Okay, let's turn now to

the organization of the visual system.

I mentioned a couple times
already that we don't need to
know a great deal about
the enormous amount of
information that's known anatomically and
electrophysiologically.
About the mammalian visual system in
all kinds of species, including humans.
And we have to, of course, know something.
And let's begin by discussing the eye and
the retina.
Here's an image of the eye, of course.
And I mentioned before that it's
the cornea that's doing the major
part of the refraction of the light that's
coming from the environment to the eye.
And that, that refraction is
refined by the lens which is.
Neurally controlled by part of
the autonomic system that makes it
thinner or fatter.
And in that way,
by adjusting the thickness of the lens,
can precisely focus on the retina.
The rays of light that
are coming into the eye,
depending on whether we're
looking at something that's.
In the middle distance at the far,
at a far distance and the near distance.
The receptors, of course,
are in the retina.
And I think what's important to
notice from a picture like this,
as well, is that there's a central
spot on the retina called the fovea.
That's F-O-V-E-A.
As you can see here.
And it's that point on the retina,
and you'll see this in a minute, in more
detail, it's that point on the retina,
that's more in line with whatever you're
looking at with a fixation point.
The point at which you are focusing your
attention, and at which the two eyes are.
By, again, the motions of the extra
ocular muscles made to converge upon.
But those are the elements
of the of the eye.
And of, of course they're critical
in making an image on the retina,
which is what we said
initiates the process.
Even though, again, I.
Underscore that the image that's on
the retina is not really an image of
the objects in the external world.
Images don't exist in the external world,

images are something that

are biologically created.
So lets take a closer look at [COUGH]
how the retina would look if You
were to look at it using an opthalmoscope
as a, as a, opthalmologists do,
look through the pupil and image directly
the retina at the back of the eye.
So, there are a number of obvious
features of the, of the retina.
One is the so called optic disk or
optic papilla.
This is the point at which the,
all of the nerves exiting the eye, or
the axons exiting the eye, are collected
together to form the optic nerve.
This is the fovea.
And the area around it is called
the macula ludea because it
has a somewhat different appearance,
as you can.
See here so
it is this point that you are aligning.
Both eyes on with the point
of fixation that your eyes
are moving together to see in space.
Obviously you can see the blood vessels
that's important for a lot of things.
Particularly pathologies like macular
degeneration a common cause of
of blindness.
So here is again, the eye, retina,
and this little box is blown
up here in diagrammatic fashion to show
you the elements that make up the retina.
And, the first thing you'll notice,
a very sort of strange counter
intuitive phenomenon that.
Light is coming to the eye
from this direction,
so the lens is out here somewhere.
Light's coming in this direction,
falling on the, on the retina.
And, what's odd about this is that
the light comes through all of
the elements that we'll
talk about in a second.
Before it falls on the,
on the receptor cells as such here
will be, here are the receptors.
Now why should that be?
There's a good reason for
this and that is rods and
cones the two types of photo receptors
that are in the retina are filled with
discs that have the pigments in
them that absorb the photons.
This is the way vision works, light comes
in, in the form of photons, it's absorbed
by the photo pigments,

that are either in rods, or cones.

And those are metabolically
turning over at a higher rate.
And it's this pigment epithelium
that absorbs the degraded atritus
of the disc in the rods and the cones,
and absent that connection with
the pigment epithelium, which is
the back of the eye as it needs to be.
Without that turning over you can't
maintain the physiological integrity of
the rods and cones.
So basically the reason for
having the receptors at
the back is because they have to be
adjacent to the pigment epithelium here.
So what is all the rest of this stuff.
This is the receptors the rods and cones
we'll talk more about this in a second.
These are horizontal cells and
amacrine cells, and bipolar cells,
are the three types in
the layers of the retina
that intercede between the light coming
in and the, and the photoreceptors.
And they're doing an immense of
amount of processing that the eye in,
in it's retinal component is
really part of a brain and
the logically it's not a peripheral
element except at it's
physical location it is neurologically
a derivative of the brain itself.
It's a piece of the brain and
these elements the horizontal cells,
the bipolar cells or I should say the
bipolar cells and the horizontal cells.
The anacrhin cells are all processing
the activity that's generated in photo,
photo receptors in ways that
are again provide some of
that detail we're not going to talk about
but still have many mysteries about.
What they are actually doing,
before they are led to the Ganging
cell which is the output cell, right?
The Ganging cell is the output
cells of the retina, and
is their axons that are going
to form the op, optic nerve.
And as you see they are running At
the surface of the retina where the light
is passing through all these elements and
even though these are degrading the light
to some degree, this is the way the retina
actually turns out to be working.
Now let's talk in more, or
detail about what's an absolutely
critical factor that distingushes, and,
sort of, answers the question of why we

