Predictive coding networks are neuroscience-inspired models with roots in both Bayesian statistics and neuroscience. Training such models, however, is quite inefficient and unstable. In this work, we show how by simply changing the temporal scheduling of the update rule for the synaptic weights leads to an algorithm that is much more efficient and stable than the original one, and has theoretical guarantees in terms of convergence. The proposed algorithm, that we call incremental predictive coding (iPC) is also more biologically plausible than the original one, as it it fully automatic. In an extensive set of experiments, we show that iPC constantly performs better than the original formulation on a large number of benchmarks for image classification, as well as for the training of both conditional and masked language models, in terms of test accuracy, efficiency, and convergence with respect to a large set of hyperparameters.

Biologically plausible learning algorithms offer a promising alternative to traditional deep learning techniques, especially in overcoming the limitations of backpropagation in fast and low-energy neuromorphic implementations. To this end, there has been extensive research in understanding what their capabilities are. In this work, we show how one of such algorithms, called predictive coding, is able to perform causal inference tasks. First, we show how a simple change in the inference process of predictive coding enables to compute interventions without the need to mutilate or redefine a causal graph. Then, we explore applications in cases where the graph is unknown, and has to be inferred from observational data. Empirically, we show how such findings can be used to improve the performance of predictive coding in image classification tasks, and conclude that such models are naturally able to perform causal inference tasks using a biologically plausible kind of message passing.

Animals can adapt their preferences for different types of reward according to physiological state, such as hunger or thirst. To explain this ability, we employ a simple multi-objective reinforcement learning model that learns multiple values according to different reward dimensions such as food or water. We show that by weighting these learned values according to the current needs, behaviour may be flexibly adapted to present preferences. This model predicts that individual dopamine neurons should encode the errors associated with some reward dimensions more than with others. To provide a preliminary test of this prediction, we reanalysed a small dataset obtained from a single primate in an experiment which to our knowledge is the only published study where the responses of dopamine neurons to stimuli predicting distinct types of rewards were recorded. We observed that in addition to subjective economic value, dopamine neurons encode a gradient of reward dimensions; some neurons respond most to stimuli predicting food rewards while the others respond more to stimuli predicting fluids. We also proposed a possible implementation of the model in the basal ganglia network, and demonstrated how the striatal system can learn values in multiple dimensions, even when dopamine neurons encode mixtures of prediction error from different dimensions. Additionally, the model reproduces the instant generalisation to new physiological states seen in dopamine responses and in behaviour. Our results demonstrate how a simple neural circuit can flexibly guide behaviour according to animals' needs.

One of the key problems the brain faces is inferring the state of the world from a sequence of dynamically changing stimuli, and it is not yet clear how the sensory system achieves this task. A well-established computational framework for describing perceptual processes in the brain is provided by the theory of predictive coding. Although the original proposals of predictive coding have discussed temporal prediction, later work developing this theory mostly focused on static stimuli, and key questions on neural implementation and computational properties of temporal predictive coding networks remain open. Here, we address these questions and present a formulation of the temporal predictive coding model that can be naturally implemented in recurrent networks, in which activity dynamics rely only on local inputs to the neurons, and learning only utilises local Hebbian plasticity. Additionally, we show that temporal predictive coding networks can approximate the performance of the Kalman filter in predicting behaviour of linear systems, and behave as a variant of a Kalman filter which does not track its own subjective posterior variance. Importantly, temporal predictive coding networks can achieve similar accuracy as the Kalman filter without performing complex mathematical operations, but just employing simple computations that can be implemented by biological networks. Moreover, when trained with natural dynamic inputs, we found that temporal predictive coding can produce Gabor-like, motion-sensitive receptive fields resembling those observed in real neurons in visual areas. In addition, we demonstrate how the model can be effectively generalized to nonlinear systems. Overall, models presented in this paper show how biologically plausible circuits can predict future stimuli and may guide research on understanding specific neural circuits in brain areas involved in temporal prediction.

An autoassociative memory model is a function that, given a set of data points, takes as input an arbitrary vector and outputs the most similar data point from the memorized set. However, popular memory models fail to retrieve images even when the corruption is mild and easy to detect for a human evaluator. This is because similarities are evaluated in the raw pixel space, which does not contain any semantic information about the images. This problem can be easily solved by computing similarities in an embedding space instead of the pixel space. We show that an effective way of computing such embeddings is via a network pretrained with a contrastive loss. As the dimension of embedding spaces is often significantly smaller than the pixel space, we also have a faster computation of similarity scores. We test this method on complex datasets such as CIFAR10 and STL10. An additional drawback of current models is the need of storing the whole dataset in the pixel space, which is often extremely large. We relax this condition and propose a class of memory models that only stores low-dimensional semantic embeddings, and uses them to retrieve similar, but not identical, memories. We demonstrate a proof of concept of this method on a simple task on the MNIST dataset.

