y2clutch / raw_transcripts /lecture_9.txt
elyx's picture
initial commit
dae990d
All right.
Good morning.
You know, I have a mini stroke or did I
say some names outside before anyone else say some names?
Oh, no.
No.
Okay.
Maybe I just had a mini stroke.
Okay.
I'm just trying to work out what happened there.
So it's really my pleasure to raise a volume.
So.
Is that better?
I can't tell from down here.
Okay.
It's my pleasure to try and take you through this
today.
When I was a young graduate student, actually, this field
had just started.
And one of the papers I'll introduce you to.
Was a real key moment in that development.
I remember talking to one of the authors of that
paper and.
Been blown away by the idea that perhaps we could
track.
The evolution of a decision in the brain.
I think we think it's fairly commonplace to look at
this now, but back then this seemed like there's something
beyond.
Our abilities.
So in the last couple of lectures, what I hope
I hope you understand is that signals from the senses,
like the eye I communicated to the brain along parallel
pathways of showing you that these signals are represented in
early brain areas, the primary cortical areas, for example, in
the form of topographic maps of the sensory periphery.
I'm telling you also the higher brain areas are called
high brain areas seem to transform those topographic maps of
the century periphery into frames or reference frames in which
we can make actions frames that are behaviour useful more
so those now probable graphic maps.
I've also shown you to some degree that Apple Motor
areas can use these spatial representations to guide movement bands.
So what we skipped over and what is still really
one of the fundamental unknowns about neuroscience is what's in
between these two things.
I illustrated to you that there are some cells that
we will call sensory motor neurones that give both sensory
input and have motor related glands activity.
They seem to be particularly important in this process.
And in this lecture we'll go through a particular class,
those neurones that sit in this whole area of the
brain called the lateral imprint area, which we discovered a
little bit in the last lecture.
And the question really is how do we decide which
action to execute?
There are many different actions we could possibly execute.
How do we decide among these options?
What I want to try and introduce you to is
the idea that we can actually practice evolution decisions.
We still don't know how we do decide, I should
say, but we're getting closer and closer to understanding that
fundamental point about behaviour.
To try and illustrate this, we have to settle on
a on a definition of a decision that we can
actually explore experimentally.
And I'm just going to take you through that in
the next couple of slides.
There are many different ways we can think about decisions,
but we need to think about ones in which we
can try and work out what is the neural basis
of this decision.
This slide represents two possible kinds of decisions that you
might make by commonly in the top in going to
a restaurant, in this case a pizza restaurant, you look
at the menu, you're trying to work out what it
is you would like to eat.
You surveyed the different options you take into account your
previous experience and biases.
You look for evidence in the form of the different
ingredients in the in the menu.
For example, you deliberate about what it is you would
like.
I'm asking my mom, in which case you just get
the same thing every time.
You learn what's available.
Understand the differences and deliberate.
This is a decision that we could perhaps explore.
Similarly, if you're a goalkeeper in football and it's your
job to try and save a penalty, then that task
to try and stop that ball going into the goal,
by the way, it takes about 0.3 of a second
for the ball to get from the penalty spot past
the goalkeeper, but not very long at all.
Your task is to try and evaluate the sensory evidence
or maybe the pattern of steps that kick the ball
is going to is using the direction that they're coming
from.
Maybe analyse that, that that particular player before and you've
worked out that they like to kick it up in
the top right of the net.
You have to try and make a rapid a very
rapid decision.
You have to execute that decision within about 50 or
80 milliseconds of holding it very, very quickly and strongly
that the evidence you have for the hypothesis that you
are testing it quickly to make that decision.
So these are two forms of decision, the latter where
there's very little action leaping left or leaping right or
staying put is an easier one to study in the
context of operating systems and the kind of decision that
we're going to try and explore in this lecture.
So a definition of the kinds of decisions that I
would like to use is that a decision, is a
commitment to a proposition or the selection of an action,
a process that results in the overt act of choosing
based on evidence of prior knowledge and belief.
Overt is easier.
Is a decisions that we can look at objectively rather
than just subjectively.
As Jeffrey saw, one of the pioneers of this field
is puts it where choice refers directly to the final
commitment, the one among the alternative actions, decisions referred most
directly to the deliberation preceding the action.
So it's not the actual action we choose, but the
process of deliberating among the possible actions that we undertake.
Or as Golden seven.
Mike Catlin is another leader, as is his former.
First of all, decision is the deliberative process that results
in the commitment to categorical propositions that the right an
apt analogy for judge or jury that may take time
to weigh evidence for Internet of interpretations and or possible
ramifications before settling on a verdict.
In these kinds of definitions is that these decisions take
time.
You accumulate evidence and you take time to make them
be deliberate.
And the fact that we're taking time to make them
means that we can look for the signals of that
process in the brain.
If there was no time in which you needed to
make that decision, you wouldn't know what to look for
in the brain.
But it takes time to accumulate evidence, and that's something
we can can't find.