have rods and cones in the first place.

We are diurnal animals,
and most mammals are.
Meaning that we operate in the daylight,
and sleep at night, and.
That's true of, of many species.
Of course, there are species
that are nocturnal as, as well.
But [COUGH] either way we have to
have receptors that are sensitive to
very low amounts of light at night.
And to orders of magnitude
the more elimination.
In the day time and along this axis, is
part of the luminance that we're capable
of seeing and you can see, again that
this varies over many order of magnitude.
12, 13 orders of magnitude
from very dimmest light that
we're capable of perceiving; that
is the threshold of human vision.
And, at the other end of this ability
to see light, for human beings, we,
are many orders of magnitude, witnessing
and responding to many orders of
magnitude, of more intense illumination,
and it is the cones that are.
Responsive to, as you can see here in
this bar that indicates the range of cone
functions that are indicating responses
to relatively high levels of light, and
it's the rods that are functioning
at low levels of light and
enabling us to see reasonably well in
starlight or moonlight or basically.
At night each of these
ranges has a different name.
It's not too important, but
in very low levels we discuss this in
terms of scotopic vision in
the middle range, mesopic.
And where the cones are operative,
in the range in which cones are operative,
we talk about that as the photopic range.
So again, there's an absolute threshold
of light that is remarkably low.
I mean the rods are extremely, well the
cones are too, they're both very efficient
responders to photon by absorbing photons
in the disc that I mentioned a minute ago.
The fig, photopigments
are enormously sensitive and.
Competent in, in absorbing photons but
the sensitivity
of codes does not do any difference in the
absorption ability of rods versus cones,
it's due to the fact that a whole
lot of receptors, rod receptors.
Converge on ganglion cells through
the retinal processing system

that I mentioned a minute ago.

And it's that convergence that
allows them to respond to just a,
about a hundred photons per
millimeter squared on the retina.
Is the is the threshold of human vision.
But the, but
the trade off that one's making in that,
is that the the resolution when you have
photons that are being absorbed
very efficiently by rods.
But the sensitivity is
coming from the convergence.
A lot of those rods onto
basically a single ganglion cell.
You are degrading the resolution.
Why?
Because you're integrating over a whole
millimeter onto a ganglion cell and
the information that passes centrally
is limited by that degree convergence.
The cones, on the other hand, which are
again, equally have an equal ability to
respond to the absorption of photons,
but they are basically organized in
the retina in a one-on-one fashion so
that their resolution is much higher.
One cone goes to one bipolar cell,
and the lateral integration that
the amicrhome cells, and horizontal
cells are doing that were not really.
Discussing here, means that
the pixiletion provided by cones,
is much, much higher,
than than that provided by rods.
And the out come of that is,
is the enormously different
sensitivity we have to light, but also
the difference in resolution that we have.
Whether we're looking at the region
of the retina that is dense in cones.
Which is the phobia or the rest of
the retina which has a much higher
level of rods and
a much lower level of cones.
So again,
let's go back to this picture and
remember that the point that we aligned.
With what we want to see in detail,
the fovea, which I indicated here in
this picture of the retina as you'd
see it with an ophthalmoscope.
If we look a that, in, and this is of
course a diagramatic illustration here,
but if we look at that, in a picture like
this, looking at the density of cones,
each one of these little elements
here is representing a cone.
The density of cones is enormously high
in the phobia allowing and if you want

