Daily Papers

Reservoir computers (RCs) provide a computationally efficient alternative to deep learning while also offering a framework for incorporating brain-inspired computational principles. By using an internal neural network with random, fixed connections-the 'reservoir'-and training only the output weights, RCs simplify the training process but remain sensitive to the choice of hyperparameters that govern activation functions and network architecture. Moreover, typical RC implementations overlook a critical aspect of neuronal dynamics: the balance between excitatory and inhibitory (E-I) signals, which is essential for robust brain function. We show that RCs characteristically perform best in balanced or slightly over-inhibited regimes, outperforming excitation-dominated ones. To reduce the need for precise hyperparameter tuning, we introduce a self-adapting mechanism that locally adjusts E/I balance to achieve target neuronal firing rates, improving performance by up to 130% in tasks like memory capacity and time series prediction compared with globally tuned RCs. Incorporating brain-inspired heterogeneity in target neuronal firing rates further reduces the need for fine-tuning hyperparameters and enables RCs to excel across linear and non-linear tasks. These results support a shift from static optimization to dynamic adaptation in reservoir design, demonstrating how brain-inspired mechanisms improve RC performance and robustness while deepening our understanding of neural computation.

This study introduces a novel approach by replacing the traditional perceptron neuron model with a biologically inspired probabilistic meta neuron, where the internal neuron parameters are jointly learned, leading to improved classification accuracy of spiking neural networks (SNNs). To validate this innovation, we implement and compare two SNN architectures: one based on standard leaky integrate-and-fire (LIF) neurons and another utilizing the proposed probabilistic meta neuron model. As a second key contribution, we present a new biologically inspired classification framework that uniquely integrates SNNs with Lempel-Ziv complexity (LZC) a measure closely related to entropy rate. By combining the temporal precision and biological plausibility of SNNs with the capacity of LZC to capture structural regularity, the proposed approach enables efficient and interpretable classification of spatiotemporal neural data, an aspect not addressed in existing works. We consider learning algorithms such as backpropagation, spike-timing-dependent plasticity (STDP), and the Tempotron learning rule. To explore neural dynamics, we use Poisson processes to model neuronal spike trains, a well-established method for simulating the stochastic firing behavior of biological neurons. Our results reveal that depending on the training method, the classifier's efficiency can improve by up to 11.00%, highlighting the advantage of learning additional neuron parameters beyond the traditional focus on weighted inputs alone.

Recently, interest has grown in exploring the hypothesis that neural activity conveys information through precise spiking motifs. To investigate this phenomenon, various algorithms have been proposed to detect such motifs in Single Unit Activity (SUA) recorded from populations of neurons. In this study, we present a novel detection model based on the inversion of a generative model of raster plot synthesis. Using this generative model, we derive an optimal detection procedure that takes the form of logistic regression combined with temporal convolution. A key advantage of this model is its differentiability, which allows us to formulate a supervised learning approach using a gradient descent on the binary cross-entropy loss. To assess the model's ability to detect spiking motifs in synthetic data, we first perform numerical evaluations. This analysis highlights the advantages of using spiking motifs over traditional firing rate based population codes. We then successfully demonstrate that our learning method can recover synthetically generated spiking motifs, indicating its potential for further applications. In the future, we aim to extend this method to real neurobiological data, where the ground truth is unknown, to explore and detect spiking motifs in a more natural and biologically relevant context.

Compared with rate-based artificial neural networks, Spiking Neural Networks (SNN) provide a more biological plausible model for the brain. But how they perform supervised learning remains elusive. Inspired by recent works of Bengio et al., we propose a supervised learning algorithm based on Spike-Timing Dependent Plasticity (STDP) for a hierarchical SNN consisting of Leaky Integrate-and-fire (LIF) neurons. A time window is designed for the presynaptic neuron and only the spikes in this window take part in the STDP updating process. The model is trained on the MNIST dataset. The classification accuracy approach that of a Multilayer Perceptron (MLP) with similar architecture trained by the standard back-propagation algorithm.

The Spiking Neural Network (SNN) has attracted more and more attention recently. It adopts binary spike signals to transmit information. Benefitting from the information passing paradigm of SNNs, the multiplications of activations and weights can be replaced by additions, which are more energy-efficient. However, its ``Hard Reset" mechanism for the firing activity would ignore the difference among membrane potentials when the membrane potential is above the firing threshold, causing information loss. Meanwhile, quantifying the membrane potential to 0/1 spikes at the firing instants will inevitably introduce the quantization error thus bringing about information loss too. To address these problems, we propose to use the ``Soft Reset" mechanism for the supervised training-based SNNs, which will drive the membrane potential to a dynamic reset potential according to its magnitude, and Membrane Potential Rectifier (MPR) to reduce the quantization error via redistributing the membrane potential to a range close to the spikes. Results show that the SNNs with the ``Soft Reset" mechanism and MPR outperform their vanilla counterparts on both static and dynamic datasets.

In recent years, deep learning has been a revolution in the field of machine learning, for computer vision in particular. In this approach, a deep (multilayer) artificial neural network (ANN) is trained in a supervised manner using backpropagation. Huge amounts of labeled examples are required, but the resulting classification accuracy is truly impressive, sometimes outperforming humans. Neurons in an ANN are characterized by a single, static, continuous-valued activation. Yet biological neurons use discrete spikes to compute and transmit information, and the spike times, in addition to the spike rates, matter. Spiking neural networks (SNNs) are thus more biologically realistic than ANNs, and arguably the only viable option if one wants to understand how the brain computes. SNNs are also more hardware friendly and energy-efficient than ANNs, and are thus appealing for technology, especially for portable devices. However, training deep SNNs remains a challenge. Spiking neurons' transfer function is usually non-differentiable, which prevents using backpropagation. Here we review recent supervised and unsupervised methods to train deep SNNs, and compare them in terms of accuracy, but also computational cost and hardware friendliness. The emerging picture is that SNNs still lag behind ANNs in terms of accuracy, but the gap is decreasing, and can even vanish on some tasks, while the SNNs typically require much fewer operations.

We propose a formal mathematical model for sparse representations and active dendrites in neocortex. Our model is inspired by recent experimental findings on active dendritic processing and NMDA spikes in pyramidal neurons. These experimental and modeling studies suggest that the basic unit of pattern memory in the neocortex is instantiated by small clusters of synapses operated on by localized non-linear dendritic processes. We derive a number of scaling laws that characterize the accuracy of such dendrites in detecting activation patterns in a neuronal population under adverse conditions. We introduce the union property which shows that synapses for multiple patterns can be randomly mixed together within a segment and still lead to highly accurate recognition. We describe simulation results that provide further insight into sparse representations as well as two primary results. First we show that pattern recognition by a neuron with active dendrites can be extremely accurate and robust with high dimensional sparse inputs even when using a tiny number of synapses to recognize large patterns. Second, equations representing recognition accuracy of a dendrite predict optimal NMDA spiking thresholds under a generous set of assumptions. The prediction tightly matches NMDA spiking thresholds measured in the literature. Our model matches many of the known properties of pyramidal neurons. As such the theory provides a mathematical framework for understanding the benefits and limits of sparse representations in cortical networks.

Why do neurons communicate through spikes? By definition, spikes are all-or-none neural events which occur at continuous times. In other words, spikes are on one side binary, existing or not without further details, and on the other can occur at any asynchronous time, without the need for a centralized clock. This stands in stark contrast to the analog representation of values and the discretized timing classically used in digital processing and at the base of modern-day neural networks. As neural systems almost systematically use this so-called event-based representation in the living world, a better understanding of this phenomenon remains a fundamental challenge in neurobiology in order to better interpret the profusion of recorded data. With the growing need for intelligent embedded systems, it also emerges as a new computing paradigm to enable the efficient operation of a new class of sensors and event-based computers, called neuromorphic, which could enable significant gains in computation time and energy consumption -- a major societal issue in the era of the digital economy and global warming. In this review paper, we provide evidence from biology, theory and engineering that the precise timing of spikes plays a crucial role in our understanding of the efficiency of neural networks.

The precise timing of spikes emitted by neurons plays a crucial role in shaping the response of efferent biological neurons. This temporal dimension of neural activity holds significant importance in understanding information processing in neurobiology, especially for the performance of neuromorphic hardware, such as event-based cameras. Nonetheless, many artificial neural models disregard this critical temporal dimension of neural activity. In this study, we present a model designed to efficiently detect temporal spiking motifs using a layer of spiking neurons equipped with heterogeneous synaptic delays. Our model capitalizes on the diverse synaptic delays present on the dendritic tree, enabling specific arrangements of temporally precise synaptic inputs to synchronize upon reaching the basal dendritic tree. We formalize this process as a time-invariant logistic regression, which can be trained using labeled data. To demonstrate its practical efficacy, we apply the model to naturalistic videos transformed into event streams, simulating the output of the biological retina or event-based cameras. To evaluate the robustness of the model in detecting visual motion, we conduct experiments by selectively pruning weights and demonstrate that the model remains efficient even under significantly reduced workloads. In conclusion, by providing a comprehensive, event-driven computational building block, the incorporation of heterogeneous delays has the potential to greatly improve the performance of future spiking neural network algorithms, particularly in the context of neuromorphic chips.

Traveling waves of neural activity have been observed throughout the brain at a diversity of regions and scales; however, their precise computational role is still debated. One physically inspired hypothesis suggests that the cortical sheet may act like a wave-propagating system capable of invertibly storing a short-term memory of sequential stimuli through induced waves traveling across the cortical surface, and indeed many experimental results from neuroscience correlate wave activity with memory tasks. To date, however, the computational implications of this idea have remained hypothetical due to the lack of a simple recurrent neural network architecture capable of exhibiting such waves. In this work, we introduce a model to fill this gap, which we denote the Wave-RNN (wRNN), and demonstrate how such an architecture indeed efficiently encodes the recent past through a suite of synthetic memory tasks where wRNNs learn faster and reach significantly lower error than wave-free counterparts. We further explore the implications of this memory storage system on more complex sequence modeling tasks such as sequential image classification and find that wave-based models not only again outperform comparable wave-free RNNs while using significantly fewer parameters, but additionally perform comparably to more complex gated architectures such as LSTMs and GRUs.

Recent years have seen significant progress in developing spiking neural networks (SNNs) as a potential solution to the energy challenges posed by conventional artificial neural networks (ANNs). However, our theoretical understanding of SNNs remains relatively limited compared to the ever-growing body of literature on ANNs. In this paper, we study a discrete-time model of SNNs based on leaky integrate-and-fire (LIF) neurons, referred to as discrete-time LIF-SNNs, a widely used framework that still lacks solid theoretical foundations. We demonstrate that discrete-time LIF-SNNs with static inputs and outputs realize piecewise constant functions defined on polyhedral regions, and more importantly, we quantify the network size required to approximate continuous functions. Moreover, we investigate the impact of latency (number of time steps) and depth (number of layers) on the complexity of the input space partitioning induced by discrete-time LIF-SNNs. Our analysis highlights the importance of latency and contrasts these networks with ANNs employing piecewise linear activation functions. Finally, we present numerical experiments to support our theoretical findings.

State-of-the-art systems neuroscience experiments yield large-scale multimodal data, and these data sets require new tools for analysis. Inspired by the success of large pretrained models in vision and language domains, we reframe the analysis of large-scale, cellular-resolution neuronal spiking data into an autoregressive spatiotemporal generation problem. Neuroformer is a multimodal, multitask generative pretrained transformer (GPT) model that is specifically designed to handle the intricacies of data in systems neuroscience. It scales linearly with feature size, can process an arbitrary number of modalities, and is adaptable to downstream tasks, such as predicting behavior. We first trained Neuroformer on simulated datasets, and found that it both accurately predicted simulated neuronal circuit activity, and also intrinsically inferred the underlying neural circuit connectivity, including direction. When pretrained to decode neural responses, the model predicted the behavior of a mouse with only few-shot fine-tuning, suggesting that the model begins learning how to do so directly from the neural representations themselves, without any explicit supervision. We used an ablation study to show that joint training on neuronal responses and behavior boosted performance, highlighting the model's ability to associate behavioral and neural representations in an unsupervised manner. These findings show that Neuroformer can analyze neural datasets and their emergent properties, informing the development of models and hypotheses associated with the brain.

Nowadays, neural networks act as a synonym for artificial intelligence. Present neural network models, although remarkably powerful, are inefficient both in terms of data and energy. Several alternative forms of neural networks have been proposed to address some of these problems. Specifically, spiking neural networks are suitable for efficient hardware implementations. However, effective learning algorithms for spiking networks remain elusive, although it is suspected that effective plasticity mechanisms could alleviate the problem of data efficiency. Here, we present a new framework for spiking neural networks - Spark - built upon the idea of modular design, from simple components to entire models. The aim of this framework is to provide an efficient and streamlined pipeline for spiking neural networks. We showcase this framework by solving the sparse-reward cartpole problem with simple plasticity mechanisms. We hope that a framework compatible with traditional ML pipelines may accelerate research in the area, specifically for continuous and unbatched learning, akin to the one animals exhibit.

Recurrent Neural Networks (RNNs) are popular models of brain function. The typical training strategy is to adjust their input-output behavior so that it matches that of the biological circuit of interest. Even though this strategy ensures that the biological and artificial networks perform the same computational task, it does not guarantee that their internal activity dynamics match. This suggests that the trained RNNs might end up performing the task employing a different internal computational mechanism, which would make them a suboptimal model of the biological circuit. In this work, we introduce a novel training strategy that allows learning not only the input-output behavior of an RNN but also its internal network dynamics, based on sparse neural recordings. We test the proposed method by training an RNN to simultaneously reproduce internal dynamics and output signals of a physiologically-inspired neural model. Specifically, this model generates the multiphasic muscle-like activity patterns typically observed during the execution of reaching movements, based on the oscillatory activation patterns concurrently observed in the motor cortex. Remarkably, we show that the reproduction of the internal dynamics is successful even when the training algorithm relies on the activities of a small subset of neurons sampled from the biological network. Furthermore, we show that training the RNNs with this method significantly improves their generalization performance. Overall, our results suggest that the proposed method is suitable for building powerful functional RNN models, which automatically capture important computational properties of the biological circuit of interest from sparse neural recordings.

Modern neural recording techniques allow neuroscientists to obtain spiking activity of multiple neurons from different brain regions over long time periods, which requires new statistical methods to be developed for understanding structure of the large-scale data. In this paper, we develop a bi-clustering method to cluster the neural spiking activity spatially and temporally, according to their low-dimensional latent structures. The spatial (neuron) clusters are defined by the latent trajectories within each neural population, while the temporal (state) clusters are defined by (populationally) synchronous local linear dynamics shared with different periods. To flexibly extract the bi-clustering structure, we build the model non-parametrically, and develop an efficient Markov chain Monte Carlo (MCMC) algorithm to sample the posterior distributions of model parameters. Validating our proposed MCMC algorithm through simulations, we find the method can recover unknown parameters and true bi-clustering structures successfully. We then apply the proposed bi-clustering method to multi-regional neural recordings under different experiment settings, where we find that simultaneously considering latent trajectories and spatial-temporal clustering structures can provide us with a more accurate and interpretable result. Overall, the proposed method provides scientific insights for large-scale (counting) time series with elongated recording periods, and it can potentially have application beyond neuroscience.