So this is one way of trying to think about
the basic components of most of these kinds of decisions.
On the top is the context in which the decision
is being made.
On the bottom is kind of the architecture of the
process that helps that decision to be made.
For example, you may need a task and motivation and
will get to a very specific task and motivation in
the moment.
You need to generate hypotheses about the world.
You need to incorporate your beliefs and prior knowledge.
That's all the context of the task.
And the context of this kind of sensory guided motor
actions and the kinds of decisions that we're going to
make require you to take some sensory input, evaluate it,
transform that sensory input into a useful form of evidence,
something that you can act upon.
You need to generate what we will call a decision
variable.
That is a point at which after which you have
committed to a particular decision.
We'll go through this in the next episode.
You need to apply that decision rule that you need
to execute the motor out.
I challenge you to describe decisions and reforms that don't
fit into this general framework.
It doesn't cover every type of decision, but most of
these things are the key component that you think, well,
one thing that shouldn't fail.
And we're going to try and fill out these boxes
in terms of specific tasks that an animal or human
might do and which they have done for the last
20 years, ad infinitum.
So what I hope we will discover in the sector
is that humans and other animals accumulate evidence for decision
over time.
That decision is made when the accumulated evidence reaches a
criterion.
Consequently, harder decisions take longer to get right.
That by adjusting bias in criterion, we can change the
process of decision making from being safe and slow growing
fast.
And I'm going to show you that there's your signatures.
That is new activity that reflects these different parts of
the process and that you see this is a key
way that evidence can be identified early in the processing
hierarchy.
And and actually, Frank, Larry, as we discover among neurones
that both respond to sensory stimuli and predict motor outputs,
that is sensory motor neurones, these are the kinds of
neurones we explored in IP.
I said to, for example, that respond in both the
sensory stimulus that was being shown to the animal and
predicted the motor action that they were got from the.
So most of this work, especially in the early stages,
has been conducted in the context of moving your eyes.
This is something we do 2 to 3 times every
second of every day that we are waiting.
We move our eyes around.
We make seconds.
These are ballistic eye movements, very rapid eye movements that
move eye around so that we can bring some kind
of growth onto the central region of our eye over
here.
And we bring it on to the central region of
the eye, because that's where the high intensity of odour
receptors are, and that's where we would like to analyse
the visual image.
So we were very good at making these eye movements.
We make them very rapidly and we make them very
frequently.
There's a lot of the brains devoted to trying to
make them in the most optimal way possible.
This is one example of a kind of pass that
a monkey might do or a human might do in
trying to search the city.
Here the ice starts off in the centre of the
image.
Their task is simply to find out, see and to
look at that to.
Now, when you look at the centre of these things
in the periphery, they're quite hard to detect.
It's very hard to tell whether something is an Al
or a T when they can be repeated in whichever
way.
And so monkeys tend to enter humans and make eye
movements to see sequentially surveil the image.
And so these little movements between each of these objects
are these two kinds.
And in this case, the monkey eventually finds that he,
in fact, actually makes an eye movement away from the
team and come back to the team.
So these are these are the kinds of movements you
make all the day when you're reading, for example, or
looking at someone's face.
My kids do it as well, and they're very good
at it just as we are.
That in itself is still a quite a hard path
to try and pick apart because the eye is moving
around all over the place.
There's many different potential stimuli that could be present.
It's still pretty tricky.
We need to find simplify that task even further.
And a lot of the field over the last ten
years is settled on this equal task and you will
encounter it a lot in the literature.
So I want to take you through the task a
little bit.
It is still a decision making process.
There will be some sensory input.
There will be a motor output and there is some
decision making process in between those two things.
However, the task in this case is going to be.
Estimate which way this field of random dots will see
them in a second reading and then make an eye
movement accordingly.
So the dots are moving to the left, making eye
movement to the left.
The dots are moving to the right, making eye movement
to the right.
That is the simplest possible task with moments involved.
So an animal or a human can be looking at
an evening monitor to see a field of dots on
a moving and then have two potential outputs.
Move left and move right.
And look at these two different dots.
And indeed, after the dance, the emotion seems has gone
away.
They're allowed to move their eyes and make that appropriate
decision.
This is perhaps the simplest task we can look at,
and it has proved enormously fruitful to understand the very
basic aspects of decision making.
So if you go back to this general context of
what a decision might look like and then we try
to put this past in that in that framework, you
get something like the of the animal, the animal or
the human.
The task is simply are the dots moving left or
right?
And the motivation for animals is reward often also the
motivation for humans.
Amazon.
Humans generate two hypotheses on the basis of the sensory
evidence hypothesis.
One dog moving that told to the moving right.
And I can bring in to this task, although we
won't discuss it equally here.
They believe some families set up example.
You could train and even or train a monkey on
a task such that the you over represent the probability
of the adults moving right and perhaps the monkey will
learn or even will learn that it's more likely to
go right.