to think about it in terms of

pixelation much greater resolution
of the image in the phobia than outside
the phobia and as you move away from
the phobia the density of cones again
these green things that are also bigger.
Outside of the fovea becomes less,
and the density of the rods increases.
There are actually no rods at
all in the middle of the fovea.
And as you move more peripherally,
as in this diagram, you really have,
end up with very few cones and quite
a large number of rods, proportionally.
And that means that your resolution
peripherally is much less
than your resolution centrally.
And you could demonstrate that I mean if
you look at a point on a piece of paper,
and make marks away from that
point on the piece of paper or
just look at line of text in,
in a book is an easy way to do it.
If you're looking at a particular word,
ask yourself how many words you
could identify to the right or
to the left of the point of the word that
you, to the word that you're fixating on.
And the answer is very few,
very quickly, your resolution falls and
you can't really identify
what the word is.
Just a few visual degrees away
from the point of fixation.
Let's look for the first time a little
bit more at the structure of the rods
and cones.
They have different structures
which give them their names.
The rods being, well, surprise, surprise,
more rod like in their appearance.
And the cones having a more conical shape.
And these are blowups of the rods and
cones.
And these discs that you see here,
are the discs that I mentioned before
that are the locusts of the photo
pigments that are actually observing.
With the photons and pigment epithelium
is up here and is essential for
the metabolism of the disc which are
turning over quite rapidly over a period
of days and so
there's a structural difference but
the key point here that you
can see is the presence.
In both rods and cones of these
discs of photopigment which is where
the absorption is really taking place.
So, this is a diagram,

here, of the distribution of rods and

cones.
This point of indicated by the zero
here is the middle of the phobia.
And as you can see.
This axis is the density of the
photoreceptor elements of in the middle
of the phobia, the density of cones
indicated in green, is extremely high,
this acts as indicates in
thousands the number of receptors.
In the area and you can see there's an
enormous density of cones in the middle of
the fovea and they fall off very rapidly,
so that just a few degrees away from
the center of the fovea the rods rise so
that the sensitivity to light.
Is much greater away from
the Here is a blow up
of a piece of retina to show you
the phobia little bit better.
And what's important here, again, is
the recognition that the other elements.
In the other layers of the eye,
are diminished in the [INAUDIBLE],
again accentuating the ability of
light to actually activate and
provide the sharpest possible vision
in this central region of the phobia.
But what this means, and
it's why this picture to the right
is I've included on this slide.
What this means is if you
want to see something sharply,
you have to move your eyes,
align your eyes.
So that whatever it is you want to
see the face on this girl.
Let's say you want to look at her,
the tip of her nose.
Is precisely aligned with
this central region that has
the greatest sensitivity because
of the high density cones there.
And the consequence of that is that,
not for all mammals, but
for humans and other mammals that
are like us, we need to move our eyes.
So that we align continually
the center of the fovea with what it
is that we want to look
at in high resolution.
So, this is work done by a Russian
physiologist named Alfred Yarbus.
I should have put his name down here.
But he was the first to use
an eye tracking system in
the 1950s to ask where is it that we look
when we just naturally inspect a scene.
So he would present observers.

With this eye tracking apparatus in place,

different kinds of scenes,
in this case the face of a young girl,
and ask over a period of several minutes.
Where were the eyes actually looking?
So each one of these dark centers here
represents a place where
the eyes were held for a period.
Longer than the intervening spaces.
And these lighter lines
connecting the black dots,
the black dots being the point of
fixation or the movements that are called
saccades that are moving the eyes between
one locus of interest and another.
And you can see that of course,
we don't inspect everything,
as you might expect, in a uniform fashion.
We focus on information bearing regions
of any scene, and when you're looking
at a face, the information regions
that are most important are the eyes,
and the mouth, and distinctions like this
region that identifies the character
of this girl as having a part in
the middle of her hair some contrast here.
But this is a fascinating exercise
that's widely used today by
people interested in making advertisements
that are maximally appearing, appealing.
And many many other scientific reasons for
using eye trackers to see what it is
that people are, are, are looking at.
And it's these saccadic eye movements
I should write that down again,
saccadic eye movements that
are continually changing at our
Focus bringing the phobia onto what it
is that we want to see in in detail.