|
Nothing like working with your kids at 9:00 in the |
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|
morning. |
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|
Can you hear me up the back there? |
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|
Is it being amplified? |
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|
Okay. |
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|
Thank you. |
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|
So in the last lecture, I tried to help you |
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|
understand convey to explain to myself that the input to |
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|
the brain from the senses is this array of signals |
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|
carried by nerve cells that represent something about the outside |
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|
world. |
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|
But that array of signals coming in through the sensory |
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|
nerves is carried, is constructed over a series of parallel |
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|
pathways, each of which carries slightly different types of information |
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|
about the outside world. |
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|
And that when you get to the cortex, these pathways |
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|
are then converge and build maps, topographic maps of the |
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|
sensory periphery. |
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|
That might be a topographic map of the retina, a |
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|
topographic map of the skin of the cochlea, etc.. |
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|
And what I hope that I got for you as |
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|
well is that those maps are distorted and distorted by |
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|
the density receptors in different parts of the skin or |
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|
the eye. |
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|
For example, those maps can be plastic, although we don't |
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|
know exactly how much and when. |
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|
Those maps aren't. |
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|
Incredibly useful for the in the world, they're very good |
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|
at representing what's happening on the surface of our body. |
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|
But if we want to make movements through the world, |
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|
then we have to do something else. |
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|
We have to transform those maps into something that is |
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|
in a coordinated frame, a frame of reference that is |
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|
behaviourally useful. |
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|
And the structure of this lecture is to try and |
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|
take you through some of the folks in the brain. |
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|
So some of the mechanisms that seem to be at |
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|
least the starting point of constructing those frames of reference. |
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|
That. |
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|
I want to reinforce in this lecture as well. |
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|
That. |
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|
Interest in the signals that come through the sensory periphery |
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|
of the cortex are carried in parallel. |
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|
These reference frames are also constructed in parallel. |
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|
We build many different reference frames at the same time |
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|
in the brain. |
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|
And what we seem to do is to try to |
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|
use a particular reference frame when we need to accomplish |
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|
a task like picking up the phone. |
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|
In that case, I might use a reference frame which |
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|
is centred on my hand because that's where the most |
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|
important part is. |
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|
So I'm going to take you through those reference frames |
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|
in the first part of this lecture and then I'm |
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|
going to use those to provide a framework to try |
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|
and understand potentially what the mechanisms are for a new |
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|
or interesting type of nerve nerve circuit in the brain, |
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|
which is that from mirror neurones, which many, many of |
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|
you may have heard about. |
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|
Now, I said to you a couple of lectures, Alexa, |
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|
ago that these lectures were designed when we were in |
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|
the pandemic and I was pre-recording them, and I'm struggling |
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|
to work out the exact pacing, and I discovered that |
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|
to my course this week. |
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|
I've what I've done is there's a section of the |
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|
notes in your slides called Controlling Movements, which I'm going |
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|
to omit from today's lecture so that we can discuss |
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|
these more interesting aspects of the brain. |
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|
I have uploaded a pre-recorded version of that sections about |
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|
10 minutes long and it's accessible for me on Moodle. |
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|
Simon and I'd like you to have a look at |
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|
it when you have a chance. |
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|
It won't be necessary for understanding the rest of the |
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|
content of this lecture, although there are a couple of |
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|
subtleties that may be apparent. |
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|
I want to be the performance of the U.S.. |
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|
Oops. |
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|
I just want to start the selection with with a |
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|
recording that I obtained some years ago from sources I |
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|
can't remember. |
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|
There's one particular reason which I want to show you |
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|
this recording, and I'll tell you that at the end |
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|
of it. |
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|
But it also introduces to the idea of spatial neglect. |
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|
Has anyone here heard of spatial neglect before? |
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|
Anyone? |
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|
Put your hand up. |
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|
Oh, in that case, you prepare for your mind to |
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|
be blown because spatial neglect is one of the most |
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|
|
interesting unsolved mysteries of brain function. |
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|
So this video, this first video should help introduce you |