That might be a prior belief.
That's not the general structure of these parts, but that's
something you might do to fiddle with the context and
biases, and I will bring it into a decision.
So that's the context of the past.
The decision making process is to analyse that visual motion,
which direction of the dots going.
Then transform that into a useful form of evidence generating
decision variable, apply a decision rule and then move your
eyes to the left or the right, and the path
to the next few slides is fine.
Go through this.
To be able to do that.
I need to tell you a couple of little things
about how we represent motion in the brain digital motion.
Then we're going to use this task, these dots that
you'll see moving into the next slide.
They've become very common.
The reason these dots are so common in these parts
is there are a large field of dots that can
move.
Either every dot moves to the left, for example, to
the right together.
That would we would call that a 100% coherence.
All the dots are moving in the same direction.
You can see that represented on the right here for
each of those dots as a little arrow and they're
all going in the same direction.
This is actually output Explorer.
So all adults can move together or only some of
the dots can move.
So you might vary the fraction moving in a particular
direction or if example, maybe 50% of the dots open
it up automatically.
Right.
And the other 50% adults.
Those dots are moving randomly.
And so those dots, they provide noise.
So by bearing the number of dots in a moving
in the same direction, we can vary the signal to
noise ratio with English.
This is very important.
Some decisions which are very easy to do and this
is very easy.
For example, 100% coherence are actually really hard to do
when there's only a small amount of signalling there.
So by adding noise, we can make this decision harder
and we can track things over a longer period of
time, for example, and we can ask how animals or
humans accumulate the evidence that they're about right.
Now, these dots were developed as a stimulus to explore
the responses of neurones in a very particular part of
the brain called Area MP, sometimes referred to as B
five.
This area, which was first discovered by senators that he
who is emeritus professor at UCL and at the time
I think was working over in Queen Square in the
history of neurology.
E contemporaneously with some researchers in the US in the
early 1970s discovered this tiny little part of the brain
that is several millimetres in size in monkeys where every
neurone in that part of the brain seemed to be
selected for the direction of motion of a visual stimulus.
In Sammy's work, this was in contradiction.
Its most effective area, he found, next to an area
that seemed to be selected for colour.
He was describing how different types of as different information
about the outside world had been encoded by visual cortex.
He found areas that responsive to motion, areas that were
supposed to be responsive to colour by not fluid depth
and other forms of information, which you will only learn
about later.
But the purpose here, our interest is in this area
empty.
By the way, the doublet empty stands for a middle
temporal area because on some in some monkeys, this part
of the brain is found in the middle temporal area.
In mechanics and humans.
It's found in a small suitcase inside of the brain.
You can't really see it here.
It's actually this little bit here.
This little area gets direct input from V1, the primary
visual cortex.
It is one of the most highly conserved parts of
the brain in primates.
Every single primate seems to have this part of the
cortex dedicated to being selected for visual motion.
And if you measure it from neurones in that part
of the brain, you get something like this activity shown
here.
If you're using a very strong stimulus, understand coherent dots.
The neurones are very selective for the direction and motion
of those dots.
These dashes here are meant to indicate the time of
appearances and action potential on one file of the stimulus
and can see that these neurones respond very well when
these talks are all moving together in one direction.
They are tuned for the direction of motion.
That is, even if the dots are moving in another
direction, they aren't responsive.
And they're also sensitive to the signal to noise ratio.
That is, if there are fewer dogs moving in the
same direction, they become progressively weaker.
And their response?
These are neurones in every empty.
They are based on neurones that we discover in a
second.
They don't seem to communicate much information about what an
animal would do with that signal.
They just represent a visual motion in the outside world.
We study these neurones that 15 or 20 years ourselves
in our lab.
It's an amazing part of the brain to report from.
What you will see here is actually the kinds of
stimuli that are used in the lab with different it's
quite hard to represent these moving stimuli.
So videos only really work.
One project is a little bit you need a high
frame rate presentation device to see it properly.
You see hopefully dots moving in one direction or another
direction down to left or up to the right, and
the fraction of dogs that are moving in the same
direction varies and find the file.
Well, you can.
Here is actually this is a video taken lots of
monkeys doing this task.
You can hear all the audio tones which indicate when
the start and finish is the monkey.
And in the background you hear the action potentials of
cells and area and to.
This.
So those are the action potentials that you hear about
recording played through an amplifier so the experiment can listen
to them.
That, by the way, is not available for the monkey
to do in the past.
It's in a soundproof chamber outside of this room.
So you can see that those neurones in area and
they respond very well when the when the stimulus goes
up into the right side, down to the left and
not at all when it goes down to the right,
but different neurones in area MP will prefer different motion
directions, some down to the left, some often to the
right, some often to the left.
Some down to the right, for example.
The second part of the of this task, that's the
sensory information that's coming in is representing an area MP.
It's representing the action potential that neurones in that area
produce.
And it's possible to train a monkey to do this
task to detect which or to report sorry which direction
in motion these dots are going in.
