Daily Papers

Selecting appropriate datasets is critical in modern computer vision. However, no general-purpose tools exist to evaluate the extent to which two datasets differ. For this, we propose representing images - and by extension datasets - using Distributions of Neuron Activations (DNAs). DNAs fit distributions, such as histograms or Gaussians, to activations of neurons in a pre-trained feature extractor through which we pass the image(s) to represent. This extractor is frozen for all datasets, and we rely on its generally expressive power in feature space. By comparing two DNAs, we can evaluate the extent to which two datasets differ with granular control over the comparison attributes of interest, providing the ability to customise the way distances are measured to suit the requirements of the task at hand. Furthermore, DNAs are compact, representing datasets of any size with less than 15 megabytes. We demonstrate the value of DNAs by evaluating their applicability on several tasks, including conditional dataset comparison, synthetic image evaluation, and transfer learning, and across diverse datasets, ranging from synthetic cat images to celebrity faces and urban driving scenes.

Large language models (LLMs) have revolutionized software development practices, yet concerns about their safety have arisen, particularly regarding hidden backdoors, aka trojans. Backdoor attacks involve the insertion of triggers into training data, allowing attackers to manipulate the behavior of the model maliciously. In this paper, we focus on analyzing the model parameters to detect potential backdoor signals in code models. Specifically, we examine attention weights and biases, activation values, and context embeddings of the clean and poisoned CodeBERT models. Our results suggest noticeable patterns in activation values and context embeddings of poisoned samples for the poisoned CodeBERT model; however, attention weights and biases do not show any significant differences. This work contributes to ongoing efforts in white-box detection of backdoor signals in LLMs of code through the analysis of parameters and activations.

Instruction tuning improves the ability of large language models (LLMs) to follow diverse human instructions, but achieving strong performance on specific target tasks remains challenging. A critical bottleneck is selecting the most relevant data to maximize task-specific performance. Existing data selection approaches include unstable influence-based methods and more stable distribution alignment methods, the latter of which critically rely on the underlying sample representation. In practice, most distribution alignment methods, from shallow features (e.g., BM25) to neural embeddings (e.g., BGE, LLM2Vec), may fail to capture how the model internally processes samples. To bridge this gap, we adopt a model-centric strategy in which each sample is represented by its neuronal activation pattern in the model, directly reflecting internal computation. However, directly using raw neuron activations leads to spurious similarity between unrelated samples due to neuron polysemanticity, where a single neuron may respond to multiple, unrelated concepts. To address this, we employ sparse autoencoders to disentangle polysemantic activations into sparse, monosemantic representations, and introduce a dedicated similarity metric for this space to better identify task-relevant data. Comprehensive experiments across multiple instruction datasets, models, tasks, and selection ratios show that our approach consistently outperforms existing data selection baselines in both stability and task-specific performance.

Safety fine-tuning algorithms are commonly used to fine-tune language models to reduce harmful outputs, but the exact internal mechanisms of how those models achieve this remain unclear. In studying direct preference optimisation (DPO) for toxicity reduction, current explanations claim that DPO works by dampening the most toxic MLP neurons to learn an offset to avert toxic regions in the residual stream. However, by ablating the most toxic neurons and applying activation patching, we find this explanation incomplete. By projecting neuron activation changes onto a toxicity probe, we find that only 31.8\% of toxicity reduction comes from dampened toxic neurons. Instead, DPO reduces toxicity by accumulating effects across multiple neuron groups, both reducing writing in the toxic direction and promoting anti-toxicity in the residual stream. Moreover, DPO gives noisy adjustments to neuron activations, with many neurons actually increasing toxicity. This indicates that DPO is a balancing process between opposing neuron effects to achieve toxicity reduction.

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.

Several studies have explored the mechanisms of large language models (LLMs) in coding tasks, but most have focused on programming languages (PLs) in a monolingual setting. In this paper, we investigate the relationship between multiple PLs and English in the concept space of LLMs. We perform a few-shot translation task on 21 PL pairs using two Llama-based models. By decoding the embeddings of intermediate layers during this task, we observe that the concept space is closer to English (including PL keywords) and assigns high probabilities to English tokens in the second half of the intermediate layers. We analyze neuron activations for 11 PLs and English, finding that while language-specific neurons are primarily concentrated in the bottom layers, those exclusive to each PL tend to appear in the top layers. For PLs that are highly aligned with multiple other PLs, identifying language-specific neurons is not feasible. These PLs also tend to have a larger keyword set than other PLs and are closer to the model's concept space regardless of the input/output PL in the translation task. Our findings provide insights into how LLMs internally represent PLs, revealing structural patterns in the model's concept space. Code is available at https://github.com/cisnlp/code-specific-neurons.

