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.