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|
to it, and then we'll discuss a little bit about |
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|
that and the mechanism why that occurs. |
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|
To be to the point of this of this the |
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|
first part, you turn to the benefit of the doubt. |
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|
Frequently you start to look at all of the actual |
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|
numbers. |
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|
Positional alignment serve the purpose of this guide to address |
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|
those with respect to state of excellence. |
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|
Hold accountable for their support for the weakest example of |
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|
the various forms of principle over interesting example. |
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|
Just the complication of that simplistic classification to perform this |
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|
simple graph, for example, statements that not appear for the |
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|
first time to read the code, to describe the patient, |
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|
etc. to make sense of this cause some features of |
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|
the text, but contextually so. |
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|
Another way of testing the directive to ask the patient |
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|
to open page because he came over to Iraq and |
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|
the patient neglected to use the right side of the |
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|
page only occasionally some consideration of extended meditation or whether |
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|
|
is something in the collection. |
|
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|
So this is not behaviour so cheerful coffee perception whether |
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|
they succeed in school. |
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|
But of course this difficult because because of excessive force |
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|
|
mistakes exceed the features which they create are not really |
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|
active. |
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|
However, you can get something set up and use that |
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|
can be completed in subject. |
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|
In this case the patient of social with a reserve |
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|
of the device. |
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|
Stick with the public, look at the same table for |
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|
the right side, etc. but for the patient safety and |
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|
|
security of the same patient Refugee Convention. |
|
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|
These findings, together with the description the patient suggests that |
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|
medical detectives actually complete the sight of a doctor, at |
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|
|
least a photo with ways for them to gather, to |
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|
look at symmetrically to take. |
|
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|
Okay, So I want to share that video, particularly for |
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|
|
the verbal description from the patient who has neglected completing |
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|
those letters on the page. |
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|
And you can sense from from that description how automatic |
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|
recollect or her description of the scene is. |
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|
It's not like she's struggling through all that, something we |
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|
think that she does perceive through the letters that she |
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|
is aware of. |
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|
So this neglect is a really strange and powerful thing. |
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|
It's very rare. |
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|
It seems to entail the distortion of a perceived space, |
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|
|
and it seems to, although not described particularly here, it |
|
|
|
can actually occur in more than one of different reference |
|
|
|
frames. |
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|
Two of them were were illustrated here, those reference frames, |
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|
one with the left side of visual space, and the |
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|
other was the left side of an object. |
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|
I'll get back to the second one in a minute. |
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|
And in fact, it's a very complex film. |
|
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|
We don't understand it very well. |
|
|
|
A lot of this work and some of these references |
|
|
|
in the slides actually come from colleagues at the Institute |
|
|
|
of Cognitive Neuroscience, which is about 120 metres that way |
|
|
|
in Queens Square particularly. |
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|
Don't drive it. |
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|
It's now unfortunately dead. |
|
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|
But a lot of the descriptions came from studying patients |
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|
|
with particular types of lesions that we'll get to in |
|
|
|
a second. |
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|
This is a very automatic absence of awareness of a |
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|
|
part of your visual field or part of an object |
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|
|
or part of something. |
|
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|
It really suggests that when we construct and model our |
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|
|
vision now interpretation of the outside world, we're using different |
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|
|
types of representations to do that. |
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|
And depending on whether or not they are all intact, |
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|
|
we have the capacity to access an appropriate one for |
|
|
|
the task at hand. |
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|
So as was alluded to in that description, neglect is |
|
|
|
not simply an absence of awareness, at least not in |
|
|
|
some simple way of being blind. |
|
|
|
So, for example, these are two particular examples of how |
|
|
|
you might test neglect in some patients, and in this |
|
|
|
case, a person, a patient with a left hemisphere stroke |
|
|
|
was asked across that sort. |
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|
I think there should be right hemisphere stroke should actually |
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|
cross out the components of a navel figure. |
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|
And they both figure as these figures which have a |
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|
structure but are actually made up of themselves of little |
|
|
|
structures, in this case, letters like A's. |
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|
And so the patient in this case crosses them all |
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|
the right hand side of the figure. |
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|
However, when when asked to describe the figure can actually |
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|
|
say it's a square or a circle. |
|
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|
So there's some strange things going on in this neglect. |
|
|
|
And similarly, down here on the on the bottom, this |
|
|
|
is a really lovely experiment from John Driver, where they |
|
|
|
did something akin to what you see in the in |
|
|
|
the video. |
|
|
|
Here's what the patient is asked to to draw. |
|
|
|
You can see that when asked to draw a right |
|
|
|
hand side cap, they can reproduce that fairly well. |