Large Language Models have demonstrated impressive fluency across diverse tasks, yet their tendency to produce toxic content remains a critical challenge for AI safety and public trust. Existing toxicity mitigation approaches primarily manipulate individual neuron activations, but these methods suffer from instability, context dependence, and often compromise the model's core language abilities. To address these shortcomings, we investigate three key questions: the stability of neuron-level toxicity indicators, the advantages of structural (layer-wise) representations, and the interpretability of mechanisms driving toxic generation. Through extensive experiments on Jigsaw and ToxiCN datasets, we show that aggregated layer-wise features provide more robust signals than single neurons. Moreover, we observe conceptual limitations in prior works that conflate toxicity detection experts and generation experts within neuron-based interventions. To mitigate this, we propose a novel principled intervention technique, EigenShift, based on eigen-decomposition of the language model's final output layer. This method selectively targets generation-aligned components, enabling precise toxicity suppression without impairing linguistic competence. Our method requires no additional training or fine-tuning, incurs minimal computational cost, and is grounded in rigorous theoretical analysis.

Despite strong performance in audio perception tasks, large audio-language models (AudioLLMs) remain opaque to interpretation. A major factor behind this lack of interpretability is that individual neurons in these models frequently activate in response to several unrelated concepts. We introduce the first mechanistic interpretability framework for AudioLLMs, leveraging sparse autoencoders (SAEs) to disentangle polysemantic activations into monosemantic features. Our pipeline identifies representative audio clips, assigns meaningful names via automated captioning, and validates concepts through human evaluation and steering. Experiments show that AudioLLMs encode structured and interpretable features, enhancing transparency and control. This work provides a foundation for trustworthy deployment in high-stakes domains and enables future extensions to larger models, multilingual audio, and more fine-grained paralinguistic features. Project URL: https://townim-faisal.github.io/AutoInterpret-AudioLLM/

Recent advancements have scaled neural networks to unprecedented sizes, achieving remarkable performance across a wide range of tasks. However, deploying these large-scale models on resource-constrained devices poses significant challenges due to substantial storage and computational requirements. Neural network pruning has emerged as an effective technique to mitigate these limitations by reducing model size and complexity. In this paper, we introduce an intuitive and interpretable pruning method based on activation statistics, rooted in information theory and statistical analysis. Our approach leverages the statistical properties of neuron activations to identify and remove weights with minimal contributions to neuron outputs. Specifically, we build a distribution of weight contributions across the dataset and utilize its parameters to guide the pruning process. Furthermore, we propose a Pruning-aware Training strategy that incorporates an additional regularization term to enhance the effectiveness of our pruning method. Extensive experiments on multiple datasets and network architectures demonstrate that our method consistently outperforms several baseline and state-of-the-art pruning techniques.

Facial sketch synthesis (FSS) aims to generate a vivid sketch portrait from a given facial photo. Existing FSS methods merely rely on 2D representations of facial semantic or appearance. However, professional human artists usually use outlines or shadings to covey 3D geometry. Thus facial 3D geometry (e.g. depth map) is extremely important for FSS. Besides, different artists may use diverse drawing techniques and create multiple styles of sketches; but the style is globally consistent in a sketch. Inspired by such observations, in this paper, we propose a novel Human-Inspired Dynamic Adaptation (HIDA) method. Specially, we propose to dynamically modulate neuron activations based on a joint consideration of both facial 3D geometry and 2D appearance, as well as globally consistent style control. Besides, we use deformable convolutions at coarse-scales to align deep features, for generating abstract and distinct outlines. Experiments show that HIDA can generate high-quality sketches in multiple styles, and significantly outperforms previous methods, over a large range of challenging faces. Besides, HIDA allows precise style control of the synthesized sketch, and generalizes well to natural scenes and other artistic styles. Our code and results have been released online at: https://github.com/AiArt-HDU/HIDA.