The computational principles adopted by the hippocampus in associative memory (AM) tasks have been one of the most studied topics in computational and theoretical neuroscience. Recent theories suggested that AM and the predictive activities of the hippocampus could be described within a unitary account, and that predictive coding underlies the computations supporting AM in the hippocampus. Following this theory, a computational model based on classical hierarchical predictive networks was proposed and was shown to perform well in various AM tasks. However, this fully hierarchical model did not incorporate recurrent connections, an architectural component of the CA3 region of the hippocampus that is crucial for AM. This makes the structure of the model inconsistent with the known connectivity of CA3 and classical recurrent models such as Hopfield Networks, which learn the covariance of inputs through their recurrent connections to perform AM. Earlier PC models that learn the covariance information of inputs explicitly via recurrent connections seem to be a solution to these issues. Here, we show that although these models can perform AM, they do it in an implausible and numerically unstable way. Instead, we propose alternatives to these earlier covariance-learning predictive coding networks, which learn the covariance information implicitly and plausibly, and can use dendritic structures to encode prediction errors. We show analytically that our proposed models are perfectly equivalent to the earlier predictive coding model learning covariance explicitly, and encounter no numerical issues when performing AM tasks in practice. We further show that our models can be combined with hierarchical predictive coding networks to model the hippocampo-neocortical interactions. Our models provide a biologically plausible approach to modelling the hippocampal network, pointing to a potential computational mechanism during hippocampal memory formation and recall, which employs both predictive coding and covariance learning based on the recurrent network structure of the hippocampus.

Predictive coding is an influential model of cortical neural activity. It proposes that perceptual beliefs are furnished by sequentially minimising "prediction errors"-the differences between predicted and observed data. Implicit in this proposal is the idea that successful perception requires multiple cycles of neural activity. This is at odds with evidence that several aspects of visual perception-including complex forms of object recognition-arise from an initial "feedforward sweep" that occurs on fast timescales which preclude substantial recurrent activity. Here, we propose that the feedforward sweep can be understood as performing amortized inference (applying a learned function that maps directly from data to beliefs) and recurrent processing can be understood as performing iterative inference (sequentially updating neural activity in order to improve the accuracy of beliefs). We propose a hybrid predictive coding network that combines both iterative and amortized inference in a principled manner by describing both in terms of a dual optimization of a single objective function. We show that the resulting scheme can be implemented in a biologically plausible neural architecture that approximates Bayesian inference utilising local Hebbian update rules. We demonstrate that our hybrid predictive coding model combines the benefits of both amortized and iterative inference-obtaining rapid and computationally cheap perceptual inference for familiar data while maintaining the context-sensitivity, precision, and sample efficiency of iterative inference schemes. Moreover, we show how our model is inherently sensitive to its uncertainty and adaptively balances iterative and amortized inference to obtain accurate beliefs using minimum computational expense. Hybrid predictive coding offers a new perspective on the functional relevance of the feedforward and recurrent activity observed during visual perception and offers novel insights into distinct aspects of visual phenomenology.

A large number of neural network models of associative memory have been proposed in the literature. These include the classical Hopfield networks (HNs), sparse distributed memories (SDMs), and more recently the modern continuous Hopfield networks (MCHNs), which possess close links with self-attention in machine learning. In this paper, we propose a general framework for understanding the operation of such memory networks as a sequence of three operations: similarity, separation, and projection. We derive all these memory models as instances of our general framework with differing similarity and separation functions. We extend the mathematical framework of Krotov et al (2020) to express general associative memory models using neural network dynamics with local computation, and derive a general energy function that is a Lyapunov function of the dynamics. Finally, using our framework, we empirically investigate the capacity of using different similarity functions for these associative memory models, beyond the dot product similarity measure, and demonstrate empirically that Euclidean or Manhattan distance similarity metrics perform substantially better in practice on many tasks, enabling a more robust retrieval and higher memory capacity than existing models.

The backpropagation of error algorithm (BP) used to train deep neural networks has been fundamental to the successes of deep learning. However, it requires sequential backwards updates and non-local computations which make it challenging to parallelize at scale and is unlike how learning works in the brain. Neuroscience-inspired learning algorithms, however, such as \emph{predictive coding} which utilize local learning have the potential to overcome these limitations and advance beyond deep learning technologies in the future. While predictive coding originated in theoretical neuroscience as a model of information processing in the cortex, recent work has developed the idea into a general-purpose algorithm able to train neural networks using only local computations. In this survey, we review works that have contributed to this perspective and demonstrate the close connection between predictive coding and backpropagation in terms of generalization quality, as well as works that highlight the multiple advantages of using predictive coding models over backprop-trained neural networks. Specifically, we show the substantially greater flexibility of predictive coding networks against equivalent deep neural networks, which can function as classifiers, generators, and associative memories simultaneously, and can be defined on arbitrary graph topologies. Finally, we review direct benchmarks of predictive coding networks on machine learning classification tasks, as well as its close connections to control theory and applications in robotics.