It has long been known in both neuroscience and AI that ``binding'' between neurons leads to a form of competitive learning where representations are compressed in order to represent more abstract concepts in deeper layers of the network. More recently, it was also hypothesized that dynamic (spatiotemporal) representations play an important role in both neuroscience and AI. Building on these ideas, we introduce Artificial Kuramoto Oscillatory Neurons (AKOrN) as a dynamical alternative to threshold units, which can be combined with arbitrary connectivity designs such as fully connected, convolutional, or attentive mechanisms. Our generalized Kuramoto updates bind neurons together through their synchronization dynamics. We show that this idea provides performance improvements across a wide spectrum of tasks such as unsupervised object discovery, adversarial robustness, calibrated uncertainty quantification, and reasoning. We believe that these empirical results show the importance of rethinking our assumptions at the most basic neuronal level of neural representation, and in particular show the importance of dynamical representations.

The out-of-distribution (OOD) problem generally arises when neural networks encounter data that significantly deviates from the training data distribution, i.e., in-distribution (InD). In this paper, we study the OOD problem from a neuron activation view. We first formulate neuron activation states by considering both the neuron output and its influence on model decisions. Then, to characterize the relationship between neurons and OOD issues, we introduce the neuron activation coverage (NAC) -- a simple measure for neuron behaviors under InD data. Leveraging our NAC, we show that 1) InD and OOD inputs can be largely separated based on the neuron behavior, which significantly eases the OOD detection problem and beats the 21 previous methods over three benchmarks (CIFAR-10, CIFAR-100, and ImageNet-1K). 2) a positive correlation between NAC and model generalization ability consistently holds across architectures and datasets, which enables a NAC-based criterion for evaluating model robustness. Compared to prevalent InD validation criteria, we show that NAC not only can select more robust models, but also has a stronger correlation with OOD test performance.

The human brain is autonomously active, being characterized by a self-sustained neural activity which would be present even in the absence of external sensory stimuli. Here we study the interrelation between the self-sustained activity in autonomously active recurrent neural nets and external sensory stimuli. There is no a priori semantical relation between the influx of external stimuli and the patterns generated internally by the autonomous and ongoing brain dynamics. The question then arises when and how are semantic correlations between internal and external dynamical processes learned and built up? We study this problem within the paradigm of transient state dynamics for the neural activity in recurrent neural nets, i.e. for an autonomous neural activity characterized by an infinite time-series of transiently stable attractor states. We propose that external stimuli will be relevant during the sensitive periods, {\it viz} the transition period between one transient state and the subsequent semi-stable attractor. A diffusive learning signal is generated unsupervised whenever the stimulus influences the internal dynamics qualitatively. For testing we have presented to the model system stimuli corresponding to the bars and stripes problem. We found that the system performs a non-linear independent component analysis on its own, being continuously and autonomously active. This emergent cognitive capability results here from a general principle for the neural dynamics, the competition between neural ensembles.

As one of the energy-efficient alternatives of conventional neural networks (CNNs), spiking neural networks (SNNs) have gained more and more interest recently. To train the deep models, some effective batch normalization (BN) techniques are proposed in SNNs. All these BNs are suggested to be used after the convolution layer as usually doing in CNNs. However, the spiking neuron is much more complex with the spatio-temporal dynamics. The regulated data flow after the BN layer will be disturbed again by the membrane potential updating operation before the firing function, i.e., the nonlinear activation. Therefore, we advocate adding another BN layer before the firing function to normalize the membrane potential again, called MPBN. To eliminate the induced time cost of MPBN, we also propose a training-inference-decoupled re-parameterization technique to fold the trained MPBN into the firing threshold. With the re-parameterization technique, the MPBN will not introduce any extra time burden in the inference. Furthermore, the MPBN can also adopt the element-wised form, while these BNs after the convolution layer can only use the channel-wised form. Experimental results show that the proposed MPBN performs well on both popular non-spiking static and neuromorphic datasets. Our code is open-sourced at https://github.com/yfguo91/MPBN{MPBN}.

High-resolution brain imaging can now capture not just synapse locations but their molecular composition, with the cost of such mapping falling exponentially. Yet such ultrastructural data has so far told us little about local neuronal physiology - specifically, the parameters (e.g., synaptic efficacies, local conductances) that govern neural dynamics. We propose to translate molecularly annotated ultrastructure into physiology, introducing the concept of an ultrastructure-to-dynamics compiler: a learned mapping from molecularly annotated ultrastructure to simulator-ready, uncertainty-aware physiological parameters. The requirement is paired training data, with jointly acquired ultrastructure from imaging, and dynamical responses to perturbations from physiological experiments. With this data we can train models that predict local physiology directly from structure. Such a compiler would support biophysical simulations by turning anatomical maps into models of circuit dynamics, shifting structure-to-function from a descriptive program to a predictive one and opening routes to understanding neural computation and forecasting intervention effects.

We show that in a drive-response coupling framework extreme events are suppressed in the response system by the dominance of a single driving signal. We validate this approach across three distinct response network topologies, namely (i) a pair of coupled neurons, (ii) a monolayer network of N coupled neurons and (iii) a two-layer multiplex network each composed of FitzHugh-Nagumo neuronal units. The response networks inherently exhibit extreme events. Our results demonstrate that influencing just one neuron in the response network with an appropriately tuned driving signal is sufficient to control extreme events across all three configurations. In the two-neuron case, suppression of extreme events occurs due to the breaking of phase-locking between the driving neuron and the targeted response neuron. In the case of monolayer and multiplex networks, suppression of extreme events results from the disruption of protoevent frequency dynamics and a subsequent frequency decoupling of the driven neuron from the rest of the network. We also observe that when the size of the neurons in response network connected to the drive increases, the onset of control occurs earlier indicating a scaling advantage of the method.

Recent advancements in large language models (LLMs) with billions of parameters have improved performance in various applications, but their inference processes demand significant energy and computational resources. In contrast, the human brain, with approximately 86 billion neurons, is much more energy-efficient than LLMs with similar parameters. Inspired by this, we redesign 7sim70 billion parameter LLMs using bio-plausible spiking mechanisms, emulating the efficient behavior of the human brain. We propose the first spiking large language model, SpikeLLM. Coupled with the proposed model, two essential approaches are proposed to improve spike training efficiency: Generalized Integrate-and-Fire (GIF) neurons to compress spike length from T to T{L} log_2 L bits, and an Optimal Brain Spiking framework to divide outlier channels and allocate different T for GIF neurons, which further compresses spike length to approximate log_2T bits. The necessity of spike-driven LLM is proved by comparison with quantized LLMs with similar operations. In the OmniQuant pipeline, SpikeLLM reduces 11.01% WikiText2 perplexity and improves 2.55% accuracy of common scene reasoning on a LLAMA-7B W4A4 model. In the GPTQ pipeline, SpikeLLM achieves direct additive in linear layers, significantly exceeding PB-LLMs.

Artificial Neural Networks (ANN) have gained significant popularity thanks to their ability to learn using the well-known backpropagation algorithm. Conversely, Spiking Neural Networks (SNNs), despite having broader capabilities than ANNs, have always posed challenges in the training phase. This paper shows a new method to perform supervised learning on SNNs, using Spike Timing Dependent Plasticity (STDP) and homeostasis, aiming at training the network to identify spatial patterns. Spatial patterns refer to spike patterns without a time component, where all spike events occur simultaneously. The method is tested using the SpiNNaker digital architecture. A SNN is trained to recognise one or multiple patterns and performance metrics are extracted to measure the performance of the network. Some considerations are drawn from the results showing that, in the case of a single trained pattern, the network behaves as the ideal detector, with 100% accuracy in detecting the trained pattern. However, as the number of trained patterns on a single network increases, the accuracy of identification is linked to the similarities between these patterns. This method of training an SNN to detect spatial patterns may be applied to pattern recognition in static images or traffic analysis in computer networks, where each network packet represents a spatial pattern. It will be stipulated that the homeostatic factor may enable the network to detect patterns with some degree of similarity, rather than only perfectly matching patterns.The principles outlined in this article serve as the fundamental building blocks for more complex systems that utilise both spatial and temporal patterns by converting specific features of input signals into spikes.One example of such a system is a computer network packet classifier, tasked with real-time identification of packet streams based on features within the packet content

This article is about the neural conundrum behind the slowness of human behavior. The information throughput of a human being is about 10 bits/s. In comparison, our sensory systems gather data at ~10^9 bits/s. The stark contrast between these numbers remains unexplained and touches on fundamental aspects of brain function: What neural substrate sets this speed limit on the pace of our existence? Why does the brain need billions of neurons to process 10 bits/s? Why can we only think about one thing at a time? The brain seems to operate in two distinct modes: the "outer" brain handles fast high-dimensional sensory and motor signals, whereas the "inner" brain processes the reduced few bits needed to control behavior. Plausible explanations exist for the large neuron numbers in the outer brain, but not for the inner brain, and we propose new research directions to remedy this.

Spiking neural networks (SNNs) are well suited for spatio-temporal learning and implementations on energy-efficient event-driven neuromorphic processors. However, existing SNN error backpropagation (BP) methods lack proper handling of spiking discontinuities and suffer from low performance compared with the BP methods for traditional artificial neural networks. In addition, a large number of time steps are typically required to achieve decent performance, leading to high latency and rendering spike-based computation unscalable to deep architectures. We present a novel Temporal Spike Sequence Learning Backpropagation (TSSL-BP) method for training deep SNNs, which breaks down error backpropagation across two types of inter-neuron and intra-neuron dependencies and leads to improved temporal learning precision. It captures inter-neuron dependencies through presynaptic firing times by considering the all-or-none characteristics of firing activities and captures intra-neuron dependencies by handling the internal evolution of each neuronal state in time. TSSL-BP efficiently trains deep SNNs within a much shortened temporal window of a few steps while improving the accuracy for various image classification datasets including CIFAR10.

Legged locomotion is a challenging task in the field of robotics but a rather simple one in nature. This motivates the use of biological methodologies as solutions to this problem. Central pattern generators are neural networks that are thought to be responsible for locomotion in humans and some animal species. As for robotics, many attempts were made to reproduce such systems and use them for a similar goal. One interesting design model is based on spiking neural networks. This model is the main focus of this work, as its contribution is not limited to engineering but also applicable to neuroscience. This paper introduces a new general framework for building central pattern generators that are task-independent, biologically plausible, and rely on learning methods. The abilities and properties of the presented approach are not only evaluated in simulation but also in a robotic experiment. The results are very promising as the used robot was able to perform stable walking at different speeds and to change speed within the same gait cycle.

Models of neural responses to stimuli with complex spatiotemporal correlation structure often assume that neurons are only selective for a small number of linear projections of a potentially high-dimensional input. Here we explore recent modeling approaches where the neural response depends on the quadratic form of the input rather than on its linear projection, that is, the neuron is sensitive to the local covariance structure of the signal preceding the spike. To infer this quadratic dependence in the presence of arbitrary (e.g. naturalistic) stimulus distribution, we review several inference methods, focussing in particular on two information-theory-based approaches (maximization of stimulus energy or of noise entropy) and a likelihood-based approach (Bayesian spike-triggered covariance, extensions of generalized linear models). We analyze the formal connection between the likelihood-based and information-based approaches to show how they lead to consistent inference. We demonstrate the practical feasibility of these procedures by using model neurons responding to a flickering variance stimulus.

The current article introduces a supervised learning algorithm for multilayer spiking neural networks. The algorithm presented here overcomes some limitations of existing learning algorithms as it can be applied to neurons firing multiple spikes and it can in principle be applied to any linearisable neuron model. The algorithm is applied successfully to various benchmarks, such as the XOR problem and the Iris data set, as well as complex classifications problems. The simulations also show the flexibility of this supervised learning algorithm which permits different encodings of the spike timing patterns, including precise spike trains encoding.

The neurons of artificial neural networks were originally invented when much less was known about biological neurons than is known today. Our work explores a modification to the core neuron unit to make it more parallel to a biological neuron. The modification is made with the knowledge that biological dendrites are not simply passive activation funnels, but also compute complex non-linear functions as they transmit activation to the cell body. The paper explores a novel system of "Perforated" backpropagation empowering the artificial neurons of deep neural networks to achieve better performance coding for the same features they coded for in the original architecture. After an initial network training phase, additional "Dendrite Nodes" are added to the network and separately trained with a different objective: to correlate their output with the remaining error of the original neurons. The trained Dendrite Nodes are then frozen, and the original neurons are further trained, now taking into account the additional error signals provided by the Dendrite Nodes. The cycle of training the original neurons and then adding and training Dendrite Nodes can be repeated several times until satisfactory performance is achieved. Our algorithm was successfully added to modern state-of-the-art PyTorch networks across multiple domains, improving upon original accuracies and allowing for significant model compression without a loss in accuracy.

Learning and decision making in the brain are key processes critical to survival, and yet are processes implemented by non-ideal biological building blocks which can impose significant error. We explore quantitatively how the brain might cope with this inherent source of error by taking advantage of two ubiquitous mechanisms, redundancy and synchronization. In particular we consider a neural process whose goal is to learn a decision function by implementing a nonlinear gradient dynamics. The dynamics, however, are assumed to be corrupted by perturbations modeling the error which might be incurred due to limitations of the biology, intrinsic neuronal noise, and imperfect measurements. We show that error, and the associated uncertainty surrounding a learned solution, can be controlled in large part by trading off synchronization strength among multiple redundant neural systems against the noise amplitude. The impact of the coupling between such redundant systems is quantified by the spectrum of the network Laplacian, and we discuss the role of network topology in synchronization and in reducing the effect of noise. A range of situations in which the mechanisms we model arise in brain science are discussed, and we draw attention to experimental evidence suggesting that cortical circuits capable of implementing the computations of interest here can be found on several scales. Finally, simulations comparing theoretical bounds to the relevant empirical quantities show that the theoretical estimates we derive can be tight.

We present a Cortical Neural Pool (CNP) architecture featuring a high-speed, resource-efficient CORDIC-based Hodgkin Huxley (RCHH) neuron model. Unlike shared CORDIC-based DNN approaches, the proposed neuron leverages modular and performance-optimised CORDIC stages with a latency-area trade-off. The FPGA implementation of the RCHH neuron shows 24.5% LUT reduction and 35.2% improved speed, compared to SoTA designs, with 70% better normalised root mean square error (NRMSE). Furthermore, the CNP exhibits 2.85x higher throughput (12.69 GOPS) compared to a functionally equivalent CORDIC-based DNN engine, with only a 0.35% accuracy drop compared to the DNN counterpart on the MNIST dataset. The overall results indicate that the design shows biologically accurate, low-resource spiking neural network implementations for resource-constrained edge AI applications.