The alignment of vision-language representations endows current Vision-Language Models (VLMs) with strong multi-modal reasoning capabilities. However, the interpretability of the alignment component remains uninvestigated due to the difficulty in mapping the semantics of multi-modal representations into a unified concept set. To address this problem, we propose VL-SAE, a sparse autoencoder that encodes vision-language representations into its hidden activations. Each neuron in its hidden layer correlates to a concept represented by semantically similar images and texts, thereby interpreting these representations with a unified concept set. To establish the neuron-concept correlation, we encourage semantically similar representations to exhibit consistent neuron activations during self-supervised training. First, to measure the semantic similarity of multi-modal representations, we perform their alignment in an explicit form based on cosine similarity. Second, we construct the VL-SAE with a distance-based encoder and two modality-specific decoders to ensure the activation consistency of semantically similar representations. Experiments across multiple VLMs (e.g., CLIP, LLaVA) demonstrate the superior capability of VL-SAE in interpreting and enhancing the vision-language alignment. For interpretation, the alignment between vision and language representations can be understood by comparing their semantics with concepts. For enhancement, the alignment can be strengthened by aligning vision-language representations at the concept level, contributing to performance improvements in downstream tasks, including zero-shot image classification and hallucination elimination. Codes are available at https://github.com/ssfgunner/VL-SAE.

We analyze the storage and recall of factual associations in autoregressive transformer language models, finding evidence that these associations correspond to localized, directly-editable computations. We first develop a causal intervention for identifying neuron activations that are decisive in a model's factual predictions. This reveals a distinct set of steps in middle-layer feed-forward modules that mediate factual predictions while processing subject tokens. To test our hypothesis that these computations correspond to factual association recall, we modify feed-forward weights to update specific factual associations using Rank-One Model Editing (ROME). We find that ROME is effective on a standard zero-shot relation extraction (zsRE) model-editing task, comparable to existing methods. To perform a more sensitive evaluation, we also evaluate ROME on a new dataset of counterfactual assertions, on which it simultaneously maintains both specificity and generalization, whereas other methods sacrifice one or another. Our results confirm an important role for mid-layer feed-forward modules in storing factual associations and suggest that direct manipulation of computational mechanisms may be a feasible approach for model editing. The code, dataset, visualizations, and an interactive demo notebook are available at https://rome.baulab.info/

Large language models (LLMs) have achieved impressive results in natural language processing but are prone to memorizing portions of their training data, which can compromise evaluation metrics, raise privacy concerns, and limit generalization. Traditional methods for detecting memorization rely on output probabilities or loss functions, often lacking precision due to confounding factors like common language patterns. In this paper, we introduce an analytical method that precisely detects memorization by examining neuron activations within the LLM. By identifying specific activation patterns that differentiate between memorized and not memorized tokens, we train classification probes that achieve near-perfect accuracy. The approach can also be applied to other mechanisms, such as repetition, as demonstrated in this study, highlighting its versatility. Intervening on these activations allows us to suppress memorization without degrading overall performance, enhancing evaluation integrity by ensuring metrics reflect genuine generalization. Additionally, our method supports large-scale labeling of tokens and sequences, crucial for next-generation AI models, improving training efficiency and results. Our findings contribute to model interpretability and offer practical tools for analyzing and controlling internal mechanisms in LLMs.

An important issue with Large Language Models (LLMs) is their undesired ability to generate toxic language. In this work, we show that the neurons responsible for toxicity can be determined by their power to discriminate toxic sentences, and that toxic language can be mitigated by reducing their activation levels proportionally to this power. We propose AUROC adaptation (AurA), an intervention that can be applied to any pre-trained LLM to mitigate toxicity. As the intervention is proportional to the ability of each neuron to discriminate toxic content, it is free of any model-dependent hyperparameters. We show that AurA can achieve up to 2.2 times reduction in toxicity with only a 0.72 perplexity increase. We also show that AurA is effective with models of different scale (from 1.5B to 40B parameters), and its effectiveness in mitigating toxic language, while preserving common-sense zero-shot abilities, holds across all scales. AurA can be combined with pre-prompting strategies, boosting its average mitigation potential from 1.28times to 2.35times. Moreover, AurA can counteract adversarial pre-prompts that maliciously elicit toxic content, making it an effective method for deploying safer and less toxic models.