|
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|
However, when I draw a left hand side, a very |
|
|
|
poor reproduction of that cat, this is a very interesting |
|
|
|
way of asking the question Is it part of the |
|
|
|
left visual field or left hemisphere part of my vision, |
|
|
|
or is it the left part of an object? |
|
|
|
Very simple test. |
|
|
|
Ingenious. |
|
|
|
To come up with this, ask the patient simply to |
|
|
|
draw. |
|
|
|
We draw each of these two identical objects and you |
|
|
|
can see that in both cases, the left hand side |
|
|
|
of the object is emitted, not the left hand part |
|
|
|
of the visual field. |
|
|
|
So when the object is tilted to the right, it's |
|
|
|
still the left hand side of the object that is |
|
|
|
emitted. |
|
|
|
So neglect is a really strange and worrying for the |
|
|
|
patient, but intriguing for the experiment. |
|
|
|
A form of brain damage. |
|
|
|
And a lot of that research still goes on at |
|
|
|
the Institute of Neurology and the Institute of Cognitive Neuroscience |
|
|
|
just down the road. |
|
|
|
So what does neglect tell us about representation of the |
|
|
|
world in our brain? |
|
|
|
As I said, neglect entails an inability to perceive the |
|
|
|
relationships of things within a particular frame of reference, a |
|
|
|
part of the visual field upon the object and other |
|
|
|
ones that I haven't gone through here. |
|
|
|
Depending on the spread and the focus of the brain, |
|
|
|
there is the damage the affected frame of reference can |
|
|
|
be. |
|
|
|
Egocentric, olive centric, extra or personal. |
|
|
|
And I hope that you understand each of those terms |
|
|
|
by the end of this lecture. |
|
|
|
It suggests that there isn't one single spatial reference frame |
|
|
|
in which the world is viewed through which we see |
|
|
|
the world. |
|
|
|
Instead, there are several, each with their own neural representation, |
|
|
|
each of which can be deployed depending on the task |
|
|
|
at hand. |
|
|
|
And we expect that it should be possible to identify |
|
|
|
the different frames of reference in the brain. |
|
|
|
And I should note here, by the way, that most |
|
|
|
people that show neglect also show other forms of deficits. |
|
|
|
And the reason for that, generally speaking, is that the |
|
|
|
brain damage that leads to neglect often spreads to other |
|
|
|
areas and other functions. |
|
|
|
So to understand then what we're gonna be talking about, |
|
|
|
I need to describe to you the different types of |
|
|
|
frames of reference that we can describe. |
|
|
|
And this picture summarises them. |
|
|
|
The two classes are really egocentric for Ellis and egocentric. |
|
|
|
It's very easy to think about. |
|
|
|
It's just with respect to the ego, to the body. |
|
|
|
So, for example, with respect my body, this is my |
|
|
|
right and this is my left. |
|
|
|
If I turn around, that's to my right and that's |
|
|
|
to my left hand to my eye. |
|
|
|
If I'm looking at you, then that's part of the |
|
|
|
visual world in my right, part of the world on |
|
|
|
my left foot. |
|
|
|
If I look over there, there's two parts of visual |
|
|
|
world to my right and my left. |
|
|
|
But who was in the left field now is on |
|
|
|
the right hand side of my. |
|
|
|
That's an egocentric form of representation, something that's with respect |
|
|
|
to my body. |
|
|
|
It could be my head, my retina. |
|
|
|
It could also be other forms that discover in a |
|
|
|
second. |
|
|
|
The other major class of frames of reference is our |
|
|
|
essential world centric spaces. |
|
|
|
So, for example, a GPS gives you coordinates in north, |
|
|
|
south, east, west. |
|
|
|
That's a world centre in space. |
|
|
|
It doesn't depend on the direction I'm facing. |
|
|
|
Again, if we go right here, this is my right. |
|
|
|
That's my left I'm facing. |
|
|
|
So it's my right and it's my left. |
|
|
|
I'm facing east now, So although east and south have |
|
|
|
not I worked with and south have not changed in |
|
|
|
the other central continents, by my figures, in accordance has |
|
|
|
changed. |
|
|
|
So the Alessandra coordinates base that will based or even |
|
|
|
sometimes object based reference frames. |
|
|
|
These are things that are independent of our and our |
|
|
|
own body position and depend only on the structure of |
|
|
|
the world outside. |
|
|
|
It turns out that that's the most stable representation to |
|
|
|
use because that doesn't depend on where I'm moving. |
|
|
|
But we do choose when we describe something in the |
|
|
|
world to other people, we often choose to use egocentric |
|
|
|
reference frames. |
|
|
|
So for example, if I was to describe how to |
|
|
|
get out of the building, in my case, I got |
|
|
|
that turn left, turn left again, turn right and then |
|
|
|
go straight ahead. |
|
|
|
That's an egocentric description. |
|
|
|
I could say that third, south and east and slightly |
|
|
|
north and east again. |
|
|
|
And that would be another centric description of the same |
|
|
|
towns. |
|
|
|
I won't describe much about either centric, but I will |
|
|
|
just introduce you to the idea that these two things |
|
|
|
can be distinguished in development and in perception. |
|
|
|
These really elegant experiments show you that kids originally formed |
|
|
|
egocentric representations of the world around them and then gradually |
|
|
|
formed these worlds and centric presentations. |
|
|
|
So in one in one experiment shown on the top |
|
|
|
infants, less than one year, I placed in a room |
|
|
|
and the there's two doors to the room and they're |
|
|
|
sitting around the table with the experimenter, and their mother |
|
|
|
appears at one door and then the child is rotated |
|
|
|
in the room. |
|
|
|
The room doesn't change. |
|
|
|
The charges were rotated around the table. |
|
|
|
And the question is, where does the child expect the |
|
|
|
mother to peer from from the same door that she |
|
|
|
was at before or from the door that is on |
|
|
|
the right hand side of that child, which is the |
|
|
|
egocentric reference point. |
|
|
|
And the answer is that at least in young infants, |
|
|
|
they will prefer to look and to expect that for |
|
|
|
the mother to come from the right hand or that |
|
|
|
is the wrong doors. |
|
|
|
But the one that is on their right hand side |
|
|
|
is to their egocentric preference example. |
|
|
|
Now this Dallas centric discapacidad show which World Baseball and |
|
|
|
its fans develop very slowly. |
|
|
|
And indeed, some people remain pretty poor even when they |
|
|
|
get into adulthood, including myself. |
|
|
|
So this task is illustrative of that. |
|
|
|
In this task, one would ask a child or a |
|
|
|
toddler to indicate on the right here what the view |
|
|
|
from this horse is of this pattern of events here. |
|
|
|
Is it A, B, C, or D, just have a |
|
|
|
loophole here. |
|
|
|
Who thinks it's A, The view of the horse is |
|
|
|
represented. |
|
|
|
Okay. |
|
|
|
What about B? |
|
|
|
A couple of people see. |
|
|
|
Few people. |
|
|
|
They have a few people. |
|
|
|
The answer is B, with the red, blue and yellow. |
|
|
|
Proceed from left to right in the view of the |
|
|
|
horse would be to see it from the left to |
|
|
|
the right. |
|
|
|
So that's an that's a reimagining or interpret the world |
|
|
|
from another point of view. |
|
|
|
And that's interpreting what from another point of view requires |
|
|
|
that kind of understanding, because you have to build that |
|
|
|
representation in a world based audience based on your own |
|
|
|
context. |
|
|
|
So these two abilities, egocentric and polycentric reference frames develop |
|
|
|
at different rates and the egocentric comes first. |
|
|
|
As I said, I'm not going to talk much about |
|
|
|
our centric map here because Hugo is actually going to |
|
|
|
take you through that quite a bit. |
|
|
|
And I feel like this time when we talk about |
|
|
|
spatial memory and other aspects of historical function, but the |
|
|
|
fundamental basis of these other centric maps, these cognitive maps, |
|
|
|
was actually discovered at UCL by John O'Keefe, whose building |
|
|
|
is in the anatomy building. |
|
|
|
I think somewhere in the top drawer. |
|
|
|
He's still there, still researching. |
|
|
|
He must be 80 now. |
|
|
|
He won the Nobel Prize a few years ago, but |
|
|
|
it hasn't stopped him. |
|
|
|
And he discovered, as Hugo described to you, that if |
|
|
|
you recall from the hippocampus of a mouse or a |
|
|
|
rat wandering around a small wooden box, that the cells |
|
|
|
in that hippocampus will often fire in a particular location |
|
|
|
in that particular part of the puzzle. |
|
|
|
And further experiments show that that the position in the |
|
|
|
box where they respond to the red dots and the |
|
|
|
thing, the black things, the trajectory down through that space, |
|
|
|
red dots, when the neurone fires, we can show you |
|
|
|