Spiking neural networks offer low energy consumption due to their event-driven nature. Beyond binary spike outputs, their intrinsic floating-point dynamics merit greater attention. Neuronal threshold levels and reset modes critically determine spike count and timing. Hard reset cause information loss, while soft reset apply uniform treatment to neurons. To address these issues, we design an adaptive reset neuron that establishes relationships between inputs, outputs, and reset, while integrating a simple yet effective threshold adjustment strategy. Experimental results demonstrate that our method achieves excellent performance while maintaining lower energy consumption. In particular, it attains state-of-the-art accuracy on Tiny-ImageNet and CIFAR10-DVS. Codes are available at https://github.com/2ephyrus/AR-LIF.

Emergence of deep neural networks (DNNs) has raised enormous attention towards artificial neural networks (ANNs) once again. They have become the state-of-the-art models and have won different machine learning challenges. Although these networks are inspired by the brain, they lack biological plausibility, and they have structural differences compared to the brain. Spiking neural networks (SNNs) have been around for a long time, and they have been investigated to understand the dynamics of the brain. However, their application in real-world and complicated machine learning tasks were limited. Recently, they have shown great potential in solving such tasks. Due to their energy efficiency and temporal dynamics there are many promises in their future development. In this work, we reviewed the structures and performances of SNNs on image classification tasks. The comparisons illustrate that these networks show great capabilities for more complicated problems. Furthermore, the simple learning rules developed for SNNs, such as STDP and R-STDP, can be a potential alternative to replace the backpropagation algorithm used in DNNs.

Spiking Neural Networks (SNNs) are a promising research direction for building power-efficient information processing systems, especially for temporal tasks such as speech recognition. In SNNs, delays refer to the time needed for one spike to travel from one neuron to another. These delays matter because they influence the spike arrival times, and it is well-known that spiking neurons respond more strongly to coincident input spikes. More formally, it has been shown theoretically that plastic delays greatly increase the expressivity in SNNs. Yet, efficient algorithms to learn these delays have been lacking. Here, we propose a new discrete-time algorithm that addresses this issue in deep feedforward SNNs using backpropagation, in an offline manner. To simulate delays between consecutive layers, we use 1D convolutions across time. The kernels contain only a few non-zero weights - one per synapse - whose positions correspond to the delays. These positions are learned together with the weights using the recently proposed Dilated Convolution with Learnable Spacings (DCLS). We evaluated our method on three datasets: the Spiking Heidelberg Dataset (SHD), the Spiking Speech Commands (SSC) and its non-spiking version Google Speech Commands v0.02 (GSC) benchmarks, which require detecting temporal patterns. We used feedforward SNNs with two or three hidden fully connected layers, and vanilla leaky integrate-and-fire neurons. We showed that fixed random delays help and that learning them helps even more. Furthermore, our method outperformed the state-of-the-art in the three datasets without using recurrent connections and with substantially fewer parameters. Our work demonstrates the potential of delay learning in developing accurate and precise models for temporal data processing. Our code is based on PyTorch / SpikingJelly and available at: https://github.com/Thvnvtos/SNN-delays

Finding a code to unravel the population of neural responses that leads to a distinct animal behavior has been a long-standing question in the field of neuroscience. With the recent advances in machine learning, it is shown that the hierarchically Deep Neural Networks (DNNs) perform optimally in decoding unique features out of complex datasets. In this study, we utilize the power of a DNN to explore the computational principles in the mammalian brain by exploiting the Neuropixel data from Allen Brain Institute. We decode the neural responses from mouse visual cortex to predict the presented stimuli to the animal for natural (bear, trees, cheetah, etc.) and artificial (drifted gratings, orientated bars, etc.) classes. Our results indicate that neurons in mouse visual cortex encode the features of natural and artificial objects in a distinct manner, and such neural code is consistent across animals. We investigate this by applying transfer learning to train a DNN on the neural responses of a single animal and test its generalized performance across multiple animals. Within a single animal, DNN is able to decode the neural responses with as much as 100% classification accuracy. Across animals, this accuracy is reduced to 91%. This study demonstrates the potential of utilizing the DNN models as a computational framework to understand the neural coding principles in the mammalian brain.

We propose a novel nonlinear bidirectionally coupled heterogeneous chain network whose dynamics evolve in discrete time. The backbone of the model is a pair of popular map-based neuron models, the Chialvo and the Rulkov maps. This model is assumed to proximate the intricate dynamical properties of neurons in the widely complex nervous system. The model is first realized via various nonlinear analysis techniques: fixed point analysis, phase portraits, Jacobian matrix, and bifurcation diagrams. We observe the coexistence of chaotic and period-4 attractors. Various codimension-1 and -2 patterns for example saddle-node, period-doubling, Neimark-Sacker, double Neimark-Sacker, flip- and fold-Neimark Sacker, and 1:1 and 1:2 resonance are also explored. Furthermore, the study employs two synchronization measures to quantify how the oscillators in the network behave in tandem with each other over a long number of iterations. Finally, a time series analysis of the model is performed to investigate its complexity in terms of sample entropy.

The static synaptic connectivity of neuronal circuits stands in direct contrast to the dynamics of their function. As in changing community interactions, different neurons can participate actively in various combinations to effect behaviors at different times. We introduce an unsupervised approach to learn the dynamic affinities between neurons in live, behaving animals, and to reveal which communities form among neurons at different times. The inference occurs in two major steps. First, pairwise non-linear affinities between neuronal traces from brain-wide calcium activity are organized by non-negative tensor factorization (NTF). Each factor specifies which groups of neurons are most likely interacting for an inferred interval in time, and for which animals. Finally, a generative model that allows for weighted community detection is applied to the functional motifs produced by NTF to reveal a dynamic functional connectome. Since time codes the different experimental variables (e.g., application of chemical stimuli), this provides an atlas of neural motifs active during separate stages of an experiment (e.g., stimulus application or spontaneous behaviors). Results from our analysis are experimentally validated, confirming that our method is able to robustly predict causal interactions between neurons to generate behavior. Code is available at https://github.com/dyballa/dynamic-connectomes.

There is renewed interest in modeling and understanding the nervous system of the nematode Caenorhabditis elegans (C. elegans), as this small model system provides a path to bridge the gap between nervous system structure (connectivity) and function (physiology). However, existing physiology datasets, whether involving passive recording or stimulation, are in distinct formats, and connectome datasets require preprocessing before analysis can commence. Here we compile and homogenize datasets of neural activity and connectivity. Our neural activity dataset is derived from 11 C. elegans neuroimaging experiments, while our connectivity dataset is compiled from 9 connectome annotations based on 3 primary electron microscopy studies and 1 signal propagation study. Physiology datasets, collected under varying protocols, measure calcium fluorescence in labeled subsets of the worm's 300 neurons. Our preprocessing pipeline standardizes these datasets by consistently ordering labeled neurons and resampling traces to a common sampling rate, yielding recordings from approximately 900 worms and 250 uniquely labeled neurons. The connectome datasets, collected from electron microscopy reconstructions, represent the entire nervous system as a graph of connections. Our collection is accessible on HuggingFace, facilitating analysis of the structure-function relationship in biology using modern neural network architectures and enabling cross-lab and cross-animal comparisons.

Spiking neural networks (SNNs) possess energy-efficient potential due to event-based computation. However, supervised training of SNNs remains a challenge as spike activities are non-differentiable. Previous SNNs training methods can be generally categorized into two basic classes, i.e., backpropagation-like training methods and plasticity-based learning methods. The former methods are dependent on energy-inefficient real-valued computation and non-local transmission, as also required in artificial neural networks (ANNs), whereas the latter are either considered to be biologically implausible or exhibit poor performance. Hence, biologically plausible (bio-plausible) high-performance supervised learning (SL) methods for SNNs remain deficient. In this paper, we proposed a novel bio-plausible SNN model for SL based on the symmetric spike-timing dependent plasticity (sym-STDP) rule found in neuroscience. By combining the sym-STDP rule with bio-plausible synaptic scaling and intrinsic plasticity of the dynamic threshold, our SNN model implemented SL well and achieved good performance in the benchmark recognition task (MNIST dataset). To reveal the underlying mechanism of our SL model, we visualized both layer-based activities and synaptic weights using the t-distributed stochastic neighbor embedding (t-SNE) method after training and found that they were well clustered, thereby demonstrating excellent classification ability. Furthermore, to verify the robustness of our model, we trained it on another more realistic dataset (Fashion-MNIST), which also showed good performance. As the learning rules were bio-plausible and based purely on local spike events, our model could be easily applied to neuromorphic hardware for online training and may be helpful for understanding SL information processing at the synaptic level in biological neural systems.

Few algorithms for supervised training of spiking neural networks exist that can deal with patterns of multiple spikes, and their computational properties are largely unexplored. We demonstrate in a set of simulations that the ReSuMe learning algorithm can be successfully applied to layered neural networks. Input and output patterns are encoded as spike trains of multiple precisely timed spikes, and the network learns to transform the input trains into target output trains. This is done by combining the ReSuMe learning algorithm with multiplicative scaling of the connections of downstream neurons. We show in particular that layered networks with one hidden layer can learn the basic logical operations, including Exclusive-Or, while networks without hidden layer cannot, mirroring an analogous result for layered networks of rate neurons. While supervised learning in spiking neural networks is not yet fit for technical purposes, exploring computational properties of spiking neural networks advances our understanding of how computations can be done with spike trains.

Spiking neural networks (SNNs) are brain-inspired machine learning algorithms with merits such as biological plausibility and unsupervised learning capability. Previous works have shown that converting Artificial Neural Networks (ANNs) into SNNs is a practical and efficient approach for implementing an SNN. However, the basic principle and theoretical groundwork are lacking for training a non-accuracy-loss SNN. This paper establishes a precise mathematical mapping between the biological parameters of the Linear Leaky-Integrate-and-Fire model (LIF)/SNNs and the parameters of ReLU-AN/Deep Neural Networks (DNNs). Such mapping relationship is analytically proven under certain conditions and demonstrated by simulation and real data experiments. It can serve as the theoretical basis for the potential combination of the respective merits of the two categories of neural networks.

Spiking Neural Networks (SNNs) have been widely praised for their high energy efficiency and immense potential. However, comprehensive research that critically contrasts and correlates SNNs with quantized Artificial Neural Networks (ANNs) remains scant, often leading to skewed comparisons lacking fairness towards ANNs. This paper introduces a unified perspective, illustrating that the time steps in SNNs and quantized bit-widths of activation values present analogous representations. Building on this, we present a more pragmatic and rational approach to estimating the energy consumption of SNNs. Diverging from the conventional Synaptic Operations (SynOps), we champion the "Bit Budget" concept. This notion permits an intricate discourse on strategically allocating computational and storage resources between weights, activation values, and temporal steps under stringent hardware constraints. Guided by the Bit Budget paradigm, we discern that pivoting efforts towards spike patterns and weight quantization, rather than temporal attributes, elicits profound implications for model performance. Utilizing the Bit Budget for holistic design consideration of SNNs elevates model performance across diverse data types, encompassing static imagery and neuromorphic datasets. Our revelations bridge the theoretical chasm between SNNs and quantized ANNs and illuminate a pragmatic trajectory for future endeavors in energy-efficient neural computations.

A neuroscience method to understanding the brain is to find and study the preferred stimuli that highly activate an individual cell or groups of cells. Recent advances in machine learning enable a family of methods to synthesize preferred stimuli that cause a neuron in an artificial or biological brain to fire strongly. Those methods are known as Activation Maximization (AM) or Feature Visualization via Optimization. In this chapter, we (1) review existing AM techniques in the literature; (2) discuss a probabilistic interpretation for AM; and (3) review the applications of AM in debugging and explaining networks.

We put forward the dynamical study of a novel higher-order small network of Chialvo neurons arranged in a ring-star topology, with the neurons interacting via linear diffusive couplings. This model is perceived to imitate the nonlinear dynamical properties exhibited by a realistic nervous system where the neurons transfer information through higher-order multi-body interactions. We first analyze our model using the tools from nonlinear dynamics literature: fixed point analysis, Jacobian matrix, and bifurcation patterns. We observe the coexistence of chaotic attractors, and also an intriguing route to chaos starting from a fixed point, to period-doubling, to cyclic quasiperiodic closed invariant curves, to ultimately chaos. We numerically observe the existence of codimension-1 bifurcation patterns: saddle-node, period-doubling, and Neimark Sacker. We also qualitatively study the typical phase portraits of the system and numerically quantify chaos and complexity using the 0-1 test and sample entropy measure respectively. Finally, we study the collective behavior of the neurons in terms of two synchronization measures: the cross-correlation coefficient, and the Kuramoto order parameter.

Spiking Neural Networks (SNNs) are well known as a promising energy-efficient alternative to conventional artificial neural networks. Subject to the preconceived impression that SNNs are sparse firing, the analysis and optimization of inherent redundancy in SNNs have been largely overlooked, thus the potential advantages of spike-based neuromorphic computing in accuracy and energy efficiency are interfered. In this work, we pose and focus on three key questions regarding the inherent redundancy in SNNs. We argue that the redundancy is induced by the spatio-temporal invariance of SNNs, which enhances the efficiency of parameter utilization but also invites lots of noise spikes. Further, we analyze the effect of spatio-temporal invariance on the spatio-temporal dynamics and spike firing of SNNs. Then, motivated by these analyses, we propose an Advance Spatial Attention (ASA) module to harness SNNs' redundancy, which can adaptively optimize their membrane potential distribution by a pair of individual spatial attention sub-modules. In this way, noise spike features are accurately regulated. Experimental results demonstrate that the proposed method can significantly drop the spike firing with better performance than state-of-the-art SNN baselines. Our code is available in https://github.com/BICLab/ASA-SNN.

Parkinsons disease (PD) alters cortical neural dynamics, yet reliable non-invasive electrophysiological biomarkers remain elusive. This study examined whether interpretable EEG features capturing complementary aspects of neural dynamics can discriminate Parkinsonian neural states. A comprehensive set of interpretable features was extracted and grouped into Standard descriptors (spectral power, phase synchronization, time-domain statistics) and Dynamical descriptors (aperiodic activity, cross-frequency coupling, scale-free dynamics, neuronal avalanche statistics, and instantaneous frequency measures). A multi-head attention transformer classifier was trained using strict LOSO validation. Group-level comparisons were performed to identify electrophysiological differences associated with disease and medication state. Standard feature sets achieved strongest performance in discriminating medication states (PDoff vs PDon), whereas Dynamical performed competitively in contrasts between PD patients and healthy controls. Random feature ablation analyses indicated that Dynamical descriptors provide complementary information distributed across features while correlation analysis revealed low redundancy within both feature sets. Group-level comparisons revealed medication-sensitive reductions in delta power and voltage variance, modulation of neuronal avalanche statistics, persistent increases in theta phase synchronization in PD patients, and disease-related alterations in cross-frequency interactions. Traditional spectral and synchronization features primarily reflect medication-related neural modulation, whereas dynamical descriptors reveal broader alterations in cortical network organization associated with disease but also with medication. These findings support multivariate EEG representations as a promising framework for developing non-invasive biomarkers of PD.

