Professional Documents
Culture Documents
Subjectivist View: There is no inherent organization to the world, but rather, our
brains organize our perceptionstherefore we believe the world is, itself, organized.
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.
The sensory systems of your brain must make some kind of inference
because you do not have the opportunity to perceive the world directly.
o
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
Neurons
Structure: dendrite, cell body/soma, axon (electric signal goes down axon starting
from cell body)
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
Action potential starts at a neurons soma and travels down the axon
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
Neurons in the different parts of the brain are selective for different
things.
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
e.g. blind sight (residual ability to discriminate visual stimuli when forced
to guess, even though the subject has no conscious experience of the
stimuli)
Non-invasive method for measuring activity in the human brain, and for
investigating the relationship between brain activity and behaviour
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.
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
Magnentoencephalogram (MEG)
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.
o
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
Neuroethics
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
P= perceived magnitude
S= stimulus intensity
K= constant
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
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
Sound
This brief disturbance is the only information available to you about the
hand clap
Hearing sensitivity to different frequencies: for very low/very high frequencies, pure
tone must have a very high amplitude to be heard
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
equalizers divide up the signal into many frequencies to give you finer
control
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
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 energy in the fluid is largely dissipated by the time it reaches the
helicotrema
The arches of corti are a complicated set of structures sitting on the basilar
membrane. The main parts were interested in are:
Transduction
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
bit.
This is the key motion that converts the mechanical motions into an
electrical signal used by the rest of the nervous system.
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
potential
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
direction
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:
Additivity: S1+S2=R1+R2
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
impulses.
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:
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
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
Then, take our input stimulus and use the Fourier Transform to compute
the values of the coefficients in the Fourier Series expansion
o
Can predict the systems response by adding the responses for all the
component sinusoids
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
tones
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.
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
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
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
perception.
Loudness Perception and Critical
Bands
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:
Total energy: summed energy of all pure tone components, which is the
area under the Fourier spectrum curve
Distance
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
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
frequencies
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
LSO:
MSO:
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
difference
How?
o
Anatomy of MSO:
o
The neurons are connected in a way that looks like Jeffries wiring
diagram
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
You can rotate your head so that one ear is directly in front of the source
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
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
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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
Pupil: hole
Iris: Coloured part, has muscles that change its shapechanging pupil
size
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 nerve: made up of ganglion cell axons that exit through the optic
disc
Cornea is living tissue but must be transparentno blood vessels running through it
Glaucoma: without blood vessels, nutrients are passed to the cornea and
lens through a continual flow of liquids through the eye
o
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
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
retina.
Accommodation
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
Failures of focusing:
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
fixation
Retinal Implants:
E.g. computer and camera glasses to direct light onto healthy part of
retina
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.
Ganglion cells (with axons that reach the brain) create action potentials
Anatomically distinct:
Parasol dendtritic trees are big, combine inputs from many cones
In fovea, midget ganglion cells receive synapses from only one bipolar cell
each which in turn receive synapses from only one cone
Complete coverage:
Recombine:
Physiologically distinct:
Transduction
Data compression
Light adaption
Spatial filtering
Wavelength encoding
Tapetum: white reflective surface at the back of alligators and cats eyes, can see
bleaching.
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
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
dark
At twilight light levels (mesopic) both cones and rods are available for vision
Afterimages
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
Contrast
The percent change in the light intensity in an image relative to the average light
intensity
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
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
Receptive field:
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:
inhibits cells response. Little/no response to large spot that covers both
regionsexcitation in centre cancels inhibition from surroundlateral
inhibition
Retina does not simply record light intensities, retinal responses depend on the
surrounding context (centre-surround receptive field)
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
Colour
Physics of Colour/Wavelength
Spectral Power Distribution (SPD) is a plot of energy vs wavelength.
Subject can see test light next to 3 primary lights and asked to adjust 3
intensities to make it match test light
Results:
o
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
Application: Colour TV
Explanation for colour matching experiment is that there are three types of cone
photoreceptors
All rods have the same photopigment (rhodopsin)have the same spectral
sensitivity.
Principle of univariance: the response of a photoreceptor is a function of just one
variablethe number of photons absorbed
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
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)
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
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
Dendritic tree of bistratified GC branches into two separate layers of the retina
Eye adapts to compensate for the colour SPD of the light source
Retinal image adjusts to compensate not only for the overall intensity of the light
source, but also for the colour of the light source
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.
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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.
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:
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:
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
GC response to light intensity in the neural image has spikes (batman head)
because of how it responds to edges
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:
Has 6 layers
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
There is an entire map of the visual hemi-field in each layer of the LGN
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
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
Orientation selective
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
bars/edges
Complex cells
Orientation selective
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
dominance)
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
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
To fix in baby, cover good eye for a few hours everyday until end of
critical period
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)
Highest spatial frequency you can see (high frequency cutoff) determines
your spatial acuityfinest spatial patterns you can see. Acuity limit
decreases with age
High spatial frequency filters respond to the fine spatial structure of the
world (small objects, detail)
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
Computing 3D shape
Recognising actionsmovements
Optic flow: physical motion in an image while an observer moves through the
environment
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
CAVEAT: small close object when youre moving slow looks same as far
big object when youre moving fast
Object motion:
Motion provides info about shape even in absence of other shape cues
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
motion.
Depth, Size, and Shape
Stereo Vision
Stereopsis: greek for solid sight
Relative distances are different in each eyerelative positions in the two retinae are
disparate
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
perspective
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 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
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:
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.
You can have diplopia, suppression, and binocular rivalry at the same
time in different parts of the visual field
Recognition
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:
Etc.
Summary: Evidence for brain areas that are functionally specialized for face
recognition:
Face selective cells in inferior temporal cortex (IT) and in human medial
temporal lobe (MTL)
Face columns are intermingled with other feature columns in most areas
of the monkey brain where face-selectivity has been found
After lots of training with greebles, fMRI activity shifts from object area to
face area in the brain
Idea is that we first recognise each of the constituent geons and then the
spatial relationship between them.
Grouping: integrate visual information across different patches (fill in the missing
parts)
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
fMRI
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:
Similarity
Proximity
Common region
Good continuation
Symmetry
Closure
Common fate
Different brightness
Differences in texture
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,
topography)
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
up
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
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
deficits.
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.
Symptoms:
Neglect: patient ignores half of their body, get dressed on one side, put
makeup on one side of their face.
Performance falls off smoothly with distance when cued to the wrong
location
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
correctly
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
to.
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
etc.)
fMRI response is bigger in the face area when imagining a face and bigger
in place area when imaging a place
Seen how brain activity is linked with perceptioneven in cases for which the
percept is entirely illusory:
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