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Grid cells in the medial entorhinal cortex (MEC) of the mammalian brain exhibit a strikingly regular hexagonal firing field over space. These cells are learned after birth and are thought to support spatial navigation but also more abstract computations. Although various computational models, including those based on artificial neural networks, have been proposed to explain the formation of grid cells, the process through which the MEC circuit ${\it learns}$ to develop grid cells remains unclear. In this study, we argue that predictive coding, a biologically plausible plasticity rule known to learn visual representations, can also train neural networks to develop hexagonal grid representations from spatial inputs. We demonstrate that grid cells emerge robustly through predictive coding in both static and dynamic environments, and we develop an understanding of this grid cell learning capability by analytically comparing predictive coding with existing models. Our work therefore offers a novel and biologically plausible perspective on the learning mechanisms underlying grid cells. Moreover, it extends the predictive coding theory to the hippocampal formation, suggesting a unified learning algorithm for diverse cortical representations.

In this work, we tackle the problems of efficiency and scalability for predictive coding networks in machine learning. To do so, we first propose a library called PCX, whose focus lies on performance and simplicity, and provides a user-friendly, deep-learning oriented interface. Second, we use PCX to implement a large set of benchmarks for the community to use for their experiments. As most works propose their own tasks and architectures, do not compare one against each other, and focus on small-scale tasks, a simple and fast open-source library adopted by the whole community would address all of these concerns. Third, we perform extensive benchmarks using multiple algorithms, setting new state-of-the-art results in multiple tasks and datasets, as well as highlighting limitations inherent to PC that should be addressed. Thanks to the efficiency of PCX, we are able to analyze larger architectures than commonly used, providing baselines to galvanize community efforts towards one of the main open problems in the field: scalability. The code for PCX is available at \textit{https://github.com/liukidar/pcax}., Comment: 33 pages, 25 figures

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 \emph{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., Comment: 8 Pages, 4 Figures, accepted for publication at ECAI 2023

Forming accurate memory of sequential stimuli is a fundamental function of biological agents. However, the computational mechanism underlying sequential memory in the brain remains unclear. Inspired by neuroscience theories and recent successes in applying predictive coding (PC) to \emph{static} memory tasks, in this work we propose a novel PC-based model for \emph{sequential} memory, called \emph{temporal predictive coding} (tPC). We show that our tPC models can memorize and retrieve sequential inputs accurately with a biologically plausible neural implementation. Importantly, our analytical study reveals that tPC can be viewed as a classical Asymmetric Hopfield Network (AHN) with an implicit statistical whitening process, which leads to more stable performance in sequential memory tasks of structured inputs. Moreover, we find that tPC exhibits properties consistent with behavioral observations and theories in neuroscience, thereby strengthening its biological relevance. Our work establishes a possible computational mechanism underlying sequential memory in the brain that can also be theoretically interpreted using existing memory model frameworks., Comment: 37th Conference on Neural Information Processing Systems (NeurIPS 2023)

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., Comment: Change of title and abstract, that now reflect the version accepted for publication. One co-author also added, that performed the additional experiments

Predictive coding (PC) is an influential theory in computational neuroscience, which argues that the cortex forms unsupervised world models by implementing a hierarchical process of prediction error minimization. PC networks (PCNs) are trained in two phases. First, neural activities are updated to optimize the network's response to external stimuli. Second, synaptic weights are updated to consolidate this change in activity -- an algorithm called \emph{prospective configuration}. While previous work has shown how in various limits, PCNs can be found to approximate backpropagation (BP), recent work has demonstrated that PCNs operating in this standard regime, which does not approximate BP, nevertheless obtain competitive training and generalization performance to BP-trained networks while outperforming them on tasks such as online, few-shot, and continual learning, where brains are known to excel. Despite this promising empirical performance, little is understood theoretically about the properties and dynamics of PCNs in this regime. In this paper, we provide a comprehensive theoretical analysis of the properties of PCNs trained with prospective configuration. We first derive analytical results concerning the inference equilibrium for PCNs and a previously unknown close connection relationship to target propagation (TP). Secondly, we provide a theoretical analysis of learning in PCNs as a variant of generalized expectation-maximization and use that to prove the convergence of PCNs to critical points of the BP loss function, thus showing that deep PCNs can, in theory, achieve the same generalization performance as BP, while maintaining their unique advantages., Comment: 21/07/22 initial upload (finally); 03/08/22 revisions