via various manipulations that that representation of space that is |
|
|
|
embodied in that Nero's place cells activity is actually half |
|
|
|
a century. |
|
|
|
I want to spend the next few slides discussing taking |
|
|
|
through some of the egocentric reference frame that emerged in |
|
|
|
Cortex. |
|
|
|
So I've discussed already. |
|
|
|
One might be an extended work from reference. |
|
|
|
That is, if you move your eyes from this red |
|
|
|
dot here. |
|
|
|
If you look at this on the red dot, Wall-E |
|
|
|
is on the right hand side of your visual field. |
|
|
|
If you then transfer your case to the right to |
|
|
|
the dot on the right one is now on the |
|
|
|
left hand side of the visual field. |
|
|
|
That's an eye centred reference frame your eye moves and |
|
|
|
therefore consequently the position of the objects in the world |
|
|
|
moves with respect to the centre of gaze and with |
|
|
|
respect to your retina, even though they haven't changed position |
|
|
|
in the world. |
|
|
|
Similarly, this hits into reference frames, things which with respect |
|
|
|
my head. |
|
|
|
One example of that is audition sounds that have arrived |
|
|
|
at the head and are encoded with respect to the |
|
|
|
direction of the head and the eyes don't move unlike |
|
|
|
the eyes. |
|
|
|
And so that reference frame is based. |
|
|
|
There's other frames of reference, for example, joint frames of |
|
|
|
reference again for reaching out and grasping this thing. |
|
|
|
I might like to include the world in the context |
|
|
|
of the joints that are required to pick up this |
|
|
|
and handle this environment to tell me that the 20. |
|
|
|
So a good deal of work in monkeys that are |
|
|
|
described in humans has shown that these special frames of |
|
|
|
reference I'm almost certainly built to start to be built |
|
|
|
in the parietal cortex and prior to the cortex, the |
|
|
|
visual cortex, back of the brain. |
|
|
|
Frontal lobes are here, temporal lobes are down here in |
|
|
|
the proper lobes here, and it's parietal cortex, or at |
|
|
|
least this bit of the parietal cortex is important in |
|
|
|
generating representing the spatial reference frames found in the particular |
|
|
|
part of the parietal cortex around the parietal sulcus. |
|
|
|
What this image shows here is a summary of many |
|
|
|
studies looking at neglect with left sided neglect and the |
|
|
|
density, the colour of the blobs on the side of |
|
|
|
the brain that represent the probability effectively that damage in |
|
|
|
that part of the brain would have been associated with |
|
|
|
neglect. |
|
|
|
You can see there's very high concentration around this issue |
|
|
|
prior to Super Square. |
|
|
|
If you have damage, then you get this form of |
|
|
|
neglect or some form of neglect. |
|
|
|
So if we we Inhumans, we find out it's quite |
|
|
|
difficult to look in the exact new machinery in this |
|
|
|
part of the brain. |
|
|
|
But it is possible in monkeys. |
|
|
|
And it turns out that the newer machinery there in |
|
|
|
monkeys looks pretty similar to what we might expect from |
|
|
|
brain imaging in humans or from these lesions. |
|
|
|
And so we can use that information, these recordings, from |
|
|
|
a way about having monkeys be trained to do a |
|
|
|
task. |
|
|
|
Monkeys can usually perform a much more complicated task. |
|
|
|
Another animal such as flies or rodents. |
|
|
|
And we can use those recordings to try and work |
|
|
|
out what's actually going on in these little brain areas, |
|
|
|
particularly those around this practice focus area that's received most |
|
|
|
attention so far is the lateral improperly lip might be |
|
|
|
coming back with in the next class and even more |
|
|
|
involved questions about how we decide to make movements. |
|
|
|
There is some evidence that this area, this little area |
|
|
|
called LIP, is at least partially involved in starting to |
|
|
|
generate or start to move as you place an eye |
|
|
|
based reference frame, which is the kind of topographic map |
|
|
|
you find in primary visual cortex or in early parts |
|
|
|
of visual cortex. |
|
|
|
If we want to change that reference frame into something |
|
|
|
that is not dependent on when we are looking, we |
|
|
|
need to start making some changes to the representations that |
|
|
|
we encode this graph. |
|
|
|
We actually do a little bit about how why we |
|
|
|
think that is involved in starting to transform this visual |
|
|
|
tropic retina topic representation to something that does not depend |
|
|
|
on the direction of gaze. |
|
|
|
I mean, explain what these graphs all show first, because |
|
|
|
you're going to see a field in the next few |
|
|
|
slides on the x axis. |
|
|
|
Here is the time, and you can see here it's |
|
|
|
about this little black bar is about 200 milliseconds or |
|
|
|
one fifth of a second. |
|
|
|
There's several different things on the on the y axis |
|
|
|
here on the top is the position of the animal. |
|
|
|
Now, he's actually hit fixed in this case, which head |
|
|
|
is restrained, but he's able to move his eyes around |
|
|
|
because that is the vertical movements of the eye and |
|
|
|
the horizontal movements of the eye. |
|
|
|
The next block shows you, when the stimulus appears, indicates |
|
|
|
the times when the stimulus is on these little rows. |
|
|
|
Here are what we call a Rasta plot. |
|
|
|
And each row is one trial. |
|
|
|
One time the animal performed this task. |
|
|
|
And each of those dots is the time of occurrence |
|
|
|
of a single action potential from a neurone in IP. |
|
|
|
Now, you may repeat that trial many times in this |
|
|
|
case, say 15 or 20 times, and you get pretty |
|
|
|
similar activity across each trial. |
|
|
|
And so when you average that activity, you get something |
|
|
|
like this black box below which we call a pair |
|
|
|
of stimulus time histograms. |
|
|
|
And the height of that box basically reflects the magnitude |
|
|
|
of the response of the neurone at that point in |
|
|
|
time. |
|
|
|
The number of spikes the neurone is discharging on each |
|
|
|
trial at that point in time. |
|
|
|
And so in each of these cases, you can see |
|
|
|
that just after the stimulus comes on, the neurones respond |
|
|
|
very short latency about 50 or 80 milliseconds, less than |
|
|
|
1/10 of a second after the appearance of stimulus. |
|
|
|
That's how long it takes the visual information to get |
|
|
|
from the retina to this part of the brain. |
|
|
|
Now there's three different trials shown here. |
|
|
|
These are all recordings from the same nerve cell likely |
|
|
|
in the monkey doing this task. |
|
|
|
And the monkey's task is simply to find look at |
|
|
|
the dot bit like when you're looking at Wall-E. |
|
|
|
But we're not trying to look. |
|
|
|
But look at the dot in the letters, the thing |
|
|
|
that's all the monkeys required to do. |
|
|
|
In the other two trials, the monkey is required to |
|
|
|
move the eyes from that dot to another dot that |
|
|
|
appears. |
|
|
|
So you can see here, that's what's indicated by the |
|
|
|
arrow. |
|
|
|
And you can see that the monkey is successfully doing |
|
|
|
that because it's horizontal traits which are the sort of |
|
|
|
the right position, horizontal and position changes just after the |
|
|
|
stock comes on. |
|
|
|
And that comes on at this line here. |
|
|
|
And then a stimulus has been displayed to the animal |
|
|
|
at a particular location on the TV screen that the |
|
|
|
dog is also placed on. |
|
|
|
So in this particular case on the left here, this |
|
|
|
is what we would expect from a neurone that is |
|
|
|
simply representing visual stimuli. |
|
|
|
Responding to a visual stimulus, Stan was fixating flecks of |
|
|
|
light appears neurone stops responding. |
|
|
|
Pretty simple. |
|
|
|
In the second case here, this is a bit more |
|
|
|
of a funky task in this case. |
|
|
|
Just before the animal makes nine movement, the stimulus appears |
|
|
|
on the right hand side. |
|
|
|
And here you can see what happens is that the |
|
|
|
neurone still responds to the stimulus that appears when when |
|
|
|
the eye move to that location. |
|
|
|
So that that is also consistent perhaps with the with |
|
|
|
the new on including the position or appearance of an |
|
|
|
object at a particular position with respect to its iris. |
|
|
|
However, if you display the stimulus just briefly at its |
|
|
|
new location here, the same written location with respect to |
|
|
|
the days before the animal makes the second you actually |
|
|
|
see that response physically or even though that stimulus was |
|
|
|
placed in the part of the visual field, it's not |
|
|
|
normally effective for the mirror. |
|
|
|
In other words, this new on CTV anticipate. |
|
|
|
In fact, the animal is about to make a second |
|
|
|
to this new location in middle place, and it seems |
|
|
|
to be reaching out to that new location to see |
|
|
|
if there's anything there already. |
|
|
|
