Okay so that covers the eye and

the retina just in a very sort of general,

preliminary way.
But of course the rest of the visual
system is just as important as the eye.
Many would say more important, I suppose.
But the argument that you'll see
emerging from this whole series
of talks is that basically the eye,
the retina, the rest of
the visual system that we're going to talk
about now are all doing the same thing.
They're all trying to get around
this fundamental problem of,
of not having the information directly
about the physical world that
we need to have to act in it.
The visual system from the retina to
the occipital cortex in the back of
the brain that I'll show you in a second
is called the primary visual pathway.
And let's take a look at that.
It's a little bit complicated to explain,
but again these are just the basics that
I think you'll need, and probably all
that you'll need to think about to
understand more or less what we're
talking about in the course.
So, imagine that this is a screen that
you are looking at in front of you and
that this x in the middle of
the screen is the point of fixation.
That is, it's the point, as the dotted
lines show, where you're converging
the two eyes to look at a particular
position of interest in the visual field.
And the reason that the left side of
the screen is colored one way and
the right side of the screen another
is to indicate by color coding that
this side of the visual field will end up
being processed in this side of the brain,
the opposite side of the brain and
this visual field will end up
being processed over here.
And there are reasons for that,
that are not terribly well understood that
have to do with the evolution of
vertebrate visual motor systems, but
this is just the way that the anatomy
of the human visual system is.
And what's very important for
you to recognize is
that it's the so-called
temporal side of the retina from this eye.
And the nasal side of the retina
from this eye that are looking at
this part of the visual field and
going to this occipital cortex,
the occipital cortex referring to the

occipital lobe in the back of the brain,

which is as we'll see in a minute, is the
locus of the cortical visual processing.
At least that's where the cortical
visual processing begins.
And the opposite side of the retina
shown here in purple on this side,
the so-called nasal side,
and on this side the, the,
the temporal side of this retina
are combining by crossing at
what's called the optic chiasm
ending up on this side of the brain.
This is complicated anatomy.
It's not entirely rational, but it's
the way we've evolved to see, as I say for
reasons that probably have to do with
primitive visual motor coordination.
But in any event, this defines
the primary visual pathway from
the retina to what's called the thalamus,
which is a nucleus in,
more or less, the middle
subcortical region of the brain.
The thalamus is a walnut-sized
anatomical entity that contains
a large number of nuclei that process
different forms of sensory information.
Including one subnucleus called
the lateral lenticular or
dorsal lateral lenticular nucleus.
That's the nucleus that's
specifically concerned with vision.
So that's the, sort of,
next way station from the retina for
the information that is coming
from the visual field and
eventually ending up for
processing in the visual cortex.
The axons from the neurons here,
just like the axons from the retina
project to the thalamus.
The axons from the thalamus are going
to project to the visual cortex in
what's called the optic radiation.
And this region of the visual cortex that
the input comes to, not input directly
from the retina, but via the thalamus
is called a primary visual cortex,
because as we'll see in a second, there
are other extrastriate regions, and for
regions we'll talk about in a minute, that
are secondary, tersiary and are higher
order regions of the cortex that
are adjacent to these primary regions.
So why is this called
the primary visual cortex?
It's called the primary visual
cortex because it's the pathway that
really concerns perception.

It's the information going from the world,

through these different stages
of the primary pathway to
the occipital cortex and
the primary visual cortex in particular.
That's generating what we see.
But as, is apparent here
there are other aspects there
are other projections from the rat lymph
to other extremely important regions.
So, [COUGH] one of these uh,regions
that's shown here in the dotted line
is to the so-called, superior colliculus,
not so-called, I mean,
that's just an anatomical name
that's been given to it for
many decades now, that's concerned
with moving the head and the eyes.
So obviously, as we said a minute ago, the
eyes have to work in coordinate fashion to
converge on a point of space that
you're interested in seeing.
And the superior colliculus is
concerned with input from the retinas
that mediate the coordination of eyes
through the visual motor system.
There's a whole nother aspect is that
we kind of glossed over before, but
that's the pupil of the eye.
That also needs to be
controlled autonomically.
And so they're additional
autonomically related nuclei here
that are controlling the,
the diameter of the pupil as you
look at a brighter scene,
the pupil contracts, as you know.
You can easily see this just
by looking into a mirror and
shining a light in your eye.
And that information, again, is critically
provided by the, the, the retinas.
But it doesn't involve perception,
it involves the autonomic system in
control of the diameter of the pupil so
that you adjust the amount of
light that's coming into the eye,
depending on the ambient illumination that
is being provided by the, the [INAUDIBLE].
So that's the primary visual pathway.
Thalamus, retinal to thalamus to
visual cortex primary visual cortex.
And these other elements
are enormously important.
The superior colliculus, and
the control of pupillary diameter.
And other things that have to do with
phenomena, like circadian rhythms.
The adaptation of your
physiology to the light,