A network-based approach is presented to investigate the cerebrovascular flow patterns during atrial fibrillation (AF) with respect to normal sinus rhythm (NSR). AF, the most common cardiac arrhythmia with faster and irregular beating, has been recently and independently associated with the increased risk of dementia. However, the underlying hemodynamic mechanisms relating the two pathologies remain mainly undetermined so far; thus the contribution of modeling and refined statistical tools is valuable. Pressure and flow rate temporal series in NSR and AF are here evaluated along representative cerebral sites (from carotid arteries to capillary brain circulation), exploiting reliable artificially built signals recently obtained from an in silico approach. The complex network analysis evidences, in a synthetic and original way, a dramatic signal variation towards the distal/capillary cerebral regions during AF, which has no counterpart in NSR conditions. At the large artery level, networks obtained from both AF and NSR hemodynamic signals exhibit elongated and chained features, which are typical of pseudo-periodic series. These aspects are almost completely lost towards the microcirculation during AF, where the networks are topologically more circular and present random-like characteristics. As a consequence, all the physiological phenomena at microcerebral level ruled by periodicity - such as regular perfusion, mean pressure per beat, and average nutrient supply at cellular level - can be strongly compromised, since the AF hemodynamic signals assume irregular behaviour and random-like features. Through a powerful approach which is complementary to the classical statistical tools, the present findings further strengthen the potential link between AF hemodynamic and cognitive decline.

It is desirable for statistical models to detect signals of interest independently of their position. If the data is generated by some smooth process, this additional structure should be taken into account. We introduce a new class of neural networks that are shift invariant and preserve smoothness of the data: functional neural networks (FNNs). For this, we use methods from functional data analysis (FDA) to extend multi-layer perceptrons and convolutional neural networks to functional data. We propose different model architectures, show that the models outperform a benchmark model from FDA in terms of accuracy and successfully use FNNs to classify electroencephalography (EEG) data.

Transformer-based language models excel at both recall (retrieving memorized facts) and reasoning (performing multi-step inference), but whether these abilities rely on distinct internal mechanisms remains unclear. Distinguishing recall from reasoning is crucial for predicting model generalization, designing targeted evaluations, and building safer interventions that affect one ability without disrupting the other.We approach this question through mechanistic interpretability, using controlled datasets of synthetic linguistic puzzles to probe transformer models at the layer, head, and neuron level. Our pipeline combines activation patching and structured ablations to causally measure component contributions to each task type. Across two model families (Qwen and LLaMA), we find that interventions on distinct layers and attention heads lead to selective impairments: disabling identified "recall circuits" reduces fact-retrieval accuracy by up to 15\% while leaving reasoning intact, whereas disabling "reasoning circuits" reduces multi-step inference by a comparable margin. At the neuron level, we observe task-specific firing patterns, though these effects are less robust, consistent with neuronal polysemanticity.Our results provide the first causal evidence that recall and reasoning rely on separable but interacting circuits in transformer models. These findings advance mechanistic interpretability by linking circuit-level structure to functional specialization and demonstrate how controlled datasets and causal interventions can yield mechanistic insights into model cognition, informing safer deployment of large language models.

Spiking Neural Networks (SNN) are a class of bio-inspired neural networks that promise to bring low-power and low-latency inference to edge devices through asynchronous and sparse processing. However, being temporal models, SNNs depend heavily on expressive states to generate predictions on par with classical artificial neural networks (ANNs). These states converge only after long transient periods, and quickly decay without input data, leading to higher latency, power consumption, and lower accuracy. This work addresses this issue by initializing the state with an auxiliary ANN running at a low rate. The SNN then uses the state to generate predictions with high temporal resolution until the next initialization phase. Our hybrid ANN-SNN model thus combines the best of both worlds: It does not suffer from long state transients and state decay thanks to the ANN, and can generate predictions with high temporal resolution, low latency, and low power thanks to the SNN. We show for the task of event-based 2D and 3D human pose estimation that our method consumes 88% less power with only a 4% decrease in performance compared to its fully ANN counterparts when run at the same inference rate. Moreover, when compared to SNNs, our method achieves a 74% lower error. This research thus provides a new understanding of how ANNs and SNNs can be used to maximize their respective benefits.

Mapping behavioral actions to neural activity is a fundamental goal of neuroscience. As our ability to record large neural and behavioral data increases, there is growing interest in modeling neural dynamics during adaptive behaviors to probe neural representations. In particular, neural latent embeddings can reveal underlying correlates of behavior, yet, we lack non-linear techniques that can explicitly and flexibly leverage joint behavior and neural data. Here, we fill this gap with a novel method, CEBRA, that jointly uses behavioral and neural data in a hypothesis- or discovery-driven manner to produce consistent, high-performance latent spaces. We validate its accuracy and demonstrate our tool's utility for both calcium and electrophysiology datasets, across sensory and motor tasks, and in simple or complex behaviors across species. It allows for single and multi-session datasets to be leveraged for hypothesis testing or can be used label-free. Lastly, we show that CEBRA can be used for the mapping of space, uncovering complex kinematic features, and rapid, high-accuracy decoding of natural movies from visual cortex.

Interpreting individual neurons or directions in activations space is an important component of mechanistic interpretability. As such, many algorithms have been proposed to automatically produce neuron explanations, but it is often not clear how reliable these explanations are, or which methods produce the best explanations. This can be measured via crowd-sourced evaluations, but they can often be noisy and expensive, leading to unreliable results. In this paper, we carefully analyze the evaluation pipeline and develop a cost-effective and highly accurate crowdsourced evaluation strategy. In contrast to previous human studies that only rate whether the explanation matches the most highly activating inputs, we estimate whether the explanation describes neuron activations across all inputs. To estimate this effectively, we introduce a novel application of importance sampling to determine which inputs are the most valuable to show to raters, leading to around 30x cost reduction compared to uniform sampling. We also analyze the label noise present in crowd-sourced evaluations and propose a Bayesian method to aggregate multiple ratings leading to a further ~5x reduction in number of ratings required for the same accuracy. Finally, we use these methods to conduct a large-scale study comparing the quality of neuron explanations produced by the most popular methods for two different vision models.

We introduce MABe22, a large-scale, multi-agent video and trajectory benchmark to assess the quality of learned behavior representations. This dataset is collected from a variety of biology experiments, and includes triplets of interacting mice (4.7 million frames video+pose tracking data, 10 million frames pose only), symbiotic beetle-ant interactions (10 million frames video data), and groups of interacting flies (4.4 million frames of pose tracking data). Accompanying these data, we introduce a panel of real-life downstream analysis tasks to assess the quality of learned representations by evaluating how well they preserve information about the experimental conditions (e.g. strain, time of day, optogenetic stimulation) and animal behavior. We test multiple state-of-the-art self-supervised video and trajectory representation learning methods to demonstrate the use of our benchmark, revealing that methods developed using human action datasets do not fully translate to animal datasets. We hope that our benchmark and dataset encourage a broader exploration of behavior representation learning methods across species and settings.

We propose a novel algorithm called Backpropagation Neural Tree (BNeuralT), which is a stochastic computational dendritic tree. BNeuralT takes random repeated inputs through its leaves and imposes dendritic nonlinearities through its internal connections like a biological dendritic tree would do. Considering the dendritic-tree like plausible biological properties, BNeuralT is a single neuron neural tree model with its internal sub-trees resembling dendritic nonlinearities. BNeuralT algorithm produces an ad hoc neural tree which is trained using a stochastic gradient descent optimizer like gradient descent (GD), momentum GD, Nesterov accelerated GD, Adagrad, RMSprop, or Adam. BNeuralT training has two phases, each computed in a depth-first search manner: the forward pass computes neural tree's output in a post-order traversal, while the error backpropagation during the backward pass is performed recursively in a pre-order traversal. A BNeuralT model can be considered a minimal subset of a neural network (NN), meaning it is a "thinned" NN whose complexity is lower than an ordinary NN. Our algorithm produces high-performing and parsimonious models balancing the complexity with descriptive ability on a wide variety of machine learning problems: classification, regression, and pattern recognition.

Recurrent neural networks (RNNs) provide a powerful approach in neuroscience to infer latent dynamics in neural populations and to generate hypotheses about the neural computations underlying behavior. However, past work has focused on relatively simple, input-driven, and largely deterministic behaviors - little is known about the mechanisms that would allow RNNs to generate the richer, spontaneous, and potentially stochastic behaviors observed in natural settings. Modeling with Hidden Markov Models (HMMs) has revealed a segmentation of natural behaviors into discrete latent states with stochastic transitions between them, a type of dynamics that may appear at odds with the continuous state spaces implemented by RNNs. Here we first show that RNNs can replicate HMM emission statistics and then reverse-engineer the trained networks to uncover the mechanisms they implement. In the absence of inputs, the activity of trained RNNs collapses towards a single fixed point. When driven by stochastic input, trajectories instead exhibit noise-sustained dynamics along closed orbits. Rotation along these orbits modulates the emission probabilities and is governed by transitions between regions of slow, noise-driven dynamics connected by fast, deterministic transitions. The trained RNNs develop highly structured connectivity, with a small set of "kick neurons" initiating transitions between these regions. This mechanism emerges during training as the network shifts into a regime of stochastic resonance, enabling it to perform probabilistic computations. Analyses across multiple HMM architectures - fully connected, cyclic, and linear-chain - reveal that this solution generalizes through the modular reuse of the same dynamical motif, suggesting a compositional principle by which RNNs can emulate complex discrete latent dynamics.

Artificial neural networks for motor control usually adopt generic architectures like fully connected MLPs. While general, these tabula rasa architectures rely on large amounts of experience to learn, are not easily transferable to new bodies, and have internal dynamics that are difficult to interpret. In nature, animals are born with highly structured connectivity in their nervous systems shaped by evolution; this innate circuitry acts synergistically with learning mechanisms to provide inductive biases that enable most animals to function well soon after birth and learn efficiently. Convolutional networks inspired by visual circuitry have encoded useful biases for vision. However, it is unknown the extent to which ANN architectures inspired by neural circuitry can yield useful biases for other AI domains. In this work, we ask what advantages biologically inspired ANN architecture can provide in the domain of motor control. Specifically, we translate C. elegans locomotion circuits into an ANN model controlling a simulated Swimmer agent. On a locomotion task, our architecture achieves good initial performance and asymptotic performance comparable with MLPs, while dramatically improving data efficiency and requiring orders of magnitude fewer parameters. Our architecture is interpretable and transfers to new body designs. An ablation analysis shows that constrained excitation/inhibition is crucial for learning, while weight initialization contributes to good initial performance. Our work demonstrates several advantages of biologically inspired ANN architecture and encourages future work in more complex embodied control.

Brain-computer interfaces (BCIs), transform neural signals in the brain into in-structions to control external devices. However, obtaining sufficient training data is difficult as well as limited. With the advent of advanced machine learning methods, the capability of brain-computer interfaces has been enhanced like never before, however, these methods require a large amount of data for training and thus require data augmentation of the limited data available. Here, we use spiking neural networks (SNN) as data generators. It is touted as the next-generation neu-ral network and is considered as one of the algorithms oriented to general artifi-cial intelligence because it borrows the neural information processing from bio-logical neurons. We use the SNN to generate neural spike information that is bio-interpretable and conforms to the intrinsic patterns in the original neural data. Ex-periments show that the model can directly synthesize new spike trains, which in turn improves the generalization ability of the BCI decoder. Both the input and output of the spiking neural model are spike information, which is a brain-inspired intelligence approach that can be better integrated with BCI in the future.

Biological cortical neurons are remarkably sophisticated computational devices, temporally integrating their vast synaptic input over an intricate dendritic tree, subject to complex, nonlinearly interacting internal biological processes. A recent study proposed to characterize this complexity by fitting accurate surrogate models to replicate the input-output relationship of a detailed biophysical cortical pyramidal neuron model and discovered it needed temporal convolutional networks (TCN) with millions of parameters. Requiring these many parameters, however, could stem from a misalignment between the inductive biases of the TCN and cortical neuron's computations. In light of this, and to explore the computational implications of leaky memory units and nonlinear dendritic processing, we introduce the Expressive Leaky Memory (ELM) neuron model, a biologically inspired phenomenological model of a cortical neuron. Remarkably, by exploiting such slowly decaying memory-like hidden states and two-layered nonlinear integration of synaptic input, our ELM neuron can accurately match the aforementioned input-output relationship with under ten thousand trainable parameters. To further assess the computational ramifications of our neuron design, we evaluate it on various tasks with demanding temporal structures, including the Long Range Arena (LRA) datasets, as well as a novel neuromorphic dataset based on the Spiking Heidelberg Digits dataset (SHD-Adding). Leveraging a larger number of memory units with sufficiently long timescales, and correspondingly sophisticated synaptic integration, the ELM neuron displays substantial long-range processing capabilities, reliably outperforming the classic Transformer or Chrono-LSTM architectures on LRA, and even solving the Pathfinder-X task with over 70% accuracy (16k context length).

The deep neural nets of modern artificial intelligence (AI) have not achieved defining features of biological intelligence, including abstraction, causal learning, and energy-efficiency. While scaling to larger models has delivered performance improvements for current applications, more brain-like capacities may demand new theories, models, and methods for designing artificial learning systems. Here, we argue that this opportunity to reassess insights from the brain should stimulate cooperation between AI research and theory-driven computational neuroscience (CN). To motivate a brain basis of neural computation, we present a dynamical view of intelligence from which we elaborate concepts of sparsity in network structure, temporal dynamics, and interactive learning. In particular, we suggest that temporal dynamics, as expressed through neural synchrony, nested oscillations, and flexible sequences, provide a rich computational layer for reading and updating hierarchical models distributed in long-term memory networks. Moreover, embracing agent-centered paradigms in AI and CN will accelerate our understanding of the complex dynamics and behaviors that build useful world models. A convergence of AI/CN theories and objectives will reveal dynamical principles of intelligence for brains and engineered learning systems. This article was inspired by our symposium on dynamical neuroscience and machine learning at the 6th Annual US/NIH BRAIN Initiative Investigators Meeting.