How the brain performs credit assignment is a fundamental unsolved problem in neuroscience. Many `biologically plausible' algorithms have been proposed, which compute gradients that approximate those computed by backpropagation (BP), and which operate in ways that more closely satisfy the constraints imposed by neural circuitry. Many such algorithms utilize the framework of energy-based models (EBMs), in which all free variables in the model are optimized to minimize a global energy function. However, in the literature, these algorithms exist in isolation and no unified theory exists linking them together. Here, we provide a comprehensive theory of the conditions under which EBMs can approximate BP, which lets us unify many of the BP approximation results in the literature (namely, predictive coding, equilibrium propagation, and contrastive Hebbian learning) and demonstrate that their approximation to BP arises from a simple and general mathematical property of EBMs at free-phase equilibrium. This property can then be exploited in different ways with different energy functions, and these specific choices yield a family of BP-approximating algorithms, which both includes the known results in the literature and can be used to derive new ones., Comment: 31/05/22 initial upload; 22/06/22 change corresponding author; 03/08/22 revisions

The backpropagation of error algorithm used to train deep neural networks has been fundamental to the successes of deep learning. However, it requires sequential backward 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 current deep learning technologies. 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 theoretical connections between predictive coding and backpropagation, as well as works that highlight the multiple advantages of using predictive coding models over backpropagation-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., Comment: 18/02/22 initial upload

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 possesses 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 only second-order interactions between neurons, 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., Comment: 09/02/22 initial upload; 17/06/222 camera ready version upload

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, and investigate how the topology of the graph influences the final performance. We conclude by comparing against simple baselines trained~with~BP., Comment: 15 pages, 11 figures

Associative memories in the brain receive and store patterns of activity registered by the sensory neurons, and are able to retrieve them when necessary. Due to their importance in human intelligence, computational models of associative memories have been developed for several decades now. They include autoassociative memories, which allow for storing data points and retrieving a stored data point $s$ when provided with a noisy or partial variant of $s$, and heteroassociative memories, able to store and recall multi-modal data. In this paper, we present a novel neural model for realizing associative memories, based on a hierarchical generative network that receives external stimuli via sensory neurons. This model is trained using predictive coding, an error-based learning algorithm inspired by information processing in the cortex. To test the capabilities of this model, we perform multiple retrieval experiments from both corrupted and incomplete data points. In an extensive comparison, we show that this new model outperforms in retrieval accuracy and robustness popular associative memory models, such as autoencoders trained via backpropagation, and modern Hopfield networks. In particular, in completing partial data points, our model achieves remarkable results on natural image datasets, such as ImageNet, with a surprisingly high accuracy, even when only a tiny fraction of pixels of the original images is presented. Furthermore, we show that this method is able to handle multi-modal data, retrieving images from descriptions, and vice versa. We conclude by discussing the possible impact of this work in the neuroscience community, by showing that our model provides a plausible framework to study learning and retrieval of memories in the brain, as it closely mimics the behavior of the hippocampus as a memory index and generative model., Comment: 24 pages, 18 figures

Circadian and other physiological rhythms play a key role in both normal homeostasis and disease processes. Such is the case of circadian and infradian seizure patterns observed in epilepsy. In this paper we explore a new implantable stimulator that implements chronotherapy as a feedforward input to supplement both open-loop and closed-loop methods. This integrated algorithm allows for stimulation to be adjusted to the ultradian, circadian and infradian patterns observed in patients through slowly-varying temporal adjustments of stimulation and algorithm sub-components, while also enabling adaption of stimulation based on immediate physiological needs such as a breakthrough seizure or change of posture. Embedded physiological sensors in the stimulator can be used to refine the baseline stimulation circadian pattern as a "digital zeitgeber". This approach is tested on a canine with severe drug-resistant idiopathic generalized epilepsy exhibiting a diurnal pattern correlated with sleep-wake cycles. Prior to implantation, the canine's cluster seizures evolved to status epilepticus (SE) and required emergency pharmacological intervention. The cranially-mounted system was fully-implanted bilaterally into the centromedian nucleus of the thalamus. Using time-based modulation, thalamocortical rhythm-specific tuning of frequency parameters as well as fast-adaptive modes based on activity, the canine experienced no further SE events post-implant as of the time of writing (seven months). Importantly, no significant cluster seizures have been observed either, allowing the reduction of rescue medication. The use of chronotherapy as a feedforward signal to augment adaptive neurostimulators could prove a useful method in conditions where sensitivity to temporal patterns are characteristics of the disease state, providing a novel mechanism for tailoring a more patient-specific therapy approach., Comment: 6 text pages, 4 main figures, Fig 1 has 4 panels, Fig 2 has 3 panels, Fig 4 has 2 panels. 2 text pages of supplementary material, 1 supplementary figure