And this was something that was accomplished first in the
late 1980s and now has become very sound parts of
monkeys and humans.
And you can see here that if we put on
the x axis, the number of the fraction of Dr.
moving coherently at 1% or 10% or 100%.
This is the proportion correct on an author's choice task
where the animal has to report to the left, to
the right, example or two up to the right and
down to the left.
That as those as the number of dots moving coherently
increases, so does the fraction of times that the animal
gets its decision.
Right.
These animals are highly motivated because they are actually water
regulated and they're working for juice.
They just have very well, I've tried to do this
task many times.
I'm nowhere near as good as these monkeys.
These monkeys are getting almost 100% correct, about ten or
20%.
That's moving in the same direction.
To me, it takes 30 or 40%, 0% for.
But his monkeys are highly motivated.
So this is what we would call a psychometric function.
It simply says that the monkey gets better as the
numbers don't move in the same direction also increases.
All right.
So I thought then about the fact that this area
empty, which seems to be responsive to these dots, provides
potentially action potentials or neural activity that they may represent
the motion direction of these dots.
And we know that the monkey can actually detect which
motion direction is going in.
So then between these two things, between the sensory input
and this motor output.
So now I need to tell you that one of
the reasons we're looking at Area A is it has
a not only does it have a strong input from
the primary visual cortex, it also has a strong apple
to this whole area of the frontal cortex over the
lateral impropriety area.
And so what researchers thought back in the late 1990s,
and this is when I started to see this field.
Since the neurones in their empty seem to be encoding
information about the visual stimulus, not about monkey's behaviour.
What we should do is look at one of the
areas, the area empty sensing.
In this case there might be and see whether or
not the same as bear.
Maybe those neurones that are getting input from these neurones
in MP are actually closer to the decision making process
than those neurones in area.
So if I was to summarise the activity of neurones
in every MP, looks like then as a function of
time you get something on the left.
This is a schematic.
When the stimulus comes on the activity, those neurones increases.
And when the stimulus turns off, the activity resume decreases.
So this neurone is responsive to the visual stimulus.
And the amplitude of this on the number of action
potentials a neurone produces depends on the emotion strength that
stimulus.
So there's only a few dots moving in the right
direction.
And if you continue to produce, if there are lots
of dots moving the wrong direction, a lot of attention
to produce.
So these neurones encode both the direction of motion and
the signal strength of the incoming stimulus.
In Aria Lippi, however, he finds something quite different.
Now when the stimulus comes on, instead of an immediate
change in the activity of these neurones, you get a
slow ramping up of activity.
The slope of that ramp seems depend on how much
signal there is in the visuals inlets.
And that activity is actually sustained even after the stimulus
is going on.
So these neurones are quite different to the sensory neurones
in our MP which are providing input to them.
They're not as closely linked to the onset of this
stimulus.
Their activity persists after the offset of the stimulus.
And in between, they seem to show this ramping behaviour,
this accumulation of activity from low levels to high levels.
That depends on the signal strength, the visual signal strength.
This is the same kind of way that we've shown
other cells in the previous lectures.
And I'll go through the presentation here again and I
can shows the activity of a real neurone.
An area like a when it's recorded from in this
kind of past.
It's quite a busy slide, so I'll just take you
through its early.
First of all, the monkey in this case is looking
at a screen very much like what we saw before.
There are some dots moving on that screen and there
are a couple of choice targets left or right, for
example.
And it turns out that if you measure from neurones
an area like this, you find a region of the
visual fields where these neurones tend to respond to the
visual stimulus and also a region that these people field
where neurones will predict an upcoming movement.
In fact.
So it's half the dots.
Come on.
And then they go off.
When they go off, the monkey has to make an
argument to the left or the right reporting which motion
direction in which Lincoln.
In these pictures down the bottom here.
Each of these rows represents a single file in which
the animals doing this asked each of those dots the
time of occurrence of a single action Santa Cruz 11
year on and the average activity over many trials is
shown in it is found on the bottom.
So large fires mean more activity from the zero.
And you can see as shown in the schematic before
that has a seamless turns on.
In this case, there's no coherent motion and there's very
little response in the neurone does start to build up,
however.
And indeed that build up in activity sustained until the
moment that the animal makes that eye movement to the
left or the right.
Just before that movement is made, that activity goes away.
It's as if that activity is predicting when the movement
will occur.
So these neurones in our area, AYP, which is I
showing in the second have some sensory input, are also
predicting when an eye movement occur and indeed which location
visual space I'm moving towards.
So you can vary the signal as visual motion streamers
in the on the screen.
In this case, if you move it, for example, in
one direction, it's to say to the left.
Which would inform the animal that the eye movement should
be to the left that is away from the part
of visual space that these neurones represent.
Then you find actually that the activity of these neurones
is very low.
And indeed, when they make an eye movement, nothing much
changes.