A basic question within the emerging field of mechanistic interpretability is the degree to which neural networks learn the same underlying mechanisms. In other words, are neural mechanisms universal across different models? In this work, we study the universality of individual neurons across GPT2 models trained from different initial random seeds, motivated by the hypothesis that universal neurons are likely to be interpretable. In particular, we compute pairwise correlations of neuron activations over 100 million tokens for every neuron pair across five different seeds and find that 1-5\% of neurons are universal, that is, pairs of neurons which consistently activate on the same inputs. We then study these universal neurons in detail, finding that they usually have clear interpretations and taxonomize them into a small number of neuron families. We conclude by studying patterns in neuron weights to establish several universal functional roles of neurons in simple circuits: deactivating attention heads, changing the entropy of the next token distribution, and predicting the next token to (not) be within a particular set.

We analyze a family of large language models in such a lightweight manner that can be done on a single GPU. Specifically, we focus on the OPT family of models ranging from 125m to 66b parameters and rely only on whether an FFN neuron is activated or not. First, we find that the early part of the network is sparse and represents many discrete features. Here, many neurons (more than 70% in some layers of the 66b model) are "dead", i.e. they never activate on a large collection of diverse data. At the same time, many of the alive neurons are reserved for discrete features and act as token and n-gram detectors. Interestingly, their corresponding FFN updates not only promote next token candidates as could be expected, but also explicitly focus on removing the information about triggering them tokens, i.e., current input. To the best of our knowledge, this is the first example of mechanisms specialized at removing (rather than adding) information from the residual stream. With scale, models become more sparse in a sense that they have more dead neurons and token detectors. Finally, some neurons are positional: them being activated or not depends largely (or solely) on position and less so (or not at all) on textual data. We find that smaller models have sets of neurons acting as position range indicators while larger models operate in a less explicit manner.

Recurrent neural networks (RNNs) are widely used throughout neuroscience as models of local neural activity. Many properties of single RNNs are well characterized theoretically, but experimental neuroscience has moved in the direction of studying multiple interacting areas, and RNN theory needs to be likewise extended. We take a constructive approach towards this problem, leveraging tools from nonlinear control theory and machine learning to characterize when combinations of stable RNNs will themselves be stable. Importantly, we derive conditions which allow for massive feedback connections between interacting RNNs. We parameterize these conditions for easy optimization using gradient-based techniques, and show that stability-constrained "networks of networks" can perform well on challenging sequential-processing benchmark tasks. Altogether, our results provide a principled approach towards understanding distributed, modular function in the brain.

Animals ranging from rats to humans can demonstrate cognitive map capabilities. We evolved weights in a biologically plausible recurrent neural network (RNN) using an evolutionary algorithm to replicate the behavior and neural activity observed in rats during a spatial and working memory task in a triple T-maze. The rat was simulated in the Webots robot simulator and used vision, distance and accelerometer sensors to navigate a virtual maze. After evolving weights from sensory inputs to the RNN, within the RNN, and from the RNN to the robot's motors, the Webots agent successfully navigated the space to reach all four reward arms with minimal repeats before time-out. Our current findings suggest that it is the RNN dynamics that are key to performance, and that performance is not dependent on any one sensory type, which suggests that neurons in the RNN are performing mixed selectivity and conjunctive coding. Moreover, the RNN activity resembles spatial information and trajectory-dependent coding observed in the hippocampus. Collectively, the evolved RNN exhibits navigation skills, spatial memory, and working memory. Our method demonstrates how the dynamic activity in evolved RNNs can capture interesting and complex cognitive behavior and may be used to create RNN controllers for robotic applications.

Prior studies on the emergence in large models have primarily focused on how the functional capabilities of large language models (LLMs) scale with model size. Our research, however, transcends this traditional paradigm, aiming to deepen our understanding of the emergence within LLMs by placing a special emphasis not just on the model size but more significantly on the complex behavior of neuron interactions during the training process. By introducing the concepts of "self-organization" and "multifractal analysis," we explore how neuron interactions dynamically evolve during training, leading to "emergence," mirroring the phenomenon in natural systems where simple micro-level interactions give rise to complex macro-level behaviors. To quantitatively analyze the continuously evolving interactions among neurons in large models during training, we propose the Neuron-based Multifractal Analysis (NeuroMFA). Utilizing NeuroMFA, we conduct a comprehensive examination of the emergent behavior in LLMs through the lens of both model size and training process, paving new avenues for research into the emergence in large models.

Partially inspired by features of computation in visual cortex, deep neural networks compute hierarchical representations of their inputs. While these networks have been highly successful in machine learning, it remains unclear to what extent they can aid our understanding of cortical function. Several groups have developed metrics that provide a quantitative comparison between representations computed by networks and representations measured in cortex. At the same time, neuroscience is well into an unprecedented phase of large-scale data collection, as evidenced by projects such as the Allen Brain Observatory. Despite the magnitude of these efforts, in a given experiment only a fraction of units are recorded, limiting the information available about the cortical representation. Moreover, only a finite number of stimuli can be shown to an animal over the course of a realistic experiment. These limitations raise the question of how and whether metrics that compare representations of deep networks are meaningful on these datasets. Here, we empirically quantify the capabilities and limitations of these metrics due to limited image presentations and neuron samples. We find that the comparison procedure is robust to different choices of stimuli set and the level of subsampling that one might expect in a large-scale brain survey with thousands of neurons. Using these results, we compare the representations measured in the Allen Brain Observatory in response to natural image presentations to deep neural network. We show that the visual cortical areas are relatively high order representations (in that they map to deeper layers of convolutional neural networks). Furthermore, we see evidence of a broad, more parallel organization rather than a sequential hierarchy, with the primary area VISp(V1) being lower order relative to the other areas.

This paper studies the problem of training a two-layer ReLU network for binary classification using gradient flow with small initialization. We consider a training dataset with well-separated input vectors: Any pair of input data with the same label are positively correlated, and any pair with different labels are negatively correlated. Our analysis shows that, during the early phase of training, neurons in the first layer try to align with either the positive data or the negative data, depending on its corresponding weight on the second layer. A careful analysis of the neurons' directional dynamics allows us to provide an O(log n{mu}) upper bound on the time it takes for all neurons to achieve good alignment with the input data, where n is the number of data points and mu measures how well the data are separated. After the early alignment phase, the loss converges to zero at a O(1{t}) rate, and the weight matrix on the first layer is approximately low-rank. Numerical experiments on the MNIST dataset illustrate our theoretical findings.

Instance segmentation of neurons in volumetric light microscopy images of nervous systems enables groundbreaking research in neuroscience by facilitating joint functional and morphological analyses of neural circuits at cellular resolution. Yet said multi-neuron light microscopy data exhibits extremely challenging properties for the task of instance segmentation: Individual neurons have long-ranging, thin filamentous and widely branching morphologies, multiple neurons are tightly inter-weaved, and partial volume effects, uneven illumination and noise inherent to light microscopy severely impede local disentangling as well as long-range tracing of individual neurons. These properties reflect a current key challenge in machine learning research, namely to effectively capture long-range dependencies in the data. While respective methodological research is buzzing, to date methods are typically benchmarked on synthetic datasets. To address this gap, we release the FlyLight Instance Segmentation Benchmark (FISBe) dataset, the first publicly available multi-neuron light microscopy dataset with pixel-wise annotations. In addition, we define a set of instance segmentation metrics for benchmarking that we designed to be meaningful with regard to downstream analyses. Lastly, we provide three baselines to kick off a competition that we envision to both advance the field of machine learning regarding methodology for capturing long-range data dependencies, and facilitate scientific discovery in basic neuroscience.

Epilepsy is a common disease of the nervous system. Timely prediction of seizures and intervention treatment can significantly reduce the accidental injury of patients and protect the life and health of patients. This paper presents a neuromorphic Spiking Convolutional Transformer, named Spiking Conformer, to detect and predict epileptic seizure segments from scalped long-term electroencephalogram (EEG) recordings. We report evaluation results from the Spiking Conformer model using the Boston Children's Hospital-MIT (CHB-MIT) EEG dataset. By leveraging spike-based addition operations, the Spiking Conformer significantly reduces the classification computational cost compared to the non-spiking model. Additionally, we introduce an approximate spiking neuron layer to further reduce spike-triggered neuron updates by nearly 38% without sacrificing accuracy. Using raw EEG data as input, the proposed Spiking Conformer achieved an average sensitivity rate of 94.9% and a specificity rate of 99.3% for the seizure detection task, and 96.8%, 89.5% for the seizure prediction task, and needs >10x fewer operations compared to the non-spiking equivalent model.

Within the complex neuroarchitecture of the brain, astrocytes play crucial roles in development, structure, and metabolism. These cells regulate neural activity through tripartite synapses, directly impacting cognitive processes such as learning and memory. Despite the growing recognition of astrocytes' significance, traditional Spiking Neural Network (SNN) models remain predominantly neuron-centric, overlooking the profound influence of astrocytes on neural dynamics. Inspired by these biological insights, we have developed an Astrocyte-Modulated Spiking Unit (AM-SU), an innovative framework that integrates neuron-astrocyte interactions into the computational paradigm, demonstrating wide applicability across various hardware platforms. Our Astrocyte-Modulated Spiking Neural Network (AstroSNN) exhibits exceptional performance in tasks involving memory retention and natural language generation, particularly in handling long-term dependencies and complex linguistic structures. The design of AstroSNN not only enhances its biological authenticity but also introduces novel computational dynamics, enabling more effective processing of complex temporal dependencies. Furthermore, AstroSNN shows low latency, high throughput, and reduced memory usage in practical applications, making it highly suitable for resource-constrained environments. By successfully integrating astrocytic dynamics into intelligent neural networks, our work narrows the gap between biological plausibility and neural modeling, laying the groundwork for future biologically-inspired neural computing research that includes both neurons and astrocytes.

The informational synthesis of neural structures, processes, parameters and characteristics that allow a unified description and modeling as neural machines of natural and artificial neural systems is presented. The general informational parameters as the global quantitative measure of the neural systems computing potential as absolute and relative neural power were proposed. Neural information organizing and processing follows the way in which nature manages neural information by developing functions, functionalities and circuits related to different internal or peripheral components and also to the whole system through a non-deterministic memorization, fragmentation and aggregation of afferent and efferent information, deep neural information processing representing multiple alternations of fragmentation and aggregation stages. The relevant neural characteristics were integrated into a neural machine type model that incorporates unitary also peripheral or interface components as the central ones. The proposed approach allows overcoming the technical constraints in artificial computational implementations of neural information processes and also provides a more relevant description of natural ones.

The approximation capability of ANNs and their RNN instantiations, is strongly correlated with the number of parameters packed into these networks. However, the complexity barrier for human understanding, is arguably related to the number of neurons and synapses in the networks, and to the associated nonlinear transformations. In this paper we show that the use of biophysical synapses, as found in LTCs, have two main benefits. First, they allow to pack more parameters for a given number of neurons and synapses. Second, they allow to formulate the nonlinear-network transformation, as a linear system with state-dependent coefficients. Both increase interpretability, as for a given task, they allow to learn a system linear in its input features, that is smaller in size compared to the state of the art. We substantiate the above claims on various time-series prediction tasks, but we believe that our results are applicable to any feedforward or recurrent ANN.

Spiking Neural Networks (SNNs) offer energy-efficient, biologically plausible computation but suffer from non-differentiable spike generation, necessitating reliance on heuristic surrogate gradients. This paper introduces UltraLIF, a principled framework that replaces surrogate gradients with ultradiscretization, a mathematical formalism from tropical geometry providing continuous relaxations of discrete dynamics. The central insight is that the max-plus semiring underlying ultradiscretization naturally models neural threshold dynamics: the log-sum-exp function serves as a differentiable soft-maximum that converges to hard thresholding as a learnable temperature parameter eps to 0. Two neuron models are derived from distinct dynamical systems: UltraLIF from the LIF ordinary differential equation (temporal dynamics) and UltraDLIF from the diffusion equation modeling gap junction coupling across neuronal populations (spatial dynamics). Both yield fully differentiable SNNs trainable via standard backpropagation with no forward-backward mismatch. Theoretical analysis establishes pointwise convergence to classical LIF dynamics with quantitative error bounds and bounded non-vanishing gradients. Experiments on six benchmarks spanning static images, neuromorphic vision, and audio demonstrate improvements over surrogate gradient baselines, with gains most pronounced in single-timestep (T{=}1) settings on neuromorphic and temporal datasets. An optional sparsity penalty enables significant energy reduction while maintaining competitive accuracy.

Learning synaptic weights of spiking neural network (SNN) models that can reproduce target spike trains from provided neural firing data is a central problem in computational neuroscience and spike-based computing. The discovery of the optimal weight values can be posed as a supervised learning task wherein the weights of the model network are chosen to maximize the similarity between the target spike trains and the model outputs. It is still largely unknown whether optimizing spike train similarity of highly recurrent SNNs produces weight matrices similar to those of the ground truth model. To this end, we propose flexible heuristic supervised learning rules, termed Pre-Synaptic Pool Modification (PSPM), that rely on stochastic weight updates in order to produce spikes within a short window of the desired times and eliminate spikes outside of this window. PSPM improves spike train similarity for all-to-all SNNs and makes no assumption about the post-synaptic potential of the neurons or the structure of the network since no gradients are required. We test whether optimizing for spike train similarity entails the discovery of accurate weights and explore the relative contributions of local and homeostatic weight updates. Although PSPM improves similarity between spike trains, the learned weights often differ from the weights of the ground truth model, implying that connectome inference from spike data may require additional constraints on connectivity statistics. We also find that spike train similarity is sensitive to local updates, but other measures of network activity such as avalanche distributions, can be learned through synaptic homeostasis.

Neuromorphic systems, inspired by the complexity and functionality of the human brain, have gained interest in academic and industrial attention due to their unparalleled potential across a wide range of applications. While their capabilities herald innovation, it is imperative to underscore that these computational paradigms, analogous to their traditional counterparts, are not impervious to security threats. Although the exploration of neuromorphic methodologies for image and video processing has been rigorously pursued, the realm of neuromorphic audio processing remains in its early stages. Our results highlight the robustness and precision of our FPGA-based neuromorphic system. Specifically, our system showcases a commendable balance between desired signal and background noise, efficient spike rate encoding, and unparalleled resilience against adversarial attacks such as FGSM and PGD. A standout feature of our framework is its detection rate of 94%, which, when compared to other methodologies, underscores its greater capability in identifying and mitigating threats within 5.39 dB, a commendable SNR ratio. Furthermore, neuromorphic computing and hardware security serve many sensor domains in mission-critical and privacy-preserving applications.

Spiking neural networks (SNNs) are brain-inspired energy-efficient models that encode information in spatiotemporal dynamics. Recently, deep SNNs trained directly have shown great success in achieving high performance on classification tasks with very few time steps. However, how to design a directly-trained SNN for the regression task of object detection still remains a challenging problem. To address this problem, we propose EMS-YOLO, a novel directly-trained SNN framework for object detection, which is the first trial to train a deep SNN with surrogate gradients for object detection rather than ANN-SNN conversion strategies. Specifically, we design a full-spike residual block, EMS-ResNet, which can effectively extend the depth of the directly-trained SNN with low power consumption. Furthermore, we theoretically analyze and prove the EMS-ResNet could avoid gradient vanishing or exploding. The results demonstrate that our approach outperforms the state-of-the-art ANN-SNN conversion methods (at least 500 time steps) in extremely fewer time steps (only 4 time steps). It is shown that our model could achieve comparable performance to the ANN with the same architecture while consuming 5.83 times less energy on the frame-based COCO Dataset and the event-based Gen1 Dataset.