Deep learning has redefined the field of artificial intelligence (AI) thanks to the rise of artificial neural networks, which are architectures inspired by their neurological counterpart in the brain. Through the years, this dualism between AI and neuroscience has brought immense benefits to both fields, allowing neural networks to be used in dozens of applications. These networks use an efficient implementation of reverse differentiation, called backpropagation (BP). This algorithm, however, is often criticized for its biological implausibility (e.g., lack of local update rules for the parameters). Therefore, biologically plausible learning methods that rely on predictive coding (PC), a framework for describing information processing in the brain, are increasingly studied. Recent works prove that these methods can approximate BP up to a certain margin on multilayer perceptrons (MLPs), and asymptotically on any other complex model, and that zero-divergence inference learning (Z-IL), a variant of PC, is able to exactly implement BP on MLPs. However, the recent literature shows also that there is no biologically plausible method yet that can exactly replicate the weight update of BP on complex models. To fill this gap, in this paper, we generalize (PC and) Z-IL by directly defining them on computational graphs, and show that it can perform exact reverse differentiation. What results is the first biologically plausible algorithm that is equivalent to BP in the way of updating parameters on any neural network, providing a bridge between the interdisciplinary research of neuroscience and deep learning., Comment: 17 pages, 5 figures

Predictive coding networks (PCNs) are an influential model for information processing in the brain. They have appealing theoretical interpretations and offer a single mechanism that accounts for diverse perceptual phenomena of the brain. On the other hand, backpropagation (BP) is commonly regarded to be the most successful learning method in modern machine learning. Thus, it is exciting that recent work formulates inference learning (IL) that trains PCNs to approximate BP. However, there are several remaining critical issues: (i) IL is an approximation to BP with unrealistic/non-trivial requirements, (ii) IL approximates BP in single-step weight updates; whether it leads to the same point as BP after the weight updates are conducted for more steps is unknown, and (iii) IL is computationally significantly more costly than BP. To solve these issues, a variant of IL that is strictly equivalent to BP in fully connected networks has been proposed. In this work, we build on this result by showing that it also holds for more complex architectures, namely, convolutional neural networks and (many-to-one) recurrent neural networks. To our knowledge, we are the first to show that a biologically plausible algorithm is able to exactly replicate the accuracy of BP on such complex architectures, bridging the existing gap between IL and BP, and setting an unprecedented performance for PCNs, which can now be considered as efficient alternatives to BP., Comment: 18 pages, 3 figures

How the brain co-ordinates the actions of distant regions in an efficient manner is an open problem. Many believe that cross-frequency coupling between the amplitude of high frequency local field potential oscillations in one region and the phase of lower frequency signals in another may form a possible mechanism. This work provides a preliminary study from both an experimental and theoretical viewpoint, concentrating on possible coupling between the hippocampus and pre-frontal cortex in rats during tasks involving working memory, spatial navigation and decision making processes. Attempts to search for such coupling events are made using newly developed MATLAB scripts. This leads to the discovery of increased envelope-to-signal correlation (ESC) between the 1-10 Hz hippocampus theta and 30-40 Hz pre-fontal cortex gamma bands when a choice turn is approached during a T-maze task. From a theoretical perspective, a standard connectionist modelling approach is extended to allow for the formation of oscillations. Although detrimental to overall average task performance, this did lead to a reduced increase in performance variation as noise was increased, when compared to a standard non-oscillating model.