Backpropagation (BP) is the most successful and widely used algorithm in deep learning. However, the computations required by BP are challenging to reconcile with known neurobiology. This difficulty has stimulated interest in more biologically plausible alternatives to BP. One such algorithm is the inference learning algorithm (IL). IL trains predictive coding models of neural circuits and has achieved equal performance to BP on supervised and auto-associative tasks. In contrast to BP, however, the mathematical foundations of IL are not well-understood. Here, we develop a novel theoretical framework for IL. Our main result is that IL closely approximates an optimization method known as implicit stochastic gradient descent (implicit SGD), which is distinct from the explicit SGD implemented by BP. Our results further show how the standard implementation of IL can be altered to better approximate implicit SGD. Our novel implementation considerably improves the stability of IL across learning rates, which is consistent with our theory, as a key property of implicit SGD is its stability. We provide extensive simulation results that further support our theoretical interpretations and find IL achieves quicker convergence when trained with mini-batch size one while performing competitively with BP for larger mini-batches when combined with Adam.

A large amount of recent research has the far-reaching goal of finding training methods for deep neural networks that can serve as alternatives to backpropagation~(BP). A prominent example is predictive coding (PC), which is a neuroscience-inspired method that performs inference on hierarchical Gaussian generative models. These methods, however, fail to keep up with modern neural networks, as they are unable to replicate the dynamics of complex layers and activation functions. In this work, we solve this problem by generalizing PC to arbitrary probability distributions, enabling the training of architectures, such as transformers, that are hard to approximate with only Gaussian assumptions. We perform three experimental analyses. First, we study the gap between our method and the standard formulation of PC on multiple toy examples. Second, we test the reconstruction quality on variational autoencoders, where our method reaches the same reconstruction quality as BP. Third, we show that our method allows us to train transformer networks and achieve performance comparable with BP on conditional language models. More broadly, this method allows neuroscience-inspired learning to be applied to multiple domains, since the internal distributions can be flexibly adapted to the data, tasks, and architectures used.

Training with backpropagation (BP) in standard deep learning consists of two main steps: a forward pass that maps a data point to its prediction, and a backward pass that propagates the error of this prediction back through the network. This process is highly effective when the goal is to minimize a specific objective function. However, it does not allow training on networks with cyclic or backward connections. This is an obstacle to reaching brain-like capabilities, as the highly complex heterarchical structure of the neural connections in the neocortex are potentially fundamental for its effectiveness. In this paper, we show how predictive coding (PC), a theory of information processing in the cortex, can be used to perform inference and learning on arbitrary graph topologies. We experimentally show how this formulation, called PC graphs, can be used to flexibly perform different tasks with the same network by simply stimulating specific neurons. This enables the model to be queried on stimuli with different structures, such as partial images, images with labels, or images without labels. We conclude by investigating how the topology of the graph influences the final performance, and comparing against simple baselines trained with BP.

Backpropagation of error (backprop) is a powerful algorithm for training machine learning architectures through end-to-end differentiation. Recently it has been shown that backprop in multilayer perceptrons (MLPs) can be approximated using predictive coding, a biologically plausible process theory of cortical computation that relies solely on local and Hebbian updates. The power of backprop, however, lies not in its instantiation in MLPs but in the concept of automatic differentiation, which allows for the optimization of any differentiable program expressed as a computation graph. Here, we demonstrate that predictive coding converges asymptotically (and in practice, rapidly) to exact backprop gradients on arbitrary computation graphs using only local learning rules. We apply this result to develop a straightforward strategy to translate core machine learning architectures into their predictive coding equivalents. We construct predictive coding convolutional neural networks, recurrent neural networks, and the more complex long short-term memory, which include a nonlayer-like branching internal graph structure and multiplicative interactions. Our models perform equivalently to backprop on challenging machine learning benchmarks while using only local and (mostly) Hebbian plasticity. Our method raises the potential that standard machine learning algorithms could in principle be directly implemented in neural circuitry and may also contribute to the development of completely distributed neuromorphic architectures.

The expected free energy (EFE) is a central quantity in the theory of active inference. It is the quantity that all active inference agents are mandated to minimize through action, and its decomposition into extrinsic and intrinsic value terms is key to the balance of exploration and exploitation that active inference agents evince. Despite its importance, the mathematical origins of this quantity and its relation to the variational free energy (VFE) remain unclear. In this letter, we investigate the origins of the EFE in detail and show that it is not simply "the free energy in the future." We present a functional that we argue is the natural extension of the VFE but actively discourages exploratory behavior, thus demonstrating that exploration does not directly follow from free energy minimization into the future. We then develop a novel objective, the free energy of the expected future (FEEF), which possesses both the epistemic component of the EFE and an intuitive mathematical grounding as the divergence between predicted and desired futures.