Recurrent Spiking Neural Networks (RSNNs) have emerged as a computationally efficient and brain-inspired learning model. The design of sparse RSNNs with fewer neurons and synapses helps reduce the computational complexity of RSNNs. Traditionally, sparse SNNs are obtained by first training a dense and complex SNN for a target task, and, then, pruning neurons with low activity (activity-based pruning) while maintaining task performance. In contrast, this paper presents a task-agnostic methodology for designing sparse RSNNs by pruning a large randomly initialized model. We introduce a novel Lyapunov Noise Pruning (LNP) algorithm that uses graph sparsification methods and utilizes Lyapunov exponents to design a stable sparse RSNN from a randomly initialized RSNN. We show that the LNP can leverage diversity in neuronal timescales to design a sparse Heterogeneous RSNN (HRSNN). Further, we show that the same sparse HRSNN model can be trained for different tasks, such as image classification and temporal prediction. We experimentally show that, in spite of being task-agnostic, LNP increases computational efficiency (fewer neurons and synapses) and prediction performance of RSNNs compared to traditional activity-based pruning of trained dense models.

This study explores the interference in temporal processing within a dual-task paradigm from an artificial intelligence (AI) perspective. In this context, the dual-task setup is implemented as a simplified version of the Overcooked environment with two variations, single task (T) and dual task (T+N). Both variations involve an embedded time production task, but the dual task (T+N) additionally involves a concurrent number comparison task. Two deep reinforcement learning (DRL) agents were separately trained for each of these tasks. These agents exhibited emergent behavior consistent with human timing research. Specifically, the dual task (T+N) agent exhibited significant overproduction of time relative to its single task (T) counterpart. This result was consistent across four target durations. Preliminary analysis of neural dynamics in the agents' LSTM layers did not reveal any clear evidence of a dedicated or intrinsic timer. Hence, further investigation is needed to better understand the underlying time-keeping mechanisms of the agents and to provide insights into the observed behavioral patterns. This study is a small step towards exploring parallels between emergent DRL behavior and behavior observed in biological systems in order to facilitate a better understanding of both.

Implanted devices providing real-time neural activity classification and control are increasingly used to treat neurological disorders, such as epilepsy and Parkinson's disease. Classification performance is critical to identifying brain states appropriate for the therapeutic action. However, advanced algorithms that have shown promise in offline studies, in particular deep learning (DL) methods, have not been deployed on resource-restrained neural implants. Here, we designed and optimized three embedded DL models of commonly adopted architectures and evaluated their inference performance in a case study of seizure detection. A deep neural network (DNN), a convolutional neural network (CNN), and a long short-term memory (LSTM) network were designed to classify ictal, preictal, and interictal phases from the CHB-MIT scalp EEG database. After iterative model compression and quantization, the algorithms were deployed on a general-purpose, off-the-shelf microcontroller. Inference sensitivity, false positive rate, execution time, memory size, and power consumption were quantified. For seizure event detection, the sensitivity and FPR (h-1) for the DNN, CNN, and LSTM models were 87.36%/0.169, 96.70%/0.102, and 97.61%/0.071, respectively. Predicting seizures for early warnings was also feasible. The implemented compression and quantization achieved a significant saving of power and memory with an accuracy degradation of less than 0.5%. Edge DL models achieved performance comparable to many prior implementations that had no time or computational resource limitations. Generic microcontrollers can provide the required memory and computational resources, while model designs can be migrated to ASICs for further optimization. The results suggest that edge DL inference is a feasible option for future neural implants to improve classification performance and therapeutic outcomes.

Modelling the complex spatiotemporal patterns of large-scale brain dynamics is crucial for neuroscience, but traditional methods fail to capture the rich structure in modalities such as magnetoencephalography (MEG). Recent advances in deep learning have enabled significant progress in other domains, such as language and vision, by using foundation models at scale. Here, we introduce MEG-GPT, a transformer based foundation model that uses time-attention and next time-point prediction. To facilitate this, we also introduce a novel data-driven tokeniser for continuous MEG data, which preserves the high temporal resolution of continuous MEG signals without lossy transformations. We trained MEG-GPT on tokenised brain region time-courses extracted from a large-scale MEG dataset (N=612, eyes-closed rest, Cam-CAN data), and show that the learnt model can generate data with realistic spatio-spectral properties, including transient events and population variability. Critically, it performs well in downstream decoding tasks, improving downstream supervised prediction task, showing improved zero-shot generalisation across sessions (improving accuracy from 0.54 to 0.59) and subjects (improving accuracy from 0.41 to 0.49) compared to a baseline methods. Furthermore, we show the model can be efficiently fine-tuned on a smaller labelled dataset to boost performance in cross-subject decoding scenarios. This work establishes a powerful foundation model for electrophysiological data, paving the way for applications in computational neuroscience and neural decoding.

The human brain is a complex spiking neural network (SNN) that learns multimodal signals in a zero-shot manner by generalizing existing knowledge. Remarkably, the brain achieves this with minimal power consumption, using event-based signals that propagate within its structure. However, mimicking the human brain in neuromorphic hardware presents both hardware and software challenges. Hardware limitations, such as the slowdown of Moore's law and the von Neumann bottleneck, hinder the efficiency of digital computers. On the software side, SNNs are known for their difficult training, especially when learning multimodal signals. To overcome these challenges, we propose a hardware-software co-design that combines a fixed and random liquid state machine (LSM) SNN encoder with trainable artificial neural network (ANN) projections. The LSM is physically implemented using analogue resistive memory, leveraging the inherent stochasticity of resistive switching to generate random weights. This highly efficient and nanoscale in-memory computing approach effectively addresses the von Neumann bottleneck and the slowdown of Moore's law. The ANN projections are implemented digitally, allowing for easy optimization using contrastive loss, which helps to overcome the difficulties associated with SNN training. We experimentally implement this co-design on a 40nm 256Kb in-memory computing macro. We first demonstrate LSM-based event encoding through supervised classification and linear probing on the N-MNIST and N-TIDIGITS datasets.

We present the Brain Treebank, a large-scale dataset of electrophysiological neural responses, recorded from intracranial probes while 10 subjects watched one or more Hollywood movies. Subjects watched on average 2.6 Hollywood movies, for an average viewing time of 4.3 hours, and a total of 43 hours. The audio track for each movie was transcribed with manual corrections. Word onsets were manually annotated on spectrograms of the audio track for each movie. Each transcript was automatically parsed and manually corrected into the universal dependencies (UD) formalism, assigning a part of speech to every word and a dependency parse to every sentence. In total, subjects heard over 38,000 sentences (223,000 words), while they had on average 168 electrodes implanted. This is the largest dataset of intracranial recordings featuring grounded naturalistic language, one of the largest English UD treebanks in general, and one of only a few UD treebanks aligned to multimodal features. We hope that this dataset serves as a bridge between linguistic concepts, perception, and their neural representations. To that end, we present an analysis of which electrodes are sensitive to language features while also mapping out a rough time course of language processing across these electrodes. The Brain Treebank is available at https://BrainTreebank.dev/

Some neurons in deep networks specialize in recognizing highly specific perceptual, structural, or semantic features of inputs. In computer vision, techniques exist for identifying neurons that respond to individual concept categories like colors, textures, and object classes. But these techniques are limited in scope, labeling only a small subset of neurons and behaviors in any network. Is a richer characterization of neuron-level computation possible? We introduce a procedure (called MILAN, for mutual-information-guided linguistic annotation of neurons) that automatically labels neurons with open-ended, compositional, natural language descriptions. Given a neuron, MILAN generates a description by searching for a natural language string that maximizes pointwise mutual information with the image regions in which the neuron is active. MILAN produces fine-grained descriptions that capture categorical, relational, and logical structure in learned features. These descriptions obtain high agreement with human-generated feature descriptions across a diverse set of model architectures and tasks, and can aid in understanding and controlling learned models. We highlight three applications of natural language neuron descriptions. First, we use MILAN for analysis, characterizing the distribution and importance of neurons selective for attribute, category, and relational information in vision models. Second, we use MILAN for auditing, surfacing neurons sensitive to human faces in datasets designed to obscure them. Finally, we use MILAN for editing, improving robustness in an image classifier by deleting neurons sensitive to text features spuriously correlated with class labels.

Originally inspired by neurobiology, deep neural network models have become a powerful tool of machine learning and artificial intelligence, where they are used to approximate functions and dynamics by learning from examples. Here we give a brief introduction to neural network models and deep learning for biologists. We introduce feedforward and recurrent networks and explain the expressive power of this modeling framework and the backpropagation algorithm for setting the parameters. Finally, we consider how deep neural networks might help us understand the brain's computations.

Large Language Models (LLMs) are composed of neurons that exhibit various behaviors and roles, which become increasingly diversified as models scale. Recent studies have revealed that not all neurons are active across different datasets, and this sparsity correlates positively with the task-specific ability, leading to advancements in model pruning and training efficiency. Traditional fine-tuning methods engage all parameters of LLMs, which is computationally expensive and may not be necessary. In contrast, Parameter-Efficient Fine-Tuning (PEFT) approaches aim to minimize the number of trainable parameters, yet they still operate at a relatively macro scale (e.g., layer-level). We introduce Neuron-Level Fine-Tuning (NeFT), a novel approach that refines the granularity of parameter training down to the individual neuron, enabling more precise and computationally efficient model updates. The experimental results show that NeFT not only exceeded the performance of full-parameter fine-tuning and PEFT but also provided insights into the analysis of neurons.

Large language models (LLMs) exhibit strong multilingual abilities, yet the neural mechanisms behind language-specific processing remain unclear. We analyze language-specific neurons in Llama-3.1-8B, Mistral-Nemo-12B, and Aya-Expanse-8B & 32B across 21 typologically diverse languages, identifying neurons that control language behavior. Using the Language Activation Probability Entropy (LAPE) method, we show that these neurons cluster in deeper layers, with non-Latin scripts showing greater specialization. Related languages share overlapping neurons, reflecting internal representations of linguistic proximity. Through language arithmetics, i.e. systematic activation addition and multiplication, we steer models to deactivate unwanted languages and activate desired ones, outperforming simpler replacement approaches. These interventions effectively guide behavior across five multilingual tasks: language forcing, translation, QA, comprehension, and NLI. Manipulation is more successful for high-resource languages, while typological similarity improves effectiveness. We also demonstrate that cross-lingual neuron steering enhances downstream performance and reveal internal "fallback" mechanisms for language selection when neurons are progressively deactivated. Our code is made publicly available at https://github.com/d-gurgurov/Language-Neurons-Manipulation.

The liquid state machine (LSM) combines low training complexity and biological plausibility, which has made it an attractive machine learning framework for edge and neuromorphic computing paradigms. Originally proposed as a model of brain computation, the LSM tunes its internal weights without backpropagation of gradients, which results in lower performance compared to multi-layer neural networks. Recent findings in neuroscience suggest that astrocytes, a long-neglected non-neuronal brain cell, modulate synaptic plasticity and brain dynamics, tuning brain networks to the vicinity of the computationally optimal critical phase transition between order and chaos. Inspired by this disruptive understanding of how brain networks self-tune, we propose the neuron-astrocyte liquid state machine (NALSM) that addresses under-performance through self-organized near-critical dynamics. Similar to its biological counterpart, the astrocyte model integrates neuronal activity and provides global feedback to spike-timing-dependent plasticity (STDP), which self-organizes NALSM dynamics around a critical branching factor that is associated with the edge-of-chaos. We demonstrate that NALSM achieves state-of-the-art accuracy versus comparable LSM methods, without the need for data-specific hand-tuning. With a top accuracy of 97.61% on MNIST, 97.51% on N-MNIST, and 85.84% on Fashion-MNIST, NALSM achieved comparable performance to current fully-connected multi-layer spiking neural networks trained via backpropagation. Our findings suggest that the further development of brain-inspired machine learning methods has the potential to reach the performance of deep learning, with the added benefits of supporting robust and energy-efficient neuromorphic computing on the edge.

This study introduces BrainTransformers, an innovative Large Language Model (LLM) implemented using Spiking Neural Networks (SNN). Our key contributions include: (1) designing SNN-compatible Transformer components such as SNNMatmul, SNNSoftmax, and SNNSiLU; (2) implementing an SNN approximation of the SiLU activation function; and (3) developing a Synapsis module to simulate synaptic plasticity. Our 3-billion parameter model, BrainTransformers-3B-Chat, demonstrates competitive performance across various benchmarks, including MMLU (63.2), BBH (54.1), ARC-C (54.3), and GSM8K (76.3), while potentially offering improved energy efficiency and biological plausibility. The model employs a three-stage training approach, including SNN-specific neuronal synaptic plasticity training. This research opens new avenues for brain-like AI systems in natural language processing and neuromorphic computing. Future work will focus on hardware optimization, developing specialized SNN fine-tuning tools, and exploring practical applications in energy-efficient computing environments.

Motivated by the celebrated discrete-time model of nervous activity outlined by McCulloch and Pitts in 1943, we propose a novel continuous-time model, the McCulloch-Pitts network (MPN), for sequence learning in spiking neural networks. Our model has a local learning rule, such that the synaptic weight updates depend only on the information directly accessible by the synapse. By exploiting asymmetry in the connections between binary neurons, we show that MPN can be trained to robustly memorize multiple spatiotemporal patterns of binary vectors, generalizing the ability of the symmetric Hopfield network to memorize static spatial patterns. In addition, we demonstrate that the model can efficiently learn sequences of binary pictures as well as generative models for experimental neural spike-train data. Our learning rule is consistent with spike-timing-dependent plasticity (STDP), thus providing a theoretical ground for the systematic design of biologically inspired networks with large and robust long-range sequence storage capacity.

This paper presents a new methodology to alleviate the fundamental trade-off between accuracy and latency in spiking neural networks (SNNs). The approach involves decoding confidence information over time from the SNN outputs and using it to develop a decision-making agent that can dynamically determine when to terminate each inference. The proposed method, Dynamic Confidence, provides several significant benefits to SNNs. 1. It can effectively optimize latency dynamically at runtime, setting it apart from many existing low-latency SNN algorithms. Our experiments on CIFAR-10 and ImageNet datasets have demonstrated an average 40% speedup across eight different settings after applying Dynamic Confidence. 2. The decision-making agent in Dynamic Confidence is straightforward to construct and highly robust in parameter space, making it extremely easy to implement. 3. The proposed method enables visualizing the potential of any given SNN, which sets a target for current SNNs to approach. For instance, if an SNN can terminate at the most appropriate time point for each input sample, a ResNet-50 SNN can achieve an accuracy as high as 82.47% on ImageNet within just 4.71 time steps on average. Unlocking the potential of SNNs needs a highly-reliable decision-making agent to be constructed and fed with a high-quality estimation of ground truth. In this regard, Dynamic Confidence represents a meaningful step toward realizing the potential of SNNs.