What is the right supervisory signal to train visual representations? Current approaches in computer vision use category labels from datasets such as ImageNet to train ConvNets. However, in case of biological agents, visual representation learning does not require millions of semantic labels. We argue that biological agents use physical interactions with the world to learn visual representations unlike current vision systems which just use passive observations (images and videos downloaded from web). For example, babies push objects, poke them, put them in their mouth and throw them to learn representations. Towards this goal, we build one of the first systems on a Baxter platform that pushes, pokes, grasps and observes objects in a tabletop environment. It uses four different types of physical interactions to collect more than 130K datapoints, with each datapoint providing supervision to a shared ConvNet architecture allowing us to learn visual representations. We show the quality of learned representations by observing neuron activations and performing nearest neighbor retrieval on this learned representation. Quantitatively, we evaluate our learned ConvNet on image classification tasks and show improvements compared to learning without external data. Finally, on the task of instance retrieval, our network outperforms the ImageNet network on recall@1 by 3%

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.

Deep Learning has revolutionized vision via convolutional neural networks (CNNs) and natural language processing via recurrent neural networks (RNNs). However, success stories of Deep Learning with standard feed-forward neural networks (FNNs) are rare. FNNs that perform well are typically shallow and, therefore cannot exploit many levels of abstract representations. We introduce self-normalizing neural networks (SNNs) to enable high-level abstract representations. While batch normalization requires explicit normalization, neuron activations of SNNs automatically converge towards zero mean and unit variance. The activation function of SNNs are "scaled exponential linear units" (SELUs), which induce self-normalizing properties. Using the Banach fixed-point theorem, we prove that activations close to zero mean and unit variance that are propagated through many network layers will converge towards zero mean and unit variance -- even under the presence of noise and perturbations. This convergence property of SNNs allows to (1) train deep networks with many layers, (2) employ strong regularization, and (3) to make learning highly robust. Furthermore, for activations not close to unit variance, we prove an upper and lower bound on the variance, thus, vanishing and exploding gradients are impossible. We compared SNNs on (a) 121 tasks from the UCI machine learning repository, on (b) drug discovery benchmarks, and on (c) astronomy tasks with standard FNNs and other machine learning methods such as random forests and support vector machines. SNNs significantly outperformed all competing FNN methods at 121 UCI tasks, outperformed all competing methods at the Tox21 dataset, and set a new record at an astronomy data set. The winning SNN architectures are often very deep. Implementations are available at: github.com/bioinf-jku/SNNs.

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.

Learning predictive models from observations using deep neural networks (DNNs) is a promising new approach to many real-world planning and control problems. However, common DNNs are too unstructured for effective planning, and current control methods typically rely on extensive sampling or local gradient descent. In this paper, we propose a new framework for integrated model learning and predictive control that is amenable to efficient optimization algorithms. Specifically, we start with a ReLU neural model of the system dynamics and, with minimal losses in prediction accuracy, we gradually sparsify it by removing redundant neurons. This discrete sparsification process is approximated as a continuous problem, enabling an end-to-end optimization of both the model architecture and the weight parameters. The sparsified model is subsequently used by a mixed-integer predictive controller, which represents the neuron activations as binary variables and employs efficient branch-and-bound algorithms. Our framework is applicable to a wide variety of DNNs, from simple multilayer perceptrons to complex graph neural dynamics. It can efficiently handle tasks involving complicated contact dynamics, such as object pushing, compositional object sorting, and manipulation of deformable objects. Numerical and hardware experiments show that, despite the aggressive sparsification, our framework can deliver better closed-loop performance than existing state-of-the-art methods.

Large-scale text-to-image diffusion models excel in generating high-quality images from textual inputs, yet concerns arise as research indicates their tendency to memorize and replicate training data, raising We also addressed the issue of memorization in diffusion models, where models tend to replicate exact training samples raising copyright infringement and privacy issues. Efforts within the text-to-image community to address memorization explore causes such as data duplication, replicated captions, or trigger tokens, proposing per-prompt inference-time or training-time mitigation strategies. In this paper, we focus on the feed-forward layers and begin by contrasting neuron activations of a set of memorized and non-memorized prompts. Experiments reveal a surprising finding: many different sets of memorized prompts significantly activate a common subspace in the model, demonstrating, for the first time, that memorization in the diffusion models lies in a special subspace. Subsequently, we introduce a novel post-hoc method for editing pre-trained models, whereby memorization is mitigated through the straightforward pruning of weights in specialized subspaces, avoiding the need to disrupt the training or inference process as seen in prior research. Finally, we demonstrate the robustness of the pruned model against training data extraction attacks, thereby unveiling new avenues for a practical and one-for-all solution to memorization.

