor signal that is transmitted to a comparatora hypothetical structure that
receives both the corollary discharge and the sensory movement signal
If visual motion signal is different from the eye movement command, then you see
Depth, Size, and Shape
Stereo Vision
Stereopsis: greek for solid sight
Relative distances are different in each eyerelative positions in the two retinae are
Binocular disparitydifference in the location of a feature between the right eyes
and left eyes image. Amount of disparity depends on the depthis a cue that the
visual system uses to infer depth.
The disparity also depends on the distance to the fixation as well, so that disparities
must be further interpreted using estimates of the fixation distance.
Horopter: imaginary 3D surface in the room in front of you that includes the object
you are fixating on and all other points in 3D space that project to corresponding
positions in the two retinae. The geometric horopter (the set of points with 0
disparity) is a circle that includes the fixation point and the optical centres of the
two eyes.
Uncrossed disparity: object further away from you than the horopter has uncrossed
disparitiesyou would have to uncross your eyes to fixate on it, it lies further to the
right from the right eyes viewpoint than from the left eyes viewpoint
Crossed disparity: object closer than the horopter has crossed disparities. Youd
have to cross your eyes to fixate on itfurther to the left from the right eyes
Stereoscope: one way to view stereo image pairs is to use a mirror stereoscopeif
you put your face in front of a pair of angled mirrors and put two slightly different
pictures off to the sides, your left eye will see the left picture, your right eye will
view the right-hand picture
Stereogram: pair of images that are viewed using a stereoscopetwo images are
slightly different with features in one image shifted to slightly different positions in
the other image. The shifts mimic differences which ordinarily would exist between
the views of genuine 3D objects
Random-dot stereogram: indicates that:

You can see depth from binocular disparity without any other depth cue
present (motion parallax, perspective etc)

You can see depth without first extracting delineated form, or a

recognizable objectbinocular combination is early, precedes processing
of recognizable forms/shapes/objects

You can determine which dot in the left eye goes with which dot in the
right eye in the presence of many potential false matcheschosen so that
the inferred 3D scene is relatively smooth and continuous

How does the visual system see depth in a random-dot stereogram?

One hypothesis is that the visual system matches up features of similar

shape, size, contrast, etc. To estimate disparity

Problem of resolving this ambiguity of matching the dots is called global

stereopsisbrain must find the correct overall (global) set of matches

Autostereogram: also known as the magic eye stimulus

Display slightly different images to the two eyes. Autostereogram works

by having repetitive patternsto see depth in an autostereogram, you
need to either cross or diverge your eyes so that they fixate separately on
two different repeats of a repetitive patterneffectively get two different
images to the two eyes.

Stereoblindness: 10% of people are stereoblind, some only blind to either crossed
or uncrossed disparities. Some caused by strabismus (wandering eye)if not fixed
at infancy binocular vision never develops properly.
Some people with strabismus end up with amblyopialazy eye
Amblyopia is a general term used for a visual deficit that has nothing to do with the
optics or eye/retina structure
Other people with untreated strabismus end up as alternate fixators who can see
with either eye but never use them both at the same timefirst look at you with
their left eye (right is diverged) then switchno binocular vision and no stereopsis
Stereovision in the brain
Some neurons in V1 are selective for particular disparities
Some neurons dont respond at all when a line is shown to one eye at a timeto
respond the line must be presented to both lines simultaneously to both eyes, have
the correct orientation, direction of motion and the correct binocular disparity
Neurons differ in the binocular disparities to which they are tuned

Many neurons are tuned for 0 disparityon the horopter, some tuned for
a range of crossed and uncrossed disparities

Fusiontwo visual fieldstwo images are combined in the brain to yield a single
unified perceptual experience
Things that can happen with 2 eyes:

Panuns fusional area: range of disparities, or equivalently the range of

depths in 3D space on either side of the horopter, over which the visual
system can successfully fuse the two views. If disparity is small enough
(within fusional area), visual system succeeds in fusing the two viewsif
disparity is too big you get diplopia or suppression or binocular rivalry

Diplopia (doube vision): when you see two of something

Suppression: what normally happens when the retinal disparity is too

bigone eyes view dominates and is perceived.