We explore the effects of various plasticity functions on assemblies of neurons. To bridge the gap between experimental and computational theories we make use of a conceptual framework, the Assembly Calculus, which is a formal system for the description of brain function based on assemblies of neurons. The Assembly Calculus includes operations for projecting, associating, and merging assemblies of neurons. Our research is focused on simulating different plasticity functions with Assembly Calculus. Our main contribution is the modification and evaluation of the projection operation. We experiment with Oja's and Spike Time-Dependent Plasticity (STDP) rules and test the effect of various hyper-parameters.

Recent advances in artificial neural networks for machine learning, and language modeling in particular, have established a family of recurrent neural network (RNN) architectures that, unlike conventional RNNs with vector-form hidden states, use two-dimensional (2D) matrix-form hidden states. Such 2D-state RNNs, known as Fast Weight Programmers (FWPs), can be interpreted as a neural network whose synaptic weights (called fast weights) dynamically change over time as a function of input observations, and serve as short-term memory storage; corresponding synaptic weight modifications are controlled or programmed by another network (the programmer) whose parameters are trained (e.g., by gradient descent). In this Primer, we review the technical foundations of FWPs, their computational characteristics, and their connections to transformers and state space models. We also discuss connections between FWPs and models of synaptic plasticity in the brain, suggesting a convergence of natural and artificial intelligence.

While neural networks can be approximated by linear models as their width increases, certain properties of wide neural networks cannot be captured by linear models. In this work we show that recently proposed Neural Quadratic Models can exhibit the "catapult phase" [Lewkowycz et al. 2020] that arises when training such models with large learning rates. We then empirically show that the behaviour of neural quadratic models parallels that of neural networks in generalization, especially in the catapult phase regime. Our analysis further demonstrates that quadratic models can be an effective tool for analysis of neural networks.

This position and survey paper identifies the emerging convergence of neuroscience, artificial general intelligence (AGI), and neuromorphic computing toward a unified research paradigm. Using a framework grounded in brain physiology, we highlight how synaptic plasticity, sparse spike-based communication, and multimodal association provide design principles for next-generation AGI systems that potentially combine both human and machine intelligences. The review traces this evolution from early connectionist models to state-of-the-art large language models, demonstrating how key innovations like transformer attention, foundation-model pre-training, and multi-agent architectures mirror neurobiological processes like cortical mechanisms, working memory, and episodic consolidation. We then discuss emerging physical substrates capable of breaking the von Neumann bottleneck to achieve brain-scale efficiency in silicon: memristive crossbars, in-memory compute arrays, and emerging quantum and photonic devices. There are four critical challenges at this intersection: 1) integrating spiking dynamics with foundation models, 2) maintaining lifelong plasticity without catastrophic forgetting, 3) unifying language with sensorimotor learning in embodied agents, and 4) enforcing ethical safeguards in advanced neuromorphic autonomous systems. This combined perspective across neuroscience, computation, and hardware offers an integrative agenda for in each of these fields.

Task-trained recurrent neural networks (RNNs) are widely used in neuroscience and machine learning to model dynamical computations. To gain mechanistic insight into how neural systems solve tasks, prior work often reverse-engineers individual trained networks. However, different RNNs trained on the same task and achieving similar performance can exhibit strikingly different internal solutions, a phenomenon known as solution degeneracy. Here, we develop a unified framework to systematically quantify and control solution degeneracy across three levels: behavior, neural dynamics, and weight space. We apply this framework to 3,400 RNNs trained on four neuroscience-relevant tasks: flip-flop memory, sine wave generation, delayed discrimination, and path integration, while systematically varying task complexity, learning regime, network size, and regularization. We find that higher task complexity and stronger feature learning reduce degeneracy in neural dynamics but increase it in weight space, with mixed effects on behavior. In contrast, larger networks and structural regularization reduce degeneracy at all three levels. These findings empirically validate the Contravariance Principle and provide practical guidance for researchers seeking to tune the variability of RNN solutions, either to uncover shared neural mechanisms or to model the individual variability observed in biological systems. This work provides a principled framework for quantifying and controlling solution degeneracy in task-trained RNNs, offering new tools for building more interpretable and biologically grounded models of neural computation.

Spiking Neural Networks (SNNs) as one of the biology-inspired models have received much attention recently. It can significantly reduce energy consumption since they quantize the real-valued membrane potentials to 0/1 spikes to transmit information thus the multiplications of activations and weights can be replaced by additions when implemented on hardware. However, this quantization mechanism will inevitably introduce quantization error, thus causing catastrophic information loss. To address the quantization error problem, we propose a regularizing membrane potential loss (RMP-Loss) to adjust the distribution which is directly related to quantization error to a range close to the spikes. Our method is extremely simple to implement and straightforward to train an SNN. Furthermore, it is shown to consistently outperform previous state-of-the-art methods over different network architectures and datasets.

Sensory systems across all modalities and species exhibit adaptation to continuously changing input statistics. Individual neurons have been shown to modulate their response gains so as to maximize information transmission in different stimulus contexts. Experimental measurements have revealed additional, nuanced sensory adaptation effects including changes in response maxima and minima, tuning curve repulsion from the adapter stimulus, and stimulus-driven response decorrelation. Existing explanations of these phenomena rely on changes in inter-neuronal synaptic efficacy, which, while more flexible, are unlikely to operate as rapidly or reversibly as single neuron gain modulations. Using published V1 population adaptation data, we show that propagation of single neuron gain changes in a recurrent network is sufficient to capture the entire set of observed adaptation effects. We propose a novel adaptive efficient coding objective with which single neuron gains are modulated, maximizing the fidelity of the stimulus representation while minimizing overall activity in the network. From this objective, we analytically derive a set of gains that optimize the trade-off between preserving information about the stimulus and conserving metabolic resources. Our model generalizes well-established concepts of single neuron adaptive gain control to recurrent populations, and parsimoniously explains experimental adaptation data.

Neural network training can be accelerated when a learnable update rule is used in lieu of classic adaptive optimizers (e.g. Adam). However, learnable update rules can be costly and unstable to train and use. A simpler recently proposed approach to accelerate training is to use Adam for most of the optimization steps and periodically, only every few steps, nowcast (predict future) parameters. We improve this approach by Neuron interaction and Nowcasting (NiNo) networks. NiNo leverages neuron connectivity and graph neural networks to more accurately nowcast parameters by learning in a supervised way from a set of training trajectories over multiple tasks. We show that in some networks, such as Transformers, neuron connectivity is non-trivial. By accurately modeling neuron connectivity, we allow NiNo to accelerate Adam training by up to 50\% in vision and language tasks.

Spiking neural networks (SNNs) are known as a typical kind of brain-inspired models with their unique features of rich neuronal dynamics, diverse coding schemes and low power consumption properties. How to obtain a high-accuracy model has always been the main challenge in the field of SNN. Currently, there are two mainstream methods, i.e., obtaining a converted SNN through converting a well-trained Artificial Neural Network (ANN) to its SNN counterpart or training an SNN directly. However, the inference time of a converted SNN is too long, while SNN training is generally very costly and inefficient. In this work, a new SNN training paradigm is proposed by combining the concepts of the two different training methods with the help of the pretrain technique and BP-based deep SNN training mechanism. We believe that the proposed paradigm is a more efficient pipeline for training SNNs. The pipeline includes pipeS for static data transfer tasks and pipeD for dynamic data transfer tasks. SOTA results are obtained in a large-scale event-driven dataset ES-ImageNet. For training acceleration, we achieve the same (or higher) best accuracy as similar LIF-SNNs using 1/10 training time on ImageNet-1K and 2/5 training time on ES-ImageNet and also provide a time-accuracy benchmark for a new dataset ES-UCF101. These experimental results reveal the similarity of the functions of parameters between ANNs and SNNs and also demonstrate the various potential applications of this SNN training pipeline.

Attending to what is relevant is fundamental to both the mammalian brain and modern machine learning models such as Transformers. Yet, determining relevance remains a core challenge, traditionally offloaded to learning algorithms like backpropagation. Inspired by recent cellular neurobiological evidence linking neocortical pyramidal cells to distinct mental states, this work shows how models (e.g., Transformers) can emulate high-level perceptual processing and awake thought (imagination) states to pre-select relevant information before applying attention. Triadic neuronal-level modulation loops among questions (Q), clues (keys, K), and hypotheses (values, V) enable diverse, deep, parallel reasoning chains at the representation level and allow a rapid shift from initial biases to refined understanding. This leads to orders-of-magnitude faster learning with significantly reduced computational demand (e.g., fewer heads, layers, and tokens), at an approximate cost of O(N), where N is the number of input tokens. Results span reinforcement learning (e.g., CarRacing in a high-dimensional visual setup), computer vision, and natural language question answering.

Activity difference based learning algorithms-such as contrastive Hebbian learning and equilibrium propagation-have been proposed as biologically plausible alternatives to error back-propagation. However, on traditional digital chips these algorithms suffer from having to solve a costly inference problem twice, making these approaches more than two orders of magnitude slower than back-propagation. In the analog realm equilibrium propagation may be promising for fast and energy efficient learning, but states still need to be inferred and stored twice. Inspired by lifted neural networks and compartmental neuron models we propose a simple energy based compartmental neuron model, termed dual propagation, in which each neuron is a dyad with two intrinsic states. At inference time these intrinsic states encode the error/activity duality through their difference and their mean respectively. The advantage of this method is that only a single inference phase is needed and that inference can be solved in layerwise closed-form. Experimentally we show on common computer vision datasets, including Imagenet32x32, that dual propagation performs equivalently to back-propagation both in terms of accuracy and runtime.

The brain understands the external world through an internal model that generates predictions and refines them based on prediction errors. A complete prediction specifies what will happen, when it will happen, and with what probability, which we refer to as a "prediction object". Existing models typically capture only what and when, omit probabilities, and rely on biologically-implausible algorithms. Here we show that a single population of spiking neurons can jointly encode the prediction object through a biologically grounded learning mechanism. We implement a heterogeneous Izhikevich spiking reservoir with readouts trained by an error-modulated, attention-gated three-factor Hebbian rule and test it on a novel paradigm that controls both the timing and probability of upcoming stimuli. By integrating real-time learning of "when" with offline consolidation of "what", the model encodes the complete prediction object, firing at the correct times with magnitudes proportional to the probabilities. Critically, it rapidly adapts to changes in both stimulus timing and probability, an ability that global least-squares methods such as FORCE lack without explicit resets. During learning, the model self-organizes its readout weights into near-orthogonal subspaces for "what" and "when," showing that multiplexed encoding arises naturally from generic recurrent dynamics under local, error-gated modulation. These results challenge the view that "what" and "when" predictions require separate modules, suggesting instead that mixed selectivity within shared populations supports flexible predictive cognition. The model also predicts phase-specific neuromodulation and overlapping neural subspaces, offering a parsimonious alternative to hierarchical predictive-coding accounts.

Understanding the inner workings of neural networks is essential for enhancing model performance and interpretability. Current research predominantly focuses on examining the connection between individual neurons and the model's final predictions. Which suffers from challenges in interpreting the internal workings of the model, particularly when neurons encode multiple unrelated features. In this paper, we propose a novel framework that transitions the focus from analyzing individual neurons to investigating groups of neurons, shifting the emphasis from neuron-output relationships to functional interaction between neurons. Our automated framework, NeurFlow, first identifies core neurons and clusters them into groups based on shared functional relationships, enabling a more coherent and interpretable view of the network's internal processes. This approach facilitates the construction of a hierarchical circuit representing neuron interactions across layers, thus improving interpretability while reducing computational costs. Our extensive empirical studies validate the fidelity of our proposed NeurFlow. Additionally, we showcase its utility in practical applications such as image debugging and automatic concept labeling, thereby highlighting its potential to advance the field of neural network explainability.

In this paper, we describe a design of a mixed signal circuit for a binary neuron (a.k.a perceptron, threshold logic gate) and a methodology for automatically embedding such cells in ASICs. The binary neuron, referred to as an FTL (flash threshold logic) uses floating gate or flash transistors whose threshold voltages serve as a proxy for the weights of the neuron. Algorithms for mapping the weights to the flash transistor threshold voltages are presented. The threshold voltages are determined to maximize both the robustness of the cell and its speed. The performance, power, and area of a single FTL cell are shown to be significantly smaller (79.4%), consume less power (61.6%), and operate faster (40.3%) compared to conventional CMOS logic equivalents. Also included are the architecture and the algorithms to program the flash devices of an FTL. The FTL cells are implemented as standard cells, and are designed to allow commercial synthesis and P&R tools to automatically use them in synthesis of ASICs. Substantial reductions in area and power without sacrificing performance are demonstrated on several ASIC benchmarks by the automatic embedding of FTL cells. The paper also demonstrates how FTL cells can be used for fixing timing errors after fabrication.

The brain solves the credit assignment problem remarkably well. For credit to be assigned across neural networks they must, in principle, wait for specific neural computations to finish. How the brain deals with this inherent locking problem has remained unclear. Deep learning methods suffer from similar locking constraints both on the forward and feedback phase. Recently, decoupled neural interfaces (DNIs) were introduced as a solution to the forward and feedback locking problems in deep networks. Here we propose that a specialised brain region, the cerebellum, helps the cerebral cortex solve similar locking problems akin to DNIs. To demonstrate the potential of this framework we introduce a systems-level model in which a recurrent cortical network receives online temporal feedback predictions from a cerebellar module. We test this cortico-cerebellar recurrent neural network (ccRNN) model on a number of sensorimotor (line and digit drawing) and cognitive tasks (pattern recognition and caption generation) that have been shown to be cerebellar-dependent. In all tasks, we observe that ccRNNs facilitates learning while reducing ataxia-like behaviours, consistent with classical experimental observations. Moreover, our model also explains recent behavioural and neuronal observations while making several testable predictions across multiple levels. Overall, our work offers a novel perspective on the cerebellum as a brain-wide decoupling machine for efficient credit assignment and opens a new avenue between deep learning and neuroscience.

How neuronal circuits achieve credit assignment remains a central unsolved question in systems neuroscience. Various studies have suggested plausible solutions for back-propagating error signals through multi-layer networks. These purely functionally motivated models assume distinct neuronal compartments to represent local error signals that determine the sign of synaptic plasticity. However, this explicit error modulation is inconsistent with phenomenological plasticity models in which the sign depends primarily on postsynaptic activity. Here we show how a plausible microcircuit model and Hebbian learning rule derived within an adaptive control theory framework can resolve this discrepancy. Assuming errors are encoded in top-down dis-inhibitory synaptic afferents, we show that error-modulated learning emerges naturally at the circuit level when recurrent inhibition explicitly influences Hebbian plasticity. The same learning rule accounts for experimentally observed plasticity in the absence of inhibition and performs comparably to back-propagation of error (BP) on several non-linearly separable benchmarks. Our findings bridge the gap between functional and experimentally observed plasticity rules and make concrete predictions on inhibitory modulation of excitatory plasticity.