So these neurones are already starting to disassociate the retina |
|
|
|
properly. |
|
|
|
Framework that is encoding really powerful visual cortex is something |
|
|
|
that doesn't depend on the location of the eye. |
|
|
|
It's not a complete disassociation to pinpoint. |
|
|
|
So that's like P going to see patent in this |
|
|
|
VR trial area eventually further down in the brain, away |
|
|
|
from the brain. |
|
|
|
And these neurones are really interesting. |
|
|
|
Again, this is a monkey performing a task. |
|
|
|
And again, the task is primarily to maintain fixation on |
|
|
|
a particular point on the screen. |
|
|
|
In this case we see the monkey and we can |
|
|
|
see from the lateral side he's looking at the screen. |
|
|
|
And his task here is simply to look at this |
|
|
|
central location. |
|
|
|
And while he's looking down, a stimulus is presented that |
|
|
|
comes either towards the mouth. |
|
|
|
From the top to the bottom, from different locations on |
|
|
|
the screen or towards the top of the head, again, |
|
|
|
from different locations on the screen. |
|
|
|
And what you should see here is that this neurone |
|
|
|
is very active for the two situations. |
|
|
|
On the left hand side here, there's lots of activity, |
|
|
|
lots of spikes, and that's when the object is moving |
|
|
|
towards the mouth and not when the object is moving |
|
|
|
towards the forest. |
|
|
|
If we then change the stimulus slightly. |
|
|
|
So then what? |
|
|
|
She has to look up here. |
|
|
|
You can see that the neurone is too responsive when |
|
|
|
the animal is when the object is coming towards the |
|
|
|
mouth, not towards the first. |
|
|
|
So the activity this neurone seems to depend on whether |
|
|
|
an object is moving towards the mouth, not on where |
|
|
|
the animal's viewpoint is. |
|
|
|
It doesn't matter where abouts in the visual field. |
|
|
|
It started as long as it was coming towards the |
|
|
|
end of the forest. |
|
|
|
So this vector into prior CO areas seems to be |
|
|
|
standing to construct this representation of objects in the world |
|
|
|
that depend on that location. |
|
|
|
With respect to the amount of fibre that's important for |
|
|
|
feeding. |
|
|
|
Exactly. |
|
|
|
And not just the math. |
|
|
|
If you look at other neurones in other parts of |
|
|
|
the head that represented. |
|
|
|
And it's also interesting you report the medial enterprise area |
|
|
|
rather than the lateral, the medial being close to the |
|
|
|
middle of the brain. |
|
|
|
We find exactly what we call rich second frames of |
|
|
|
reference. |
|
|
|
That is, that neurones are responsive when animals make a |
|
|
|
movement towards an object with their arms is also neurones |
|
|
|
in there that are also responsible to animals. |
|
|
|
Make seconds. |
|
|
|
This is a fairly complicated slide. |
|
|
|
I don't want to take too much away from it, |
|
|
|
but the point is here that this new one, some |
|
|
|
of these neurones are active when the animal makes a |
|
|
|
reaching movement, but not when it makes an eye movement. |
|
|
|
It seems to be this and other forms of evidence |
|
|
|
seem to suggest that it was using something about the |
|
|
|
coordinate space of the joints to actually represent the outside |
|
|
|
world. |
|
|
|
There's another area called the anterior inter parietal area, and |
|
|
|
we're going to discuss that in greater depth in a |
|
|
|
moment. |
|
|
|
So to summarise what I've said to you there, there |
|
|
|
are multiple frames of reference that can be used to |
|
|
|
represent the world. |
|
|
|
There's good evidence for the existence of multiple frames of |
|
|
|
reference in separate circuits in the brain. |
|
|
|
Spatial neglect means losing a representation of a specific frame |
|
|
|
of reference for at least one or two, and not |
|
|
|
all of them. |
|
|
|
But for that reason we can all. |
|
|
|
Be aware of objects in a particular coordinate frame. |
|
|
|
And the other point here is that there's multiple frames |
|
|
|
of reference represented in parallel different areas in the brain. |
|
|
|
Simply constructing in parallel with different reference frames. |
|
|
|
And the consequence of that is that the could have |
|
|
|
been executed or generated in parallel. |
|
|
|
When we go to form a task, we can select |
|
|
|
immediately or quickly which frame of reference we want to |
|
|
|
use to actually complete that task. |
|
|
|
We don't have to wait to redo the entire computation |
|
|
|
again, take the visual image on whatever I want to |
|
|
|
do with this. |
|
|
|
I want to reach there. |
|
|
|
I want to do this instead. |
|
|
|
That preference frame is already being built. |
|
|
|
We may not use that reference frame. |
|
|
|
We may select another reference frame, in which case the |
|
|
|
question becomes what happens to the new workflow detection of |
|
|
|
that reference frame that we did not use? |
|
|
|
And that, I think, is going to be something that |
|
|
|
should become clear in the next part of the lecture. |
|
|
|
Is there any questions about that particular component? |
|
|
|
Okay. |
|
|
|
So skip over a section, which is basically how the |
|
|
|
motor cortex controls the muscles. |
|
|
|
And I could say that there's a there's now a |
|
|
|
video on your little page, which is through that, and |
|
|
|
it's five people. |
|
|
|
What I want to spend. |
|
|
|
The next lecture discussion is how we control and even |
|
|
|
understand actions. |
|
|
|
And I want to take us through some of the |
|
|
|
really interesting what has happened in this field in the |
|
|
|
last ten or 15 years. |
|
|
|
We've talked about the parietal lobe. |
|
|
|
We skip over what I call the primary motor cortex, |
|
|
|
which is the actual guts of controlling the muscles. |
|
|
|
That's in the middle page. |
|
|
|
We're not going to talk too much about supplementary motor |
|
|
|
areas, but these are areas which help generate the initial |
|
|
|
plans for the muscle movements that go to the primary |
|
|
|
motor cortex. |
|
|
|
But between these areas, the premotor cortex and the prefrontal |
|
|
|
cortex seem to take information from the parietal lobe and |
|
|
|
then distribute them to the motor areas. |
|
|
|
That's a lot of what we understand about this has |
|
|
|
been done in the context of a particular term, and |
|
|
|
I think we can grasp something. |
|
|
|
Rastafarian or a monkey's case brought a little bit of |
|
|
|
food. |
|
|
|
Turns out that the circuits for this simply quite similar |
|
|
|
in monkeys and humans. |
|
|
|
The talk shows a schematic of a monkey brain, the |
|
|
|
bottom of the human brain, the areas of interest in |
|
|
|
the monkey brain of the those around the enterprise, those |
|
|
|
focus in particular the anterior area is a little area |
|
|
|
called F5 and also F1 in humans. |
|
|
|
The same areas exist. |
|
|
|
We know that from the anatomy and from the tracing |
|
|
|
of connections between pathways. |
|
|
|
We don't really know exactly how signals get from one |
|
|
|
area to another. |
|
|
|
So we're going to use the monkey to try and |
|
|
|
understand a bit of that, but also look at some |
|
|
|
of the human pathways. |
|
|
|
So this here is in the same kind of way. |
|
|
|
Actually, the previous slides, a description of several neurones in |
|
|
|
anterior into parietal area during grasping, which I find is |
|
|
|
absolutely fascinating because there's one particular type of neurone here |
|
|
|
in the bottom right, which seems to be important in |
|
|
|
taking that sensory information that's coming up from the sensory |
|
|
|
periphery through the visual cortex and starting to transform that |
|
|
|
into something that's useful for motor movements. |
|
|
|
So this slide shows three separate neurones in each row |
|
|
|
and for each new on the three different paths. |
|
|
|
The different tasks. |
|
|
|
The different columns are to perform a manipulation in light |
|
|
|
that is to create something more certain. |
|
|
|
See it? |
|
|
|
The second one is to reach out and grasp that |
|
|
|
thing in the dark. |
|
|
|
So I can't see there is no visual information. |
|
|
|
And the third thing is just to look at the |
|
|
|
object and not reach the ground. |
|
|
|
So the neurone on the top here shows falling rates |
|
|
|
like we might expect from a visual appeal if it's |
|
|
|
near or it's active. |
|
|
|
When the animals manipulate an object in the light, it's |
|
|
|
also active when it sees the object to manipulate it, |
|
|
|
but it's not at all active when it makes the |
|
|
|
muscle movements, they can't see the object. |
|
|
|
On the other hand, this middle row here is a |
|
|
|
kind of demand that we might expect to be important |
|
|
|
in controlling the movements that we're about to make so |
|
|
|
that no one is active. |
|
|
|
When we reach out across the line and is active |
|
|
|
when we reach out and draw something, the dog can |
|
|
|
see it, but it's not at all active. |
|
|
|
When we just look at the optics. |
|
|
|