If, on the other hand, you put a lot of
thoughts moving in that direction to the right, predicting that
the movement should go to the right, which is also
happens before this neurone.
The preferred location of the eye movement is a good
deal.
You find that the activity goes up substantially, is sustained
through the time by movement and then dies away.
So these neurones encoding three different things.
They seem to be encoding aspects of the visual stimulus,
the coherence of the dots and when and where the
eyes move.
They are sensory motor neurones like those we discussed in
Area 80 on Friday.
So this activity predicts the direction of the movement, as
I showed you here, which.
If the animal was making it immune to the right.
In this case, the activity is high because making my
move to the left, the activity is low.
The activity signals the direction of the eye movement.
It signals also the time of the current, the eye
movement.
And also the activity also depends on the actual strength
of the visual signal.
If there was a lot of speed in the same
direction, the activity is not.
The activity is lower.
So these new ones are really integrating lots of different
types of things, integrating visual sensory information and predicting where
an eye when and where an eye will move.
They are that boundary, the interface between sensation and motor
output.
Despite his own father.
The simple decision is whether I should be moved right
or left.
We'll hear his indistinguishable in his past.
But a visual motion into the Michael, where I've also
said to you that neurones in our IP may participate
in making decisions about where to move the ice.
And the logic, the reasons for making that claim that
these neurones in our area may participate in making these
decisions.
Is that they are not sensory neurones because their activity
build up slowly and is continued after the cessation of
the visual stimulus.
They also are not sensory neurones because their activity predicts
the time and direction of a subsequent eye movement.
Even when the sensory information is ambiguous.
I didn't show you here.
But those neurones are also not just motor neurones because
their activity depends on the visual stimulus.
So if example this number of dots moving in the
same direction changes the.
I give you the neurones that I have a sensory
representation.
And what I only alluded to here and haven't really
shown to you that the differences in their activity emerged
early on in the response way before my movement was
actually made.
So these two different sets of evidence that you can
see in this one task said if these neurones are
neither sensory neurones nor motor neurones, but something at the
interface between them.
And for that reason they are hypothesised to be closely
involved in the formation of decisions about where to move
the eyes because they integrate.
They say that the bridge between sensory and motor activity.
So we just come out of size and go back
to the semantic illustration of what these two areas are
providing.
In this particular past year.
The area Empty streets.
Visual Sensory Stimulus Responses to visual Motion.
I've suggested the area.
Let me instead show you something that's more related to
the behaviour output, or at least the interface between sensory
and behaviour.
So how does this fit into the kinds of parts
that we were describing for?
We can start to think about the activity in aerospace
as potentially representing the decision variables.
So.
I said that there needs to be a decision variable.
When we make a decision, we need to collapse that
decision onto something as simple, as simple enough space in
which we can make a decision, for example.
We may want to say simply that if I produced
at least five action potentials, that I am confident about
the sensory input and I would like to make the
decision to move my eyes around.
In that case, the number of action tools that I
produce is a decision variable.
It is not enough of them.
I won't make that decision.
It is too many of them.
There's more than enough of them.
I will make that decision.
The decision variable.
So we could argue that the activity of neurones in
LP is actually a decision variable itself.
And further, we could say that when we apply criteria
to that decision variable, that is a particular threshold, let's
say five action potentials.
Once I pass that bacteria, I will make that decision.
So we go back to this description of what a
decision might look like.
We have, as I said before, the task is moving
left and right some hypotheses that we generate some belief
and find knowledge that we're not really going into.
And here we have the analysis of visual motion and
a decision in the end to move the eyes left
or right.
It should be.
Another thing here is useful form of evidence is actually
the output of every empty decision.
Variable is potentially the activity of neurones in every LP,
and the decision rule is simply that when the activity
of neurones in area LP exceed a certain value, then
I move my direction in my eyes and the direction
indicated by those neurones.
So this is a simple architecture for making a very
simple decision.
But we start to learn some really interesting things from
this.
For example, in the next few slides.
What I want to show you is that the predictions
of this kind of model.
Are the simple decisions in the course of simple decisions
and hard decisions.
There are compromises between the speed and accuracy of the
decision.
I showed you this graph before it.
In the context of these tasks, monkeys and humans are
very capable of making correct decisions when the number of
dots moving in the right direction is enough and we
get it right 100% of the time, and when it's
not enough, we get it right on time, which is
chance.
And in between we have a graded performance.
What I didn't show you was that if you looked
at the reaction time and given all monkeys that it
takes the time it takes to make these decisions.
The report was moving left to right.
This also varies if the motion hearings.
It takes longer.
That's at the left.
It takes longer to make the decision when there's very
few dots moving in the right direction and takes less
time to make decision when there's a lot of moving.
And we might think that the difference between the minimum
amount of time it takes to make a response, it
might be simply in our time, it takes me to
trigger a motor action.
That difference between the minimum and maximum amount of time
make a decision for people thinking we're deliberating about what
the information is.