that's changing during

the course of the day.
Their whole.
Rhythms that are controlled again
in a very sophisticated way
from light coming into the retina and
a special class of ganglion cells
in the retina that deals with that,
that we're not going to talk about.
The point being that there are many
pathways, or at least several pathways
coming from the retina that have to do
with things other than our perception,
and we're going to focus of course, in the
courses you might imagine on perception.
And that means that we're going to be
talking effectively about
the primary visual pathway.
The station that again,
the respective reception people focus on
is the primary visual cortex
in the occipital lobe.
So here is the brain, and
we're looking at it from the medial side.
Let me show you a brain here.
So this is actually,
this is a, a human brain.
And if you look at it from the lateral
side this is the appearance you get.
If you divide the brain in the middle,
which is what's been done here.
Separate the two hemispheres,
then you can look at the hemisphere
from the medial side.
And that's what you see in this diagram.
This is the front of the brain and
this is the back of the brain.
And the occipital lobe, as I said,
is in the back of the brain.
And the region of the occipital lobe,
and most of the occipital lobe is
concerned with vision, but the region
that is specifically dedicated to
the primary visual cortex is in
the so-called calchrin solcus.
So this is a blow up of this
back of the brain region.
And the color coding is just to
allow you do sort of identify and
relate this to the visual field.
Remember those diagrams that we saw on
the preceding slide that have to do with
the point at which you're fixating on, and
points that are increasingly eccentric,
from that point,
that are increasingly far away in visual
space from that point, in a lateral sense.
So, this is a blow up of
the regions that we refer to as
the primary visual cortex, V1, and we'll

see that there is a V2 and V3 and so on.

We'll talk about that in a minute.
But V1 is easily identified.
This is a section that
would be cut more or
less in this dimension,
so it's a cross section.
And you can see, and
people have seen for many decades,
that this region of the brain is
identified because there is a stripe,
which is why another name for
it is the striate cortex.
And the striate cortex is
again in the human brain.
Defines the, the limits of
the primary visual cortex and that's
what allows one to draw diagram like this,
this sharp anatomical distinction.
Who's basis is really not
terribly well understood.
It comes from a predominance of,
fatty substance that envelops axons
in a particular layer of the brain, that's
what causes this different appearance,
that differentiates it from the extra
striate regions here, and here,
but That's just an anatomical landmark,
that's helpful.
Now [COUGH] excuse me what's shown
down below is a monkey brain.
So you can see that the monkey brain
is simpler than the human brain here,
the brain I'll actually just showed you.
And the point of this, is that we
can't do this in a human animal, but
we can do it in an experimental animal.
You could show a stimulus like this for
a long period of time and
ask, how is the retina
related through the thalamus.
Where, again there is strict
maintenance in neighbor relationships.
I didn't mention that, but
that's important to recognize.
How well is that neighbor relationship in
the retina maintained in the thalamus?
Very well physiologists have shown.
How well is that maintained in the cortex?
And this is a compelling indication that
is also very well maintained because you
can see that the image that's shown for
a long period of time on the retina by
metabolic technique that allows you to see
the regions of the primary visual cortex.
So all of this is primary visual
cortex in a Rhesus monkey.
You can see that this pattern is very
well preserved here, meaning that
the neighbor relationships on the retina

are preserved through the thalamus and

indeed preserved in the visual cortex,
so the primary visual cortex.
And that's an impressive
demonstration of retinatophy so
called, that is the hallmark of
the whole primary visual system.
Why that's the case again,
we'll come to later.
So, the next question I think
that one needs to ask is,
well, what about the rest of
the visual system in the cortex?
So again, this is what we
were looking at a minute ago.
This is the brain from the scene
with the hemisphere split apart.
Front of the brain here,
back of the brain here.
Everything behind this dotted
line is the occipital cortex.
And you can see that the one, the primary
visual cortex that we've been talking
about in the last slide is not, of course,
the only visual region in the brain.
There are many other
components that have names.
V2.
V3, V4.
They're very hard to see when the brain
is in its complete configuration
as it is here with these gyri
of the peaks of the valleys and
the south side which are the valleys
in between the gyri.
Is very hard to see in and
appreciate what's really there in terms
of the topographical relationships
between these areas V1, V2 and so on.
So very importantly,
in the late 1990s Anders Dale,
Bruce Fischl, and others working at
the Massachusetts General Center for
Imaging in Boston came up with a very
clever way of inflating the brain,
computationally of course not with air.
But taking a brain that's
imaged like this with
lets say electromagnetic imaging
that shows up these regions.
Inflating it so that [COUGH]
you get rid of the valleys and
and peaks, the sulci and, and, gyri.
And effectively you flatten the brain so
that these regions become
much more visible and
can be defined as they are here, seen
as they are here in a much clearer way.
So the green as we started with
the primary visual cortex here.