So it's active during the past, but when you look |
|
|
|
at it, so these visual motor neurones, it's a kind |
|
|
|
of classic distinction between sensory input and motor. |
|
|
|
And for the third part of neurone there, which we'll |
|
|
|
call a visual motor neurone, combines these two features. |
|
|
|
These neurones again are active during the regrouping task in |
|
|
|
the light. |
|
|
|
These neurones are also active when the animals cannot see |
|
|
|
the objects and they're also active when they see it, |
|
|
|
but don't perform the movement. |
|
|
|
So they have both sensory input seen in the visual |
|
|
|
only component and motor inputs in the motor and components. |
|
|
|
And they seem to combine, they seem to be able |
|
|
|
to combine these two forms into one neurone. |
|
|
|
And these kinds of neurones, these visual motor neurones can |
|
|
|
be found in different parts of the brain. |
|
|
|
The most prominent in the kind of cortex, the frontal |
|
|
|
cortex. |
|
|
|
And they are, we think, a very important interface between |
|
|
|
sensory information. |
|
|
|
They combine these two things. |
|
|
|
As we'll discover in the next lecture, You want to |
|
|
|
combine these two things to make the interesting contributions to |
|
|
|
defining about what we want to do. |
|
|
|
So this is the kind of neurones you find in |
|
|
|
the final area. |
|
|
|
You actually find is also further up the chain area. |
|
|
|
Fine. |
|
|
|
We'll get to that in a moment. |
|
|
|
These neurones that can be quite selective for the further |
|
|
|
features of the task. |
|
|
|
So this is recording from one neurone in this case, |
|
|
|
and this is a visual motor neurone that is a |
|
|
|
neurone that you can see both the visual component and |
|
|
|
motor in front of the task. |
|
|
|
In this case you can see that just in the |
|
|
|
visual component alone that the neurones are responsive. |
|
|
|
This is. |
|
|
|
This is during the top three. |
|
|
|
This is the actual during the top when they can |
|
|
|
see and make the movement. |
|
|
|
You can see that these neurones are active for particular |
|
|
|
configurations of grasp and not others. |
|
|
|
If in addition you look at the visual component alone, |
|
|
|
that is in the absence of the tons you see, |
|
|
|
these neurones are also selected for particular objects. |
|
|
|
How many of you have heard of the concept of |
|
|
|
affordable? |
|
|
|
Performance is a really interesting thing. |
|
|
|
When we design objects, the objects that work, the ones |
|
|
|
that we want to use for those are for particular |
|
|
|
actions. |
|
|
|
The chairs are for the ideas to be. |
|
|
|
I don't do that with. |
|
|
|
That means phone too, for the idea of picking them |
|
|
|
up and scrolling through them. |
|
|
|
That is the very particular structure. |
|
|
|
Those objects seems to promote the execution of particular motor |
|
|
|
planner. |
|
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Unfortunately. |
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They forward those. |
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Actions. |
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Maybe these neurones are part of that kind of affordances, |
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because when these neurones which are helping, which have both |
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sensory input and motor output, are actually representing particular types |
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of objects, people, okay, So perhaps they help us not |
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to generate the plants when we see that object. |
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One of the plants that I might execute if God |
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wants, but I might not execute one of the plants, |
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I might get a clue that that would afford that |
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particular action. |
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Indeed, if you look at human cortex now rather than |
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monkey cortex, you look at it from signals rather than |
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single unit recordings. |
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You also see a good deal of evidence for the |
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presence of areas that are effectively related to performance. |
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These are the responses or MRI responses in human cortex |
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to observed axons in the top row. |
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Here is when one observes actions without associated objects, and |
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the bottom is when objects are present with chewing, grasping |
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and kicking. |
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Again, this is a person sitting in a scanner and |
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not actually performing his actions that is viewing the action. |
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We think maybe when they're viewing these actions, maybe they're |
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kind of replaying them in their head as well. |
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Or maybe they're viewing them also for to start to |
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build their idea of a particular course of action. |
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And you see from these slides here, but I want |
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you to take away mainly is that there's a substantial |
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amount of activity not only in motor cortex, but also |
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in provider cortex consistent with what we see in the |
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monkeys that when they're looking at particular objects, neurones are |
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active so that those objects may afford particular actions. |
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Similarly, if you just look at the particular example of |
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tools, if they present a hammer to someone sitting in |
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a scanner as opposed to a house or other objects, |
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they would do not forward actions. |
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You find that in prior to Cortex as well as |
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in premotor cortex, this area equivalent to the monkey even |
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at five feet, gets a standard manner of activity. |
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Again, these parietal and motor cortical region seem to be |
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responsive when you're viewing objects that are full potential for |
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action. |
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This experiment I find really beautiful. |
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You've heard about transcranial magnetic stimulation. |
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That's the idea where you put a magnetic pulse outside |
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to stimulate a little bit to the cortex. |
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When you do that, you activate the neurones, the electrical, |
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the cortex. |
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|
You can choose the pulse and intensity that which doesn't |
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overtly cause an action, but you can measure the activity |
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|
of muscles in the relevant part of the body. |
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For example, in the hand. |
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And that's what's going on here. |
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These will pass through the amplitude of the muscle movement |
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recorded during stimulation of the cortex. |
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And these measurements are made. |
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What people are feeling for different objects. |
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These are right handed subjects. |