The evidence is that is provided by those you can
the spring.
And in the context of the model I'm showing you,
we can think of then evidence being accumulated over time.
We require a threshold to be reached, after which we'll
make the decision.
And when the sequence is moving, a lot of thoughts
are moving in the same direction.
That threshold is reached relatively quickly, or when only a
few dots are moving in the right direction, that threshold
is reached.
We slowly.
So this model, which is often called the drift diffusion
model or accumulation model or rate model, or is it
about faulty compounds or it simply predicts that hard decisions
take longer because the rate of accumulation of the decision
variable, the evidence is slower when when a stimulus has
less signal to noise ratio.
You can even step.
We can even start to dig down a bit further
into in this module about how the decisions actually might
be made.
I showed you that owns an area and keep people
in motion directions.
What?
I didn't show you.
But what I told you was that some of your
own example coercion happened to the left and some down
to the right.
Up into the right.
Down to the left, for example.
So to make this decision, what we would like to
do is compare the responses of neurones that are, say,
representing often to the right with those and in the
opposite direction.
Right versus left, for example.
So we might find you are preparing activity of neurones,
preparing right with motion activity, neurones, preparing network management.
The way to extract a decision variable, a useful form
of evidence from these neurones is simply to find a
difference in their activity.
What is one minus the other?
We just represent that with a minus sign and we
do that.
In an area of IP.
We expect that to be the difference between these neurones
activity, which initially starts off as zero stimulus and then
after stimulus turns on gradually or rapidly.
Starts to go in one direction or the other direction.
So, for example, in this case, this evidence area of
activity in area of IP will tend to go towards
evidence for a right wing motion.
So we can think of then of this area activity,
an area IP representing the accumulated evidence that is seen
as moving either right or left.
And further that we will apply criteria to that activity
generating IP such that when this activity reaches a certain
level, we will decide that the dots are moving to
the right or to the left.
And that active you accumulate over some time.
That time of accumulation will depend on the magnitude of
the evidence.
Yeah, so that's a really good question.
So the exact mechanism for how you subtract your rooms
can vary.
The simplest way to think about is you think back
a few lectures.
If you have glutamatergic outputs from some neurones and gabaergic
apples from other neurones and the Gabaergic and the glutamatergic
have different signs, one is positive, one is negative.
And so when you add those together, you actually have
a subtraction going on.
So if you inhibit activity from activity output of neurones
that are going right one direction, if you inhibit them
by the activity of neurones exerting liquid direction, you actually
have a function.
Antibody suppression.
So inhibition can do this infarction for you.
There are other ways of doing this, but that is
the most obvious way to find.
Those neurones are coming together in some form, invited together
into a light.
If some neurones are providing in addition to some neurone
defining excitation, you can subtract one from the other and
get this form of evidence.
Now the schematics are showing.
You going to show you the kind of real activity
on a trial by trial basis and every MP.
If you think about this, it's also schematic, but it's
a bit more realistic.
There's a lot of variability from moment to moment in
the activity of neurones and area and to.
So, for example, those neurones that were preferring right with
motion are no longer nice straight lines, but they will
be lines.
And you and Fred, right?
MARTIN And your friend Macklemore from Memphis.
In fact, these you still in the end get towards
the same value, but you got a lot of variance
in the game, for example, instead of having a straight
line here.
Absolutely.
Lines are important, first of all.
Now, the consequence of this noise that's happening on each
trial is variability in your firing is that sometimes you
might reach this criteria more quickly on than on other
times.
So for example, and because of that, we need to
make a decision about where we set the criterion and
what decision we're making.
It is evidence accumulates over time.
It will, in the end get to the right place.
If we make our criteria nice and high threshold, nice
and high, we will only.
Make a decision when the evidence is accumulated to a
really safe sure bet.
So we going to have a higher threshold and make
sure that we don't make the wrong decision.
If, on the other hand, we reduce our criteria, reduce
our threshold, we become sensitive to noise variability.
Some of the times that noise is fine, that variability
is fine.
So, for example, here we're still making the right decision
is going towards the right.
We're making it earlier because we're able to be more
sensitive to the early phase of the activity.
But we also occasionally make the wrong decision because actually
because that noise, that variability, the activity, the new ordinary,
empty, we're actually representing the wrong direction in motion at
that point in time.
So if we make our criteria really, really low, we're
going to be faster to make decisions because we need
less evidence to accumulate.
But we're not going to make we run the risk
of making the wrong decision.
So we transform a safe and slow decision and go
fast with every decision.
Just by simply changing the criteria which will apply to
activity of your.
The other way that we can try and change the
kind of responses we make is by adding bias to
the activity.
So, for example, we might come in with preconceptions and
don't move to the right.
We could somehow change the combinations we're making such that
the activity goes right with neurones ones closer to the
threshold that we've set.
We will then be very capable, very capable of detecting
dogs that move around very quickly.
With our bias allows us to make fast decisions.