And when the brain is inflated it ends

up being seen in a region like this.
And unlike this complicated
picture here you can much
more clearly see how V1 is related to V2,
V3 V4, and so on.
And these are additional areas
here that are called MT and
MST that I'll say a little
bit more about in a second.
So, what are these other areas doing?
Well again, as so
many things in vision, ones not
entirely sure what their functions are.
But it's clear enough that
the information that's coming into V1.
And that these so called higher
order area, have specific functions.
And the ones that are best
understood are V4, which seems to be
particularly interested in color,
not exclusively by any means.
None of these are exclusive
to any particular function.
And these areas MT and MST, which seem to
be particularly concerned with emotion.
So it's possible to,
to at least some degree, to indicate
what these areas are, are doing.
But not to a degree that again, we need
to worry too much about in this course.
Other than to realize they're there.
They're doing something.
It's not as well understood what
that something is as one would hope.
But the focus for us is going to be on the
primary visual cortex which is the entry
point to all of these higher regions and
much more is known not
surprisingly about V1, the primary visual
cortex, than the rest of the brain.
So the last thing I want to mention
very briefly is the concept of
neuronal receptive fields.
And we'll come back to this later and
talk about it in more detail.
I put this picture is,
and I'll introduce you to the characters
in it in the third module.
But the concept of neuronal receptive
fields is just the idea that if you
record from a neuron in the visual
cortex there was an area in space,
imagine this as a screen
in front of you or
an experimental animal more alike that's
being recorded from a single cell.
There's a place in the space
on the screen that
cells responsive to and they're also

characteristics of the stimuli at

that point in space that the cell's
going to be responsive to.
Those, those are called
visualceptive field properties.
And again, we'll come back and
talk more about this later, I think.
I don't want to overburden you with
the detailed information about
this [COUGH] today, but
the idea of receptive fields is
going to be important in what we talk
about and I at least wanted to mention it.
So that's I think,
more than enough for today and
let me just summarize the main points.
So, again we've talked
about a lot of things, but
I started with the idea that
visual stimuli are creations of
evolved biology and
not images of physical reality.
Images don't exist in the world.
They only exist when your eye brings
them into existence by the cornea,
the evolution of the cornea,
the evolution of the lens,
the evolution of retina photo receptors
laid out in the two dimensional way.
Or the artifact that people
have made to achieve the same
general purpose of making an image
with a camera and film chemistry.
And this fact is going to be critical for
understanding perception.
Again as I've said in the first module
the course is really focused on perception
and it's this fact of
the importance of the need
to create images by biological evolution
that's going to be critical for
us to understand what
perception is all about.
For the purposes of the course,
as I've said a couple of times already,
we don't need to know
an enormous amount of detail.
You probably think that I've
given you too much detail
already about the visual system.
But this is really a, a minimum.
But the primary visual
pathway is the route
to perception from stimuli through
the retina, through the thalamus,
to primary visual cortex and
beyond extrastriate visual cortex.
And so we just need to bare in
mind some of those basic bits of
information that I have given you.

And the elements that,

again we need to know something about,
our eye, retina,
the lateral geniculate nucleus,
and the thalamus and
the visual cortex in the occipital lobe.
V1 in particular again V1 is
synonymous with primary visual cortex,
V2 secondary visual cortex,
V3 tertially that, that, that's role.
So next time we're going to switch
gears [COUGH] quite a bit and
I'm going to tell you something
about the history of vision.
Basically how people have
put all this material that
I've been telling you
about today together.
How have they thought about vision,
how they conceived it's purpose,
it's operating principles.
And although people just don't know
the answers to those broad questions yet,
it certainly makes sense to kind of
see where the landscape lies today.
And what the major theories and, and
concepts of vision are today and
what direction they're going in.