Artificial neural networks are being proposed as models of parts of the brain. The networks are compared to recordings of biological neurons, and good performance in reproducing neural responses is considered to support the model's validity. A key question is how much this system identification approach tells us about brain computation. Does it validate one model architecture over another? We evaluate the most commonly used comparison techniques, such as a linear encoding model and centered kernel alignment, to correctly identify a model by replacing brain recordings with known ground truth models. System identification performance is quite variable; it also depends significantly on factors independent of the ground truth architecture, such as stimuli images. In addition, we show the limitations of using functional similarity scores in identifying higher-level architectural motifs.

Neuromorphic computing with spiking neural networks is promising for energy-efficient artificial intelligence (AI) applications. However, different from humans who continually learn different tasks in a lifetime, neural network models suffer from catastrophic forgetting. How could neuronal operations solve this problem is an important question for AI and neuroscience. Many previous studies draw inspiration from observed neuroscience phenomena and propose episodic replay or synaptic metaplasticity, but they are not guaranteed to explicitly preserve knowledge for neuron populations. Other works focus on machine learning methods with more mathematical grounding, e.g., orthogonal projection on high dimensional spaces, but there is no neural correspondence for neuromorphic computing. In this work, we develop a new method with neuronal operations based on lateral connections and Hebbian learning, which can protect knowledge by projecting activity traces of neurons into an orthogonal subspace so that synaptic weight update will not interfere with old tasks. We show that Hebbian and anti-Hebbian learning on recurrent lateral connections can effectively extract the principal subspace of neural activities and enable orthogonal projection. This provides new insights into how neural circuits and Hebbian learning can help continual learning, and also how the concept of orthogonal projection can be realized in neuronal systems. Our method is also flexible to utilize arbitrary training methods based on presynaptic activities/traces. Experiments show that our method consistently solves forgetting for spiking neural networks with nearly zero forgetting under various supervised training methods with different error propagation approaches, and outperforms previous approaches under various settings. Our method can pave a solid path for building continual neuromorphic computing systems.

We present PyCARL, a PyNN-based common Python programming interface for hardware-software co-simulation of spiking neural network (SNN). Through PyCARL, we make the following two key contributions. First, we provide an interface of PyNN to CARLsim, a computationally-efficient, GPU-accelerated and biophysically-detailed SNN simulator. PyCARL facilitates joint development of machine learning models and code sharing between CARLsim and PyNN users, promoting an integrated and larger neuromorphic community. Second, we integrate cycle-accurate models of state-of-the-art neuromorphic hardware such as TrueNorth, Loihi, and DynapSE in PyCARL, to accurately model hardware latencies that delay spikes between communicating neurons and degrade performance. PyCARL allows users to analyze and optimize the performance difference between software-only simulation and hardware-software co-simulation of their machine learning models. We show that system designers can also use PyCARL to perform design-space exploration early in the product development stage, facilitating faster time-to-deployment of neuromorphic products. We evaluate the memory usage and simulation time of PyCARL using functionality tests, synthetic SNNs, and realistic applications. Our results demonstrate that for large SNNs, PyCARL does not lead to any significant overhead compared to CARLsim. We also use PyCARL to analyze these SNNs for a state-of-the-art neuromorphic hardware and demonstrate a significant performance deviation from software-only simulations. PyCARL allows to evaluate and minimize such differences early during model development.

The coding mechanism of sensory memory on the neuron scale is one of the most important questions in neuroscience. We have put forward a quantitative neural network model, which is self organized, self similar, and self adaptive, just like an ecosystem following Darwin theory. According to this model, neural coding is a mult to one mapping from objects to neurons. And the whole cerebrum is a real-time statistical Turing Machine, with powerful representing and learning ability. This model can reconcile some important disputations, such as: temporal coding versus rate based coding, grandmother cell versus population coding, and decay theory versus interference theory. And it has also provided explanations for some key questions such as memory consolidation, episodic memory, consciousness, and sentiment. Philosophical significance is indicated at last.

Spiking Transformers have recently emerged as promising architectures for combining the efficiency of spiking neural networks with the representational power of self-attention. However, the lack of standardized implementations, evaluation pipelines, and consistent design choices has hindered fair comparison and principled analysis. In this paper, we introduce STEP, a unified benchmark framework for Spiking Transformers that supports a wide range of tasks, including classification, segmentation, and detection across static, event-based, and sequential datasets. STEP provides modular support for diverse components such as spiking neurons, input encodings, surrogate gradients, and multiple backends (e.g., SpikingJelly, BrainCog). Using STEP, we reproduce and evaluate several representative models, and conduct systematic ablation studies on attention design, neuron types, encoding schemes, and temporal modeling capabilities. We also propose a unified analytical model for energy estimation, accounting for spike sparsity, bitwidth, and memory access, and show that quantized ANNs may offer comparable or better energy efficiency. Our results suggest that current Spiking Transformers rely heavily on convolutional frontends and lack strong temporal modeling, underscoring the need for spike-native architectural innovations. The full code is available at: https://github.com/Fancyssc/STEP

Interpreting the information encoded in model weights remains a fundamental challenge in mechanistic interpretability. In this work, we introduce ROTATE (Rotation-Optimized Token Alignment in weighT spacE), a data-free method requiring no forward passes that disentangles MLP neurons directly in weight space. Our approach relies on a key statistical observation: neurons that encode coherent, monosemantic concepts exhibit high kurtosis when projected onto the model's vocabulary. By optimizing rotations of neuron weights to maximize their vocabulary-space kurtosis, our method recovers sparse, interpretable directions which we name vocabulary channels. Experiments on Llama-3.1-8B-Instruct and Gemma-2-2B-it demonstrate that ROTATE consistently recovers vocabulary channels that are faithful to the neuron's behavior. ablating individual channels selectively disables corresponding input activations or the promotion of specific concepts. Moreover, aggregating channel-level descriptions yields comprehensive neuron descriptions that outperform optimized activation-based baselines by 2-3x in head-to-head comparisons. By providing a data-free decomposition of neuron weights, ROTATE offers a scalable, fine-grained building block for interpreting LMs.

Hopfield neural networks are a possible basis for modelling associative memory in living organisms. After summarising previous studies in the field, we take a new look at learning rules, exhibiting them as descent-type algorithms for various cost functions. We also propose several new cost functions suitable for learning. We discuss the role of biases (the external inputs) in the learning process in Hopfield networks. Furthermore, we apply Newtons method for learning memories, and experimentally compare the performances of various learning rules. Finally, to add to the debate whether allowing connections of a neuron to itself enhances memory capacity, we numerically investigate the effects of self coupling. Keywords: Hopfield Networks, associative memory, content addressable memory, learning rules, gradient descent, attractor networks

Natural language is an appealing medium for explaining how large language models process and store information, but evaluating the faithfulness of such explanations is challenging. To help address this, we develop two modes of evaluation for natural language explanations that claim individual neurons represent a concept in a text input. In the observational mode, we evaluate claims that a neuron a activates on all and only input strings that refer to a concept picked out by the proposed explanation E. In the intervention mode, we construe E as a claim that the neuron a is a causal mediator of the concept denoted by E. We apply our framework to the GPT-4-generated explanations of GPT-2 XL neurons of Bills et al. (2023) and show that even the most confident explanations have high error rates and little to no causal efficacy. We close the paper by critically assessing whether natural language is a good choice for explanations and whether neurons are the best level of analysis.

Recent years have witnessed Spiking Neural Networks (SNNs) gaining attention for their ultra-low energy consumption and high biological plausibility compared with traditional Artificial Neural Networks (ANNs). Despite their distinguished properties, the application of SNNs in the computationally intensive field of image generation is still under exploration. In this paper, we propose the Spiking Diffusion Models (SDMs), an innovative family of SNN-based generative models that excel in producing high-quality samples with significantly reduced energy consumption. In particular, we propose a Temporal-wise Spiking Mechanism (TSM) that allows SNNs to capture more temporal features from a bio-plasticity perspective. In addition, we propose a threshold-guided strategy that can further improve the performances by up to 16.7% without any additional training. We also make the first attempt to use the ANN-SNN approach for SNN-based generation tasks. Extensive experimental results reveal that our approach not only exhibits comparable performance to its ANN counterpart with few spiking time steps, but also outperforms previous SNN-based generative models by a large margin. Moreover, we also demonstrate the high-quality generation ability of SDM on large-scale datasets, e.g., LSUN bedroom. This development marks a pivotal advancement in the capabilities of SNN-based generation, paving the way for future research avenues to realize low-energy and low-latency generative applications. Our code is available at https://github.com/AndyCao1125/SDM.

Objective: Target identification in brain-computer interface (BCI) spellers refers to the electroencephalogram (EEG) classification for predicting the target character that the subject intends to spell. When the visual stimulus of each character is tagged with a distinct frequency, the EEG records steady-state visually evoked potentials (SSVEP) whose spectrum is dominated by the harmonics of the target frequency. In this setting, we address the target identification and propose a novel deep neural network (DNN) architecture. Method: The proposed DNN processes the multi-channel SSVEP with convolutions across the sub-bands of harmonics, channels, time, and classifies at the fully connected layer. We test with two publicly available large scale (the benchmark and BETA) datasets consisting of in total 105 subjects with 40 characters. Our first stage training learns a global model by exploiting the statistical commonalities among all subjects, and the second stage fine tunes to each subject separately by exploiting the individualities. Results: Our DNN achieves impressive information transfer rates (ITRs) on both datasets, 265.23 bits/min and 196.59 bits/min, respectively, with only 0.4 seconds of stimulation. The code is available for reproducibility at https://github.com/osmanberke/Deep-SSVEP-BCI. Conclusion: The presented DNN strongly outperforms the state-of-the-art techniques as our accuracy and ITR rates are the highest ever reported performance results on these datasets. Significance: Due to its unprecedentedly high speller ITRs and flawless applicability to general SSVEP systems, our technique has great potential in various biomedical engineering settings of BCIs such as communication, rehabilitation and control.

The Backpropagation algorithm, widely used to train neural networks, has often been criticised for its lack of biological realism. In an attempt to find a more biologically plausible alternative, and avoid to back-propagate gradients in favour of using local learning rules, the recently introduced Forward-Forward algorithm replaces the traditional forward and backward passes of Backpropagation with two forward passes. In this work, we show that internal representations obtained with the Forward-Forward algorithm organize into robust, category-specific ensembles, composed by an extremely low number of active units (high sparsity). This is remarkably similar to what is observed in cortical representations during sensory processing. While not found in models trained with standard Backpropagation, sparsity emerges also in networks optimized by Backpropagation, on the same training objective of Forward-Forward. These results suggest that the learning procedure proposed by Forward-Forward may be superior to Backpropagation in modelling learning in the cortex, even when a backward pass is used.

We can better understand deep neural networks by identifying which features each of their neurons have learned to detect. To do so, researchers have created Deep Visualization techniques including activation maximization, which synthetically generates inputs (e.g. images) that maximally activate each neuron. A limitation of current techniques is that they assume each neuron detects only one type of feature, but we know that neurons can be multifaceted, in that they fire in response to many different types of features: for example, a grocery store class neuron must activate either for rows of produce or for a storefront. Previous activation maximization techniques constructed images without regard for the multiple different facets of a neuron, creating inappropriate mixes of colors, parts of objects, scales, orientations, etc. Here, we introduce an algorithm that explicitly uncovers the multiple facets of each neuron by producing a synthetic visualization of each of the types of images that activate a neuron. We also introduce regularization methods that produce state-of-the-art results in terms of the interpretability of images obtained by activation maximization. By separately synthesizing each type of image a neuron fires in response to, the visualizations have more appropriate colors and coherent global structure. Multifaceted feature visualization thus provides a clearer and more comprehensive description of the role of each neuron.

Spiking neural networks (SNN) are a biologically inspired model of neural networks with certain brain-like properties. In the past few decades, this model has received increasing attention in computer science community, owing also to the successful phenomenon of deep learning. In SNN, communication between neurons takes place through the spikes and spike trains. This differentiates these models from the ``standard'' artificial neural networks (ANN) where the frequency of spikes is replaced by real-valued signals. Spiking neural P systems (SNPS) can be considered a branch of SNN based more on the principles of formal automata, with many variants developed within the framework of the membrane computing theory. In this paper, we first briefly compare structure and function, advantages and drawbacks of SNN and SNPS. A key part of the article is a survey of recent results and applications of machine learning and deep learning models of both SNN and SNPS formalisms.

Redundancies and correlations in the responses of sensory neurons seem to waste neural resources but can carry cues about structured stimuli and may help the brain to correct for response errors. To assess how the retina negotiates this tradeoff, we measured simultaneous responses from populations of ganglion cells presented with natural and artificial stimuli that varied greatly in correlation structure. We found that pairwise correlations in the retinal output remained similar across stimuli with widely different spatio-temporal correlations including white noise and natural movies. Meanwhile, purely spatial correlations tended to increase correlations in the retinal response. Responding to more correlated stimuli, ganglion cells had faster temporal kernels and tended to have stronger surrounds. These properties of individual cells, along with gain changes that opposed changes in effective contrast at the ganglion cell input, largely explained the similarity of pairwise correlations across stimuli where receptive field measurements were possible.

Spiking Neural Networks (SNNs) are computational models inspired by the structure and dynamics of biological neuronal networks. Their event-driven nature enables them to achieve high energy efficiency, particularly when deployed on neuromorphic hardware platforms. Unlike conventional Artificial Neural Networks (ANNs), which primarily rely on layered architectures, SNNs naturally support a wide range of connectivity patterns, from traditional layered structures to small-world graphs characterized by locally dense and globally sparse connections. In this work, we introduce NeuroCoreX, an FPGA-based emulator designed for the flexible co-design and testing of SNNs. NeuroCoreX supports all-to-all connectivity, providing the capability to implement diverse network topologies without architectural restrictions. It features a biologically motivated local learning mechanism based on Spike-Timing-Dependent Plasticity (STDP). The neuron model implemented within NeuroCoreX is the Leaky Integrate-and-Fire (LIF) model, with current-based synapses facilitating spike integration and transmission . A Universal Asynchronous Receiver-Transmitter (UART) interface is provided for programming and configuring the network parameters, including neuron, synapse, and learning rule settings. Users interact with the emulator through a simple Python-based interface, streamlining SNN deployment from model design to hardware execution. NeuroCoreX is released as an open-source framework, aiming to accelerate research and development in energy-efficient, biologically inspired computing.