Binocular rivalry: when views from the two eyes are very different. One
eyes view dominates for several seconds then is replaced by the other
interesting because physical stimulus doesnt change but conscious
percept changes dramatically over time, have no conscious control over it.

There are two ways to have single vision

Small disparity yields fusion and stereopsis

Large disparity often causes one eyes view to be suppressed

You can have diplopia, suppression, and binocular rivalry at the same
time in different parts of the visual field

Binocular rivalry in the brain

Because of the dissociation of changing perceptual experience, binocular rivalry
presents a unique opportunity for studying the neural correlates of consciousness.
According to one idea, binocular rivalry occurs because neurons in the early stages
in visual processing respond to the physical stimulus of each eye, whereas neurons
in later stages of visual processing are switched on and off and cause the
perceptual alternations.
Somewhere between these early and later stages the neuronal signals conveying
one of the two stimuli are suppressed, as if there was a gate to visual
consciousnessmight be in V1 because thats the first place where the signals from
the two eyes come together.

Studies show that during a perceptual alternation, one typically perceives

a traveling wave in which the dominance of one patter emerges initially at
one location and expands progressively as it renders the other pattern

By relying on the fact that the visual cortex is retinotopically organized

(neurons at nearby locations in visual cortex respond to nearby locations
in the visual field), showed that neural activity propagated over
subregions of visual cortex in a manner that correlated with the dynamic
perceptual changes experienced during binocular rivalry.

Activity in a number of brain areas correlates with the perceptual alternations of

binocular rivalry, including higher-order visual cortical areas in the inferior temporal
lobe, and areas in the parietal and prefrontal cortex.
Pictorial Depth Cues
With only one eye open, still see with a sense of depth but there is inherent
ambiguity between size and distance.
Visual system relies on multiple cues for estimating/inferring distance, depth, and
3D shape such asrelative size, occlusion, cast shadows, shading, dynamic
shadows, aerial perspective, linear perspective,
texture perspective, and height within the image.

Most of these are based on the concept that

the size of the retinal image of an object is
proportional to the objects size, but
inversely proportional to the distance to the

Texture provides 3 cues about shape/distance:

Texture elements become more dense with distance

Texture becomes smaller with distance

Foreshortening (circles become ovals) when the surface is tilted

Linear perspective is a monocular depth cue

Monocular Physiological Cues
When we fixate on an object, accommodate to itchange the power of the lens to
bring it into focusaccommodative effort is a weak cue to depth.
Once we accommodate, objects closer/further are blurryblur is a cue that objects
are at a different distance than the accommodative distance.
Even weaker cue is image distortions due to astigmatism and chromatic aberration
Movement Cues
2 cases:

observer moves through stationary environmentresulting movement is

called motion parallaxobjects move at different speeds on your retina
depending o their distance from the observer

object moving when observer is stationaryresulting retinal velocities will

depend on the relative distance of each object feature from the
observerkinetic depth effect

Object recognition is used for a lot of thingsto recognise a particular type of
object (a moose), a particular exemplar (this moose), to recognise it (the moose I
saw yesterday), or to match it (the same as that moose)
What is recognition?

Match between visual input (processed through the ventral stream) and a
mental representation of an object

Impressive that we can rapidly categorize an object even though there are many
different kinds of ducks including drawings of ducks and rubber ducks
Three recognition themes:

Category specificityfaces and places

Nature of representation3D viewpoint-invariant or 2D viewpointdependent

Perceptual organizationgrouping and segmentation

Is face recognition special?