Our it also runs the risk of making the wrong
decision.
If, for example, these double blind gear activity would have
normally ended up being a st, the left was unfortunately
to the right.
Early on in the trial.
So again, we can transform a safe and slow decision
into a risky and fast decision, this time not by
changing the criteria that we're applying to the activity, but
by changing the bias that we put into the system
in the first place.
We might call this our five beliefs, our bias, or
whatever it is.
It's it's something that we can use to manipulate the
activities.
I want to spend so.
But hopefully outline to you there is that in this
very simple framework of understanding the decision and these very
simple, neurobiological driven models of making those decisions.
We actually have some profound insight about the process of
making decisions that we can have these safe and slow,
fast and risky ones.
We've actually been able to see how neurones that bridge
between sensation and motor activity might actually help revive that
season, although we still don't know.
How.
I just want to spend a few seconds, you know,
just describing some one of the other outcomes of that
kind of framework.
And that is one of the things we want to
do when we make decisions is learn from them.
We want to make better decisions in the future.
And.
The other thing that we want to do is and
make better decisions.
We kind of know how confident we were in those
decisions.
We also want to know.
We want to associate the decisions that we make with
the presence or absence of a reward that we get
from students.
So we all know how confident we are in the
season we're making so that we can learn from those
decisions.
We want to know if those decisions led to a
reward.
Strikingly, we've been able to make some progress in that
in the last ten years.
But this accumulation model actually predicts that a wave representing
confidence in the activity of neurones.
As I said before, we can have a decision variable
here which accumulates over time until we make a decision.
But if we look at the activity of neurones in
the brain over this period of time, we also find
another feature which would be very happy to be able
to answer in this context is that.
The certainty that we have and the variability in the
activity, those neurones changes as a function of time.
So not only are we getting a change in the
mean, that is the accumulation of evidence, we're also getting
a change in the variability of the activity of those
neurones.
And the consequence of that is that early on in
the trial, for example, we have a lot of variability
and we have much less confidence in our choices.
Much more uncertainty.
Whereas later on in time we have more certainty or
more confidence.
Now, of course, the variability of different.
We can actually measure the reduction here that we are
actually more confident later on.
Like more evidence because the variability reduces and the activity
of neurones and towards the main.
We actually measure some of these things by confidence in
humans and animals.
This is one really nice example of how to find
it in a child in this case.
We can measure the confidence the animal the child has
in the decision making without even asking.
And the rate at which a child will find very
difficult to do.
This task is really straightforward and really, really elegant.
A trial has shown two boxes through which they can
put their hands and is shown that there's a toy
in one of those boxes.
They are then required to.
A delay is then interposed between being shown that and
then being exposed to the two boxes again.
And then the account is required to indicate whether by
moving the hand towards the box which of the thing
before using.
And that's a simple task.
That's the simplest open task that the to do.
It is stunning when you do this task is to
do what you see represented here on the x axis
is the memorisation.
The way the time between being shown the and being
asked to complete the task, which ranges between three and
4 seconds in this experiment.
And the Y axis here is what we would call
the persistence.
That is how long the child leaves the hand in
the box scoring for the object.
The green dot here shows how long they leave the
hand boxes going.
By the way, he's not there.
There's not public approval.
When they make the correct decision and the read points
indicate how long they chooses to go where there may
be a decision.
Now, if the child had no representation of the confidence,
they hadn't made decisions.
These values should be the same.
You should explore as long whether you made the correct
or incorrect decision, you should explore the same outcome.
As strikingly, you find that the child exposed longer when
they've made the correct decision and when they're waiting for.
This implies that the chart has a representation of the
confidence in the decision they can somehow use.
Another elegant design is in monkeys, as shown here and
there.
And I'll just take you through this briefly.
But basically that same experimental design that we saw before
we realised the left or the right is now elaborated
slightly with one little change in the experimental design.
Now instead of just having left and right pockets to
move their eyes.
There's also another target that allows a monkey to make
a sure bet.
A sure bet is a small but consistent war.
So the monkey's unsure about the decisions that they're making.
They could take the show back because I know they'll
get a small reward if they're more confident that this
isn't the right thing.
It was more like a move to the left or
the right pocket, which will give a larger reward there
some risk.
And indeed, if you ask the monkey to do this
task, you find the data.
I won't go through the data particularly here, but it
is, as you would expect, Monkey makes more choices when
it is when the signal strength is lower and therefore
he's less likely to be concerned and less sure choices.
So the signal strength is higher is therefore likely to
be important.
So this is the probably the short target.
And the probability the short target decreases with this amount
of.
Signal strength.
Strikingly, if you look at the activity of the neurones
in this area of IP actually represent whether or not
the animal will choose the short target early on in
the trial.
This is a little bit tricky, so I'm just going
to show you this and describe it to do really.
This is the task here.
During this time here, there's a period of time here.
That's when the stimulus is on.
Now the dots come on and then they turn off.
