Emergence of belief-like representations through reinforcement learning — part 1/7
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Emergence of belief-like representations through reinforcement learning Jay A. Hennig1,2,∗, Sandra A. Romero Pinto2,3, Takahiro Yamaguchi3,4, Scott W. Linderman5,6, Naoshige Uchida2,3, Samuel J. Gershman1,2 1 Department of Psychology, Harvard University, Cambridge, MA, USA 2 Center for Brain Science, Harvard University, Cambridge, MA, USA 3 Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA, USA 4 Future Vehicle Research Department, Toyota Research Institute North America, Toyota Motor North America Inc., Ann Arbor, MI, USA 5 Wu Tsai Neurosciences Institute, Stanford University, Stanford, CA, USA 6 Department of Statistics, Stanford University, Stanford, CA, USA * Corresponding author Email: jay.a.hennig@gmail.com (JAH) 1 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint Abstract To behave adaptively, animals must learn to predict future reward, or value. To do this, animals are thought1 to learn reward predictions using reinforcement learning. However, in contrast to classical models, animals2 must learn to estimate value using only incomplete state information. Previous work suggests that animals3 estimate value in partially observable tasks by first forming “beliefs”—optimal Bayesian estimates of the4 hidden states in the task. Although this is one way to solve the problem of partial observability, it is not the5 only way, nor is it the most computationally scalable solution in complex, real-world environments. Here6 we show that a recurrent neural network (RNN) can learn to estimate value directly from observations, gen-7 erating reward prediction errors that resemble those observed experimentally, without any explicit objective8 of estimating beliefs. We integrate statistical, functional, and dynamical systems perspectives on beliefs to9 show that the RNN’s learned representation encodes belief information, but only when the RNN’s capac-10 ity is sufficiently large. These results illustrate how animals can estimate value in tasks without explicitly11 estimating beliefs, yielding a representation useful for systems with limited capacity.12 Author Summary13 Natural environments are full of uncertainty. For example, just because my fridge had food in it yester-14 day does not mean it will have food today. Despite such uncertainty, animals can estimate which states15 and actions are the most valuable. Previous work suggests that animals estimate value using a brain area16 called the basal ganglia, using a process resembling a reinforcement learning algorithm called TD learning.17 However, traditional reinforcement learning algorithms cannot accurately estimate value in environments18 with state uncertainty (e.g., when my fridge’s contents are unknown). One way around this problem is if19 agents form “beliefs,” a probabilistic estimate of how likely each state is, given any observations so far.20 However, estimating beliefs is a demanding process that may not be possible for animals in more complex21 environments. Here we show that an artificial recurrent neural network (RNN) trained with TD learning22 can estimate value from observations, without explicitly estimating beliefs. The trained RNN’s error signals23 resembled the neural activity of dopamine neurons measured during the same task. Importantly, the RNN’s24 2 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint activity resembled beliefs, but only when the RNN had enough capacity. This work illustrates how animals25 could estimate value in uncertain environments without needing to first form beliefs, which may be useful26 in environments where computing the true beliefs is too costly.27 Introduction28 One pervasive feature of animal behavior is the ability to predict future reward. For example, a dog may29 learn that when her owner picks up the leash, she is likely to be rewarded with a walk in the near future.30 In associative learning settings such as this one, animals learn to associate certain stimuli (e.g., her owner31 grabbing the leash) with future reward (e.g., a walk). The neural basis of associative learning has been32 interpreted through the lens of reinforcement learning (RL). In particular, one successful theoretical model33 posits that associative learning is driven by the activity of dopamine neurons in the midbrain, where spiking34 activity resembles the reward prediction error (RPE) signal in an RL algorithm called temporal difference35 (TD) learning [1, 2, 3, 4]. We will describe this algorithm in more detail below.36 In many real-world scenarios, effectively predicting reward may require a deeper understanding of the37 structure of the world that goes beyond associating observations and reward. To continue the example38 above, suppose the dog’s owner keeps his car keys under the leash. Now if he picks up the leash, this39 does not necessarily mean he is about to take his dog on a walk. In other words, his intention to take40 his dog on a walk is now “partially observable.” Standard RL approaches are insufficient for learning in41 partially observable environments, as these methods assume that all relevant states of the environment are42 fully observable. One way to solve this problem is by using observations to form a Bayesian posterior43 estimate of each hidden state, called a belief state [5]. Future reward can then be estimated by applying44 standard RL methods like TD learning to belief states rather than the raw observations.45 Do animals estimate future reward using belief states? Evidence for this idea is suggestive, although46 indirect. Previous experimental work has shown that the phasic activity of midbrain dopamine neurons47 resembles the RPEs of TD learning in partially observable environments, where TD learning is performed48 on belief states rather than observations [6, 7, 8, 9, 10, 11]. The brain may have dedicated machinery, perhaps49 in prefrontal cortex [12, 13], for computing belief states, which could then be provided to downstream areas,50 3 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint such as the basal ganglia, to perform standard RL algorithms such as TD learning [14]. This architectural51 division of labor resonates with the broader literature on probabilistic computation in cortex, which has52 identified several different ways in which belief states could be encoded by neural activity [15].53 There are a few difficulties in using a belief state to