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In one case of viewing a cup with the left |
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hand of one face, right hand to the other cases. |
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Left and right. |
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Broken handles. |
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|
And what you should see here is that the bar |
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is much higher when the cops are right handed. |
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When their visual object affords particular action. |
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Grasping with the right hand activity, motor cortex seems to |
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be potentiated by viewing this object consistent again with the |
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from right results because cool and consistent again with the |
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work and try to poaching monkeys. |
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So this brings me on then the mirror neurones. |
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How many people have heard of Mira Nair? |
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One of the few who you might know. |
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And if you've read the if you read the Sylvia |
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Hazily speaking, if you read the review that I've included |
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in your reading, you'll understand a lot more about them. |
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|
Their neurones have gained particular notoriety because they may, they |
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are hypothesised to be a source of understanding the actions |
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|
of others and perhaps even the higher cognitive social events |
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|
like empathy. |
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|
These are circuits to be active when viewing an object |
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|
for viewing someone doing something as opposed to actually doing |
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|
it yourself as well as when you're doing so. |
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|
So these are neurones originally that were discovered. |
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|
The reason I've talked to you about this search for |
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|
grasping is that these neurones were discovered by Richard Lockie |
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|
and colleagues when they were trying to understand these brain |
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|
circuits across that recording from monkeys. |
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|
And they noticed that and trained the monkeys to reach |
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|
out and collect food objects. |
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|
I wanted to know what was happening during the different |
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|
phases of movement and plenty of movement. |
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|
They're reporting from the pool area of the prefrontal cortex |
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|
called a client. |
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|
And they notice that a certain fraction of neurones consistently |
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|
seem to fire, not when the animal, not just when |
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|
the animal restarting something, but when the experimenter was actually |
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|
moving or reaching out or grabbing that. |
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|
And so. |
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|
So in the writing of the classroom, one of the |
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|
first neurones that they discovered the classical mirror neurones. |
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|
So these neurones are active when an animal is reaching |
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|
out and grasping an object that's shown here. |
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|
They're also active when the experimenter is reaching out and |
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|
grasping the object that's shown here. |
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|
Some of these new ones, it turns out, with finer |
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|
investigations, were active in particular cases when the animal when |
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|
the object was available to the animal's reach and when |
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|
it was not available within the animals, when it's within |
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|
the animal's reach, that's a very personal space that the |
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|
space around you when it's not within the reach of |
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|
the expert personal space. |
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|
The space is beyond that. |
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|
You can see here that some neurones seem to respond |
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|
when actions are performed outside of personal space and other |
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|
neurones respond when it's in the very personal space. |
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|
Yeah. |
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|
So the idea here is that these neurones are responsive |
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|
to when both an animal is making an action and |
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|
when the animal is viewing the action. |
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|
So that's where the idea of mirror neurones comes up |
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|
in mirroring responsive when you're viewing something as if you |
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|
were generating that they embody in your brain an interpretation |
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|
of the action potentially that took place or the accident |
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|
that person is performing. |
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|
That might be important. |
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|
For example, imitation, learning, fine imitation like children do. |
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|
If you perform an action should take it and learn |
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|
how to perform that action. |
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|
However, monkeys don't really live by invitation, so weather is |
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|
|
certainly a problem with monkeys. |
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|
Another question Immigration could also be important for trying to |
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|
predict what someone else is going to do. |
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|
If I see someone reaching out for an object and |
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|
predicting that they're going to be taking an object, I'll |
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|
be able to understand the potential intentions of the other |
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|
person. |
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|
That's the idea of the founding at five. |
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|
Why? |
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|
I wanted to introduce you to them in the context |
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|
of this pathways that you should be clear to by |
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|
now that they're not just on an incline. |
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|
They're also found an intuitive triangle area. |
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|
They also find another part of the brain. |
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|
So there seems to be a whole network of neurones |
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|
that seem to be providing not just the actions that |