Prosopagnosia (face blindness): affects a lot of peoplecant recognise faces
Object agnosia: cant recognise common objects but have no problem with faces
Double dissociation: some patients show a specific deficit for recognising faces,
other subjects show deficits for recognising all other objects. This suggests that
faces and objects are processed in separate, perhaps non-overlapping, brain areas.
Face inversion effect: theory that upright and upside-down are processed differently
Results from the experiment:

Normal subjects performed better for upright faces than invertedimplied

specialized processing for faces

Patient LH suffers from prosopagnosiadid relatively poorly compared to

control on both upright and inverted. But performed better for inverted
than for uprightimplies that an impairment of a face-specific processor
(engaged by upright but not inverted faces) used despite being

Face selectivity in IT: Recorded neurons in area IT of monkey cortexfound strong

responses to faces

In the experiment, each panel plots the response of an IT neuron in response to

each of the pictures
Top row:

Strong response to intact face

Scramble features randomlyvery little response


Controversial responseno matter how many manipulations you do, determined

skeptic says theres some kind of simple feature there that is triggering cell
Functional (columnar) architecture for faces: used optical imaging in IT to
demonstrate columnar organization
Doll face experiment shows that neurons in nearby columns respond to similar
complex feature patterns
fMRI and face selectivity: one human brain area responds most to faces
neighbouring brain area responds most to pictures of buildings and familiar places,
another to pictures of objects that are neither faces nor places
Role of expertise: subjects practiced to learn and make dine discriminations in a
novel category of objectsat first performance was was like that for objects, after
lots of practice, better at recognising.
Human single-neuron electrophysiology: patients with severe epilepsy undergo
surgery to have some electrodes implanted in their brain to record the electrical
activity in the brainstay at the hospital and wait for seizures to occur and
measurements are taken, once located, have surgery to remove diseased brain
Neurons are very selective (Jennifer Aniston)

Important caveat: this part of the brain is involved in memory, neuron

may be activated by some memory that is triggered by the photo

Summary: Evidence for brain areas that are functionally specialized for face

Infants show a tendency to track a moving face at just 30 minutes of age

Face agnosia without object agnosia in some patients

Object agnosia without face agnosia in other patients

Inversion effect: normal subjects are better at recognising upright than

inverted faces

Inverted inversion effect: prosopagnosic was better at recognising

inverted than upright faces

In fMRIfaces light up separate brain region

Face selective cells in inferior temporal cortex (IT) and in human medial
temporal lobe (MTL)

Evidence that face recognition is not functionally specialised in particular brain


Face columns are intermingled with other feature columns in most areas
of the monkey brain where face-selectivity has been found

Face agnosics often show impaired recognition/discrimination ability for

non-face objects within a category

After lots of training with greebles, fMRI activity shifts from object area to
face area in the brain

Reading and Letter Recognition

Letter recognition (like face) is believed to be handled by special-purpose circuitry
in the brain
Object Representation
We can rapidly recognise any familiar object from almost any viewpoint under
almost any lighting conditions
Recognition by parts: suggested that recognition works by decomposing objects
into a fixed set of primitive shapes (there are 36 such primitives), that he calls

Idea is that we first recognise each of the constituent geons and then the
spatial relationship between them.

Evidence? We can no longer identify an object when we occlude the

objects geons by obscuring their intersections

View-independent recognition: alternative theory

You store in your head a bunch of characteristic views (mental images) of objects
you recognise a new image by finding the closest matchyou dont use 3D shape
to recognise objects, only the 2D view of the objectsexperiment: show object
rotating a bit, test recognition of that object for similar views, different views, and
distractor objects. Results: good on similar views to those used in training, bad for
very different views
Recognition in the brain: trained monkeys in an object recognition task like the one
above, results: view selective responses were found in IT, responses to distractors
are much lower, even when the distractor contains features that are similar to the
targetobject recognition is viewpoint dependent
Perceptual Organization
Segmentation: divide the visual field into distinct parts/objects

Grouping: integrate visual information across different patches (fill in the missing
You cant recognise an object until it is properly grouped/segmented
Some V2 neurons respond to illusory contoursneuron responds to the orientation
of the illusory contour, not the orientation of the real lines that define it.
V1 neurons dont respond this waythey always respond to the orientation of the
real lines, ignoring the orientation of the illusory contour.
Responses to illusory contours have also been measured in the human brain with
Grouping and crowding: crowdingvisual system can process only a modest
number of featuresif there are too many features then it cant cope and fails to
recognise anything
Visual system sometimes struggles (trying multiple groupings and segmentations)
to perceptually organize a stimulus to figure out which of these features belong
together and which dont resulting in a dynamic percept in which different
organizations compete with one another
Grouping is based on many visual features:



Common region

Good continuation



Common fate

Different brightness

Differences in texture

Secondary Cortical Visual Areas and the What/Where Pathways

Secondary Cortical Visual Areas
Around 30 visual areas after V1most in occipital
Because surface of cortex is folded, have to open
sulci and flatten to visualize locations of the visual
Functional specialization hypothesis: main
hypothesis driving research into secondary visual

cortical areas, specific brain area are involved in specific visual functions
Defining visual cortical areas: visual cortical areas are defined and identified based
on both physiology and anatomy (PhACT: Physiology, architecture, connections,
Physiology: Neurons in different brain areas are selective for different things
Architecture: Cytoarchitecture can change from one area to the next, helping to
identify the boundaries between distinct brain regions
Connections: inject animal with tracer and then kill and stain to see where it ends
Topography: each of the separate early visual areas contains a retinotopically
organized map of the visual fieldcan measure retinotopy using fMRI in the human
brain using stimuli that traverse the visual field
Parietal and Temporal Pathways
Cortical visual areas are interconnected in a complicated way but there is one
general trend:

Two distinct streams (parallel pathways), one goes to parietal love and
other to temporal

Parietal lesions lead to deficits in spatial orientation, attention

Temporal lobe lesions lead to deficits in object recognition

Patients with temporal lobe lesions are aware that theres a problem and they
develop strategies to compensate for it, parietal patients are unaware of their
Attention and Awareness
Hemifield Neglect
Parietal lobe lesions cause deficits in attention and spatial operations, deficits are
most severe when the lesion is in the right hemisphere and most severe just after a
stroke, gets better over time.

Neglect: patient ignores half of their body, get dressed on one side, put
makeup on one side of their face.

Extinction: patient sees normally in the left hemifield, put an object on

their left and can see and identify it, if you show two objects, one in each
hemifield, they dont see the object on the leftthey deny it was there

Denial: sometimes are unaware that they have a problem, when

confronted they will deny and confabulatehold their right arm, they say
its theirs, hold their left arm, say it must be yours

Spatial orientation deficit: can be confused in familiar surroundings, know

its a drug store near their house, know its the usual bus stop, but dont
know how to get home

Summary: patients suffer from loss of some forms of awarenessespecially to do

with the relationship between their own body and the visual world.
Lots of evidence that parietal love is involved in sensory motor integrationwhich is
why these symptoms show loss of awareness of the visual world and sometimes
loss of body awareness
Patients with temporal lobe lesions (face and object agnosia) are fully aware that
theres a problemcompensate for it
Action without Awareness
Blindsight: two famous subjects: GY and DB, had brain damage to V1 in one
hemisphere leaving them totally blind in large parts of the contralateral visual
hemifieldwhen forced choice trials, get 43% correct for blind areaknow
something happened but cant see it (hatched area)
Split brain patients: cut corpus colossum to treat epilepsy. For right handed
patients, language in left hemisphere. The subject fixates at a point and a word
flashes up on one side of the point, subject reports only words flashed on right.
When instructed to pick up corresponding object to the flashed word, left hand
correctly retrieves objects for words subject doesnt report seeing
Consciousness, whats it good for? Blindsight patients residual vision allows for
rudimentary discriminations for action, but cant use their residual capacity for
thinking or imagery (mental manipulation of a percept)
Visual Attention
Eye/head movements: always moving your eyes around to foveate key visual
Over attentionmost direct way to shift attention. Poor resolution in periphery
means that you are aware primarily of things near the centre of gaze
Covert attention: you can shift attention without eye/head movements to filter out
unattended locations
Experiment: subjects perform difficult visual discrimination taskcue before each
trial that tells you where to pay attention, sometimes cue correct and sometimes
wrong. On averageperform better when cued correctly

Can shift spatial attention without eye movements

Shifts can be controlled by subjects decision to allocate attention or by

automatic attraction

Performance falls off smoothly with distance when cued to the wrong

Automatic shifts of attention are linked in time to the occurrence of the

cueperformance is worse if there is a long delay between cue and target
(sometimes called inhibition of return)