solve RL tasks. First, the belief state assumes54 knowledge of the environment’s transition and observation dynamics—something that may be challenging55 for animals to acquire via observations alone. Indeed, there are well-documented examples of animals56 failing to learn or use the correct environment model [16]. Second, the belief representation does not scale57 well to more realistic tasks with higher-dimensional state spaces, as beliefs live in a continuous space whose58 dimensionality grows with the number of discrete states in the environment. Finally, the belief state includes59 knowledge about all states in the environment, regardless of whether those states are relevant to the task at60 hand. Luckily, one can often use approximate representations of beliefs to find solutions that work well in61 practice [17]. This suggests that there may be other representations, more compact than the full belief state,62 that are sufficient for the particular task of estimating future reward [18, 19].63 To address these difficulties, here we take inspiration from deep reinforcement learning. In deep RL,64 rather than explicitly learning beliefs, an agent uses nonlinear function approximation to learn a hidden65 representation that is sufficient for performing the task [20]. Compared to the belief representation, this66 approach does not require explicit knowledge about the structure of the environment. It may also scale67 better to more complex tasks, by virtue of the agent not needing to represent any features of the environment68 that do not directly pertain to the task at hand. Because beliefs are a non-linear dynamical system, here69 we use recurrent neural networks (RNNs) as our nonlinear function approximator. This choice was also70 motivated by the observation from the machine learning literature that RNNs can perform well on complex71 partially observable tasks [21]. Previous work in computational neuroscience has explored whether RNNs72 can be used to directly compute beliefs [17]. Here, by contrast, we explore whether training RNNs in73 partially observable environments leads to their representations implicitly becoming more like beliefs.74 We show that RNNs can be trained to perform two previously studied associative learning tasks [7, 10],75 reproducing experimentally observed dopamine neuron activity. We probe the representations learned by the76 RNNs, and show that the representations resemble beliefs from statistical, functional, and dynamical systems77 perspectives. Finally, we show how an RNN’s capacity determines the degree to which its representation78 4 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint resembles beliefs, without a concurrent impact on its ability to perform the task. These results illustrate79 how animals might estimate reward in partially observable environments without requiring any explicit80 representation of beliefs.81 Results82 Reinforcement learning in partially observable environments using belief states83 The standard RL objective is to learn the expected discounted future return, or value, of each state: V (st) = E "∞X k=0 γkrt+k # (1) wherest∈S is the state of the environment at time t, 0≤γ <1 is a discount factor, andrt is the reward.84 Rewards are random variables that depend on the environment state, and E denotes an expectation over the85 potentially stochastic sequences of states and rewards. Because we are interested in modeling Pavlovian86 associative learning tasks, here we will assume there are no actions available to the agent. For notational87 simplicity, we will use the shorthandVt =V (st).88 TD learning estimates the value function by exploiting a set of Markovian assumptions about the en- vironment: the probability of state st depends only on the last state st−1, and the probability of reward rt depends only on the current statest. Under these assumptions, the value function can be decomposed into a recursive form known as the Bellman equation [22]: Vt = E[rt +γVt+1]. (2) If an agent has access to an approximation of the value function, bVt, along with sample paths of states and rewards, then it can compute an estimate of the discrepancy between the approximate and true value function by taking the difference between the two sides of the Bellman equation: δt =rt +γ bVt+1− bVt. (3) 5 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint This is the temporal difference error, the precise definition of the RPE used in TD learning. Under the Bellman equation, E[δt] = 0 when bVt = Vt. If bVt is determined by a set of adaptable parameters θ, the approximation can be improved by following the stochastic gradient of the squared TD error: ∆θ =ηδt∇θ bVt, (4) where 0<η < 1 is a learning rate, and∇θ bVt is the gradient of bVt with respect toθ. One common example is a linear function approximator: bVt = DX d=1 w(d)zt(d), (5) wherezt(d)∈ R is some feature (indexed by d) of the state, and θ =w∈ RD is a learned set of weights89 on all the features. Under this approximation,∇θ bVt =zt.90 In a partially observable Markov process, agents do not observe the statest directly, but instead observe91 only observations ot ∈O . Critically, observations are not in general Markovian, which means that TD92 learning methods cannot be naively applied to value function approximators defined over observations. One93 way of understanding this is to note that the value of an observation may depend on the long-term past [23].94 In the dog leash example from the Introduction, the value (to the dog) of her owner picking up the leash95 depends on the history of events leading up to that moment—for example, if her owner recently announced96 that his car keys were missing. Reward prediction requires a compression of this history into a “sufficient97 statistic”—in effect creating a transformed state space over which the Markov property holds. TD learning98 can then be applied to this transformed state space.99 One such transformed state is the posterior probability distribution over hidden states given the history of observations, also known as the belief state [5]: bt(st) =P (st|o1,...,o t) ∝P (ot|st) Z P (st|st−1)bt−1(st−1)∂st−1 (6) where the recursion follows from the Chapman-Kolmogorov equation, which stipulates how to update the100 6 .CC-BY 4.0 International licenseavailable under a was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made The copyright holder for this preprint (whichthis version posted April 7, 2023. ; https://doi.org/10.1101/2023.04.04.535512doi: bioRxiv preprint