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|
we are making, but when we just view those actions |
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|
|
will result in a positive step forward. |
|
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|
These circuits are collectively known as potentially very neurone circuits |
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|
and they are thought to be in some context important |
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|
|
for things like embodiment and empathy. |
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|
I do want to make clear, though, that there's there's |
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|
|
a lot of controversy. |
|
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|
ACLU review makes clear as well. |
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|
There's a lot of controversy about the role of these |
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|
neurones in these functions. |
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|
I don't think there's any controversy about the presence of |
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|
these neurones. |
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|
What is of interest and what is of concern is |
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|
whether these neurones are actually representing what another person is |
|
|
|
doing with these neurones, basically generating internal but complete the |
|
|
|
plan of action. |
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|
As I said to you before, we provide different spatial |
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|
|
reference findings in parallel in our brains, we choose which |
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|
spatial reference frame we're going to use when we execute |
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|
a particular action. |
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|
The implication of having multiple parallel reference frames, not all |
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|
|
of which are used, is that many. |
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|
Action plans are made and never executed and never whenever |
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|
|
aware of. |
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|
It's a mirror neurone that for a neurone that is |
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|
actually representing the actions of others. |
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|
Or is it a neurone that's representing an on executed |
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|
|
plans that we are protecting. |
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|
Which. |
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|
Afforded. |
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|
To us. |
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|
And that's the central question that still exists in the |
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|
mirror neurone. |
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|
Which are these neurones effectively anonymous, or are they there |
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|
to help us interpret the actions of others or not? |
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|
And these of course are not there to speak of |
|
|
|
things they could well be that they, for example, evolved |
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|
|
or arose simply to provide clues and execute plans that |
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|
|
are co-opted to then try and provide an interpretation understanding |
|
|
|
of others actions. |
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|
So I really suggest that you do read Sylvia's article. |
|
|
|
She's a leader in the field. |
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|
And at this stage, I will leave you with those |
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|
|
with one dichotomy. |
|
|
|
That's the dichotomy that exists in the field with the |
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|
|
laity who discovered these very new ones originally. |
|
|
|
His group of sceptics were neurones. |
|
|
|
He's his interpretation, as if this mirror mechanism is fundamental |
|
|
|
to understanding actions and intentions, then the classical view to |
|
|
|
the motor system has only a role in generation and |
|
|
|
by implication the central system has any role in sensation, |
|
|
|
have to be rejected and replaced by the view that |
|
|
|
motor system is also one of the major players in |
|
|
|
cognitive functions. |
|
|
|
The motor system helps us embody the actions that we |
|
|
|
view around us. |
|
|
|
The contract. |
|
|
|
The contract view is that from Hitchcock and colleagues, he |
|
|
|
spoke with the language specialists very prominent in the field, |
|
|
|
and he would state that a null hypothesis is in |
|
|
|
this area. |
|
|
|
A five is fundamentally a motor area that is capable |
|
|
|
of supporting sensory motor associations that are relevant to action |
|
|
|
selection. |
|
|
|
As I said before, one of these areas that is |
|
|
|
involved in generative frames of reference potential, the motor plans |
|
|
|
that we can execute this selection. |
|
|
|
So encourage you to read those things and we'll leave |
|
|
|
that idea of whether or not we're going to contribute |
|
|
|
fundamentally provision for thanks and how good they can click. |
|
|
|
Click, click, click here to read more effective. |
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|
|
Problems. |
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|
With you. |
|
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|
Take me back to the current context in which you |
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|
|
can contribute to that. |
|
|
|
I could take some time to. |
|
|
|
Come up with something specifically at the control. |
|
|
|
Group. |
|
|
|
That's one of the things that we suspect virtually all. |
|
|
|
Of the. |
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|
|
Idea. |
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|
|
But I don't think to. |
|
|
|
Make the point that medical technicians and. |
|
|
|
An important part of. |
|
|
|
The process. |
|
|
|
Of critical thinking about this is something that even. |
|
|
|
When I'm writing. |
|
|
|
The basic hypothesis. |
|
|
|
Is that there's a lot of activity. |
|
|
|
Quickly. |
|
|
|
Actually get. |
|
|
|
Effective. |
|
|
|
So it just doesn't make any difference whether. |
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|
|
Or not the significant. |
|
|
|
Activity in this market. |
|
|
|
Commercial activity. |
|
|
|
But you might find indications of. |
|
|
|
What it looks like. |
|
|
|
It's going to. |
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|
|
Be very difficult to keep. |
|
|
|
Up with. |
|
|
|
The idea. |
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|
That an activity. |
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|
By. |
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|
Virtue of that are active but actually taking place. |
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|
I see. |
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|
Okay. |
|
|
|
Thank you for taking my question, please. |
|
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|
Okay. |
|
|
|
I guess it's like if you read the study as |
|
|
|
well, but just. |
|
|
|
The active when it's objects moving towards the mouse. |
|
|
|
That's what I'm trying to say, but only when they're |
|
|
|
moving ahead of the target. |
|
|
|
So it's much more different from anything. |
|
|
|
Yeah, it's so interesting. |
|
|
|
Question Is there any. |
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|
|
More than what is it. |
|
|
|
Going to take that we know that we know psychologically, |
|
|
|
that whole thing for. |
|
|
|
Actions. |
|
|
|
Whether that specific you know, whether specific structure can afford |
|
|
|
to do for specific. |
|
|
|
Comparisons as any work is done in the executive there? |
|
|
|
But it's a good question. |
|
|
|
I don't think it's coming back. |
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|
But I don't know. |
|
|
|
But with all the time it takes to do something |
|
|
|
like this happens, but it seems to be generated, they're |
|
|
|
more expensive. |
|
|
|
But again, we recognise that. |
|
|
|
People. |
|
|
|
Keep working on. |
|
|
|
It could be difficult to predict what's going to happen |
|
|
|
in Texas because that's not something that's going to come |
|
|
|
to grips with the fact that people are killed by. |
|
|
|
In particular for the focus groups which are protected in |
|
|
|
the construction of the company as well as collaborations. |