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Last time, you learned about how pressure

waves
are converted into a neural signal in the
cochlea in the ear, and you also saw
that there was something missing from this
process.
Namely, that the sound wave after it
entered the ear
canal lacked any information about where
the sound came from.
And, yet we're able to localize where
sounds are coming from.
So how does the brain do this?
That's what we're going to talk about in
this video.
So, two sounds coming from two different
locations, their sound waves would merge
in
the ear canal and any information about
where those sounds came from would be
lost.
But fortunately, we're able to compute the
location of sounds.
And, we do so in part by comparing the
sound
waves that arrive at each ear and, here's
how this works.
If the sound is located over here, it will
arrive in this ear first and be slightly
delayed
to arrive in that ear because of the
longer
path that it takes for the sound to get
there.
Even though the extra distance that the
sound has to travel is fairly short, it's
enough to make a measurable difference in
how long it takes the sound to get there.
Furthermore, the difference in path link
varies smoothly with the exact angle.
So a sound that is located more straight
ahead, has a fairly small
difference in the distance of the far ear
as compared to the nearer ear.
A sound located farther to one side,
involves a bigger difference
between the, distance of the near ear and
the far ear.
These differences are referred to as
interaural time delays or ITDs for short.
So let's do a few calculations.
How long exactly are these delays?
So, to figure out those delays we need to
know the separation between the two ears.
How far apart are the two ears?
As well as the speed of sound.
And we'll do the calculation for a sound
located straight
off to the right or straight off to the
left,
where the difference in path length would
be the greatest
and so, the difference in timing would
also be the greatest.
Well, I made a few measurements for
myself.
I measured the diameter of my head by
taking off my
glasses and measuring how far apart the
two ear pieces are.
And, I only had a ruler with inches on it,
but I did some calculations.
So, my ears are about five and a half
inches apart.
That corresponds to 0.14 meters.
The speed of sound is about 340 meters per
second.
Depends a little bit on whether or not
you're
at sea level, as well as the weather
conditions.
That converts to 760 miles per hour.
When I take that 0.14 meters difference in
distance between
my two ears, as well as the speed of
sound.
I come up with 0.0041 seconds time
difference for a sound located off
to the right to arrive at the right ear
before the left ear.
So, that's 0.4 milliseconds.
That's an astonishingly short amount of
time.
Notice that that is less than the duration
of a single action potential.
Action potentials last about one
millisecond and can't occur
more rapidly than about every two
milliseconds or so.
This is about a fifth of the minimum
spacing between two action potentials.
And furthermore, that's the largest
difference that can occur.
We're actually sensitive to much smaller
differences.
We're able to distinguish between a sound
that is located straight ahead, and a
sound
that is located a little bit to the right
or a little bit to the left.
So it's really pretty astonishing that our
brains are able to detect these
differences.
It is thought that the way that they do
this involves
delaying signals from one ear in
comparison to the other ear.
And having those signals converge on a
particular neuron that responds
selectively to coincident
inputs as opposed to inputs that are
coming in out of sync with each other.
So, here are two example neurons.
So, this neuron here would be receiving
input
from the left ear and from the right ear.
But the axon from the left ear and the
axon from the right ear might be of
different lengths.
Having the axon have different lengths
would then
delay how long an action potential coming
from
one side to reach that neuron as compared
to an action potential from the other
side.
So think for a moment whether or not this
neuron would be more sensitive
to a sound coming from the left or a sound
coming from the right.
Well, I'll give you the answer to that in
just a moment but consider this neuron
down here, it would have a shorter path on
this side, a longer path on this side.
So it would also serve to be sensitive to
sounds coming preferentially from one side
versus the other.
But it would be sensitive in the opposite
pattern to this neuron here.
If you think it through, this neuron is
going to
respond better when the sound arrives in
this ear first.
And the sound arrives in this ear second,
because the, the action
potential has farther to go on this side
than on this side.
This neuron would respond better when this
ear heard
the sound first and this ear heard the
sound second.
Well, interaural timing differences are
not the only game in town.
We also compare the sound loudness across
the two ears
to infer the direction that the sound is
coming from.
And here's how this works.
So if there's a sound located off to this
side, it's going to be louder in this ear.
This ear is going to be in a kind of
shadow and the
sound volume when it reaches that ear will
be a little bit quieter.
And by comparing how loud the sound is in
one ear compared to the other.
The brain can infer the direction that the
sound is coming
from, but there's a critical problem with
both of these cues,
and you might want to pause the video here
for just a
moment and see if you can guess what the
problem is.
Well the problem is called the cone of
confusion.
And that is that there are many locations
that
can produce the exact same timing and
level differences.
So, if a sound is located all the way to
the right or
all the way to the left, that will produce
the maximum level difference signal.
But, for locations that are not quite all
the way to the left or to the right, there
are going to be many locations that have
the
same difference in path length and same
head shadow effect.
So any location, on this circle, or cone,
will produce the same timing and level
difference value.
Any location on this cone,
will have the same timing and level
difference
value as any other location on that
particular cone.
This cone is different from this cone.
But any location within this cone can't be
distinguished
on the basis of timing and level
difference cues.
So in the next lecture, I'll tell you how
we solve that cone of confusion,
and do better than would be predicted
based on timing differences and level
differences alone.