And then at some point in time after that, the
short target comes on.
And that's from Dave on this test line.
And then some time after that, the channel makes a
choice by moving their eyes.
You can ignore the street here for a moment and
just look at this bit during the presentation.
The stimulus.
It turns out if you divide those files into three
different parts, Monkey chooses the left target, choosing the right
target, which uses a short target.
The activity in this period before they even know that
the short target will be available, predicts the upcoming decision.
The implement.
This short target is only available on 57,000 unpredictable 50%
pilot.
And we did not know that that tiger will be
available when when this activity is developing.
And yet that activity sits between this activity, the left
and right eye movements, even before the animal knows the
target is available.
That is evidence of an activity representing the confidence the
animal has in the decision to bear out or.
I'm going to get this slide, but I encourage you
to write.
I read the papers.
I just want to end this by saying.
When we make a decision, we hope that that will
be the correct decision.
We hope to get reward for a couple of lecture
lectures ago, we discussed the part of the brain that
is actually important in generating rewarding signals eventual placement where.
And it turns out that that little area of the
brain provides broadcast signals about work and incorrect decisions to
the rest of the brain, including those areas like it
either involved in making these decisions.
And there's a beautiful set of data in the in
the in the technical area which shows.
When an animal is learning a task like the kind
of task of showing their.
That that the activity in the rental health area which
in start of encoding the reward that the animal gets
transitions to encoding the stimulus that will predict the reward.
And so the animal was able to use this teaching
signal from the entertainment area, its own signal to learn
how to make better decisions.
I just explain this diagram here to help you understand
what is going on.
Early on in the learning process, Dan was not provided
as it was.
It does get a reward, maybe a reward of juice,
for example.
And when that reward is provided, the activity of neurones
in the BTK monkey again increases.
The animals and learn to associate that the terms of
that reward that Jews with a previous occurrence of the
stimulus that predicts that reward.
This we were condition stimulus.
And after a long time of learning this relationship.
The animals have eaten during the VTR no longer respond
to the reward itself, but respond to the presence of
the stimulus, the conditions in this.
And indeed, if a reward is absent after presentation of
the conditions in which you see this produce and everything
approaching the EPA.
So the neurones in the mental states mental area are
also representing the outcomes of these decisions.
They're representing whether or not a stimulus will produce or
upcoming reward, and they're allowing animals to learn from that,
from that rewarding and rewarding scenario.
So what I hope I've shown you here then is
very simple decision architecture.
Face and sensory input.
Making eye movement has taught us a lot about how
to see the snake in the brain.
We've seen that some neurones that seem to sit at
the interface between sensory and motor outputs accumulate signals in
a way that is inconsistent with the idea that they
form in of themselves the activities of the client in
making this decision and that all we need to do
a set of criteria on the activity of neurones so
that we subsequently make a decision.
I've shown you that in addition seems to be representation
of confidence in the brain, a way that we can
be confident about whether or not we're making the right
decisions.
I'll show you also that in the reward circuits in
the brain, which allow us to learn about history and
experience of making those decisions.
How will these things come together?
This remains still a mystery.
How all these things are brought together, how they invade
consciousness, an awareness that only remains in these people.
Those signals are there.
This is what we've learnt over the last ten years,
and what I would find explained to you on Friday
is how those ending direct with the emotions that we
feel.
Thanks everyone.
I was.
Very.
Sure about a ton.
Of pressure placed on the monkeys by the experiment.
Oh, that's a really good question.
Right.
So.
So you could make these decisions in two context.
One is you got all the time in the world.
But make these decisions in context.
One is you have all the time in the world
and the other is you need to make it in
a set period of time.
Now, the context in this case for these animals is
provided not so much by the but sort of.
But these animals, there's two things that you might want
to try and make it fun.
First of all, the faster they make decisions, the quicker
they get.
And they figure they get to the next problem that
they have during the 7010 problem.
So that's one one plan.
The other one is that the stimulus plan.
So there's actually no further additional data from that.
So there's there's a concept that there's no additional evidence
coming in.
So you've already got all the evidence.
And if you make it faster decision, you get another
of.
So this is not like an election.
It's a what.
Is what what official news.
We don't get comparison with what I saw.
What is the model when you're on an area like
that is that when they're actually reaching a certain.
Probably about 16 foot about one in every six feet.
That is.
So whether this is some of this is certainly part
of the voting.
Record during the primaries.
What we do know is that the.
In the area like we predict.
When you.
And there is more or less in terms of the
number of actions that an optimist sits on.
Yeah, right.
And also this like I don't know how this relates
to the fact that the you're asking to have like
a preferred coin partner.
Yeah.
Sorry I struggle to work out between say so.
You're in the now and you were mysterious for a
long time when they seemed to actually.
Yeah.
So the fact that.
So when you want to do things to some kind.
Are you finding some kind of situation in which.
So it's actually.
A time.
Where the.
But then you the event.
Well.
Oh.