Visual search: Some patterns are easily discriminable (pre-attentive), but others
require something like a serial/scanning search of the visual stimulus
Difficult discriminations require attentionitem by item serial search
Reaction time is increased linearly with an increase in the number of distractors,
this increase for each irrelevant item is twice as big on trials where the target is
absent than on trials where the target is presentobserver needs to attend every
item when they need to make sure its absent
Feature integration theory (Treisman): theory for explaining which patterns are
easily discriminable and whyidea that front-end of the visual system breaks the
stimulus down into its constituent parts and separately analyses them to determine
patter, motion, shape, colour etc.
Attention is key in feature integration theory.
Feature integration theory is based on the functional specialization hypothesis, one
needs attention to bring separate neuronal representations together
Illusory conjunction: if you flash image of woman in red dress and black hair, might
recall as woman with red hairillusory conjunction is taken as evidence for feature
integration theorynot enough time for attentive processing to combine features
Change Blindness
Very large changes occurring in full view in a visual scene are not noticed, the
changes are arranged to occur simultaneously with some kind of extraneous brif
disruption in visual continuity.
These phenomena are attracting an increase amount of attention because they
suggest that humans internal representation of the visual world is much sparser
than usually thought.
Attention in the brain
Attention can affect neural responsesrecorded from neurons in parietal love while
monkeys were performing various tasks they were trained for.

Conclusion is that visual responses in cells in the secondary cortical visual areas are
gated according to the behavioural significance of the stimulus. There are neurons
in the brain that are correlated with attention, when a visual stimulus is relevant for
the task at hand, those neurons are more active.
Attentional modulation of V1 brain activity:
Classical view is that V1 acts as a passive, automatic, image processing machine.
Subjects viewed moving stimuli, one positioned to the left and one to the right of
the centre of fixation. Shape of fixation point told them which side to pay attention

Attend left condition increases brain

activity in the right hemisphere

Attend right condition increases brain

activity in the left hemisphere

Attention reaches all the way back to the

first visual area in the cortex

Mental Imagery
Consciousness is associated with the capacity for
thinking or imagery (mental manipulation of the percept)
fMRI experiments suggest that imagery and perception result in a similar pattern of
brain activity
recorded fMRI signals in the face area (that responds strongly to pictures of
faces) and the place area (that responds strongly to pictures of familiar places

fMRI response is bigger in the face area when imagining a face and bigger
in place area when imaging a place

Scientific Study of Consciousness

What is consciousness?

Not all operations of the brain correspond to conscious processes

Closely associated with attention

Involves some form of memory

Seen how brain activity is linked with perceptioneven in cases for which the
percept is entirely illusory:

Brain activity in V1 fluctuates with perceptual alternations during

binocular rivalry

Microstimulation in MT biases monkeys judgements of motion

Brain activity in MT is correlated with the perception of illusory motion

during the motion aftereffect

Neural responses in V2 are correlated with the perception of illusory


Brain activity in many visual areas including V1 modulates with shifts in


Imagining different attributes of visual stimuli (e.g. colour, motion, faces)

selectively activates the specific brain pathways involved in the perceptual
processing of those stimulus attributes

Do not yet understand specifically what distinguishes neural activity that underlies
conscious mental statesseveral hypotheses but none of them yet have strong
empirical support.
Recently, scientists have been exploring the function and organisation of the human
brain under more natural and unbounded settings.

Results of fMRI while watching a clint eastwood movie shows that 40% of
each individual brain does the same thing as other brains when watching
the same movie

Unified nature of conscious experience in fact consists of temporally

interleaved and highly selective activations in an ensemble of specialized
brain regions.

One area of the brain responds every time there is a close up of a

face, another analyses outdoor scenes and landscapes etc.

2 first midterm, 2 second midterm, 2 third section

conceptual and algebra
optimization problems
maximising preferences
compensating variation, fixed substitution effect
dont focus on continuous strategies
substitution and income effect
NASH eqm
No profit max, but yes cost minimization