Explainable recommendation systems are important to enhance transparency, accuracy, and fairness. Beyond result-level explanations, model-level interpretations can provide valuable insights that allow developers to optimize system designs and implement targeted improvements. However, most current approaches depend on specialized model designs, which often lack generalization capabilities. Given the various kinds of recommendation models, existing methods have limited ability to effectively interpret them. To address this issue, we propose RecSAE, an automatic, generalizable probing method for interpreting the internal states of Recommendation models with Sparse AutoEncoder. RecSAE serves as a plug-in module that does not affect original models during interpretations, while also enabling predictable modifications to their behaviors based on interpretation results. Firstly, we train an autoencoder with sparsity constraints to reconstruct internal activations of recommendation models, making the RecSAE latents more interpretable and monosemantic than the original neuron activations. Secondly, we automated the construction of concept dictionaries based on the relationship between latent activations and input item sequences. Thirdly, RecSAE validates these interpretations by predicting latent activations on new item sequences using the concept dictionary and deriving interpretation confidence scores from precision and recall. We demonstrate RecSAE's effectiveness on two datasets, identifying hundreds of highly interpretable concepts from pure ID-based models. Latent ablation studies further confirm that manipulating latent concepts produces corresponding changes in model output behavior, underscoring RecSAE's utility for both understanding and targeted tuning recommendation models. Code and data are publicly available at https://github.com/Alice1998/RecSAE.

Despite rapid adoption and deployment of large language models (LLMs), the internal computations of these models remain opaque and poorly understood. In this work, we seek to understand how high-level human-interpretable features are represented within the internal neuron activations of LLMs. We train k-sparse linear classifiers (probes) on these internal activations to predict the presence of features in the input; by varying the value of k we study the sparsity of learned representations and how this varies with model scale. With k=1, we localize individual neurons which are highly relevant for a particular feature, and perform a number of case studies to illustrate general properties of LLMs. In particular, we show that early layers make use of sparse combinations of neurons to represent many features in superposition, that middle layers have seemingly dedicated neurons to represent higher-level contextual features, and that increasing scale causes representational sparsity to increase on average, but there are multiple types of scaling dynamics. In all, we probe for over 100 unique features comprising 10 different categories in 7 different models spanning 70 million to 6.9 billion parameters.

While the activations of neurons in deep neural networks usually do not have a simple human-understandable interpretation, sparse autoencoders (SAEs) can be used to transform these activations into a higher-dimensional latent space which may be more easily interpretable. However, these SAEs can have millions of distinct latent features, making it infeasible for humans to manually interpret each one. In this work, we build an open-source automated pipeline to generate and evaluate natural language explanations for SAE features using LLMs. We test our framework on SAEs of varying sizes, activation functions, and losses, trained on two different open-weight LLMs. We introduce five new techniques to score the quality of explanations that are cheaper to run than the previous state of the art. One of these techniques, intervention scoring, evaluates the interpretability of the effects of intervening on a feature, which we find explains features that are not recalled by existing methods. We propose guidelines for generating better explanations that remain valid for a broader set of activating contexts, and discuss pitfalls with existing scoring techniques. We use our explanations to measure the semantic similarity of independently trained SAEs, and find that SAEs trained on nearby layers of the residual stream are highly similar. Our large-scale analysis confirms that SAE latents are indeed much more interpretable than neurons, even when neurons are sparsified using top-k postprocessing. Our code is available at https://github.com/EleutherAI/sae-auto-interp, and our explanations are available at https://huggingface.co/datasets/EleutherAI/auto_interp_explanations.

Vision-language models (VLMs) achieve strong multimodal performance, yet how computation is organized across populations of neurons remains poorly understood. In this work, we study VLMs through the lens of neural topology, representing each layer as a within-layer correlation graph derived from neuron-neuron co-activations. This view allows us to ask whether population-level structure is behaviorally meaningful, how it changes across modalities and depth, and whether it identifies causally influential internal components under intervention. We show that correlation topology carries recoverable behavioral signal; moreover, cross-modal structure progressively consolidates with depth around a compact set of recurrent hub neurons, whose targeted perturbation substantially alters model output. Neural topology thus emerges as a meaningful intermediate scale for VLM interpretability: richer than local attribution, more tractable than full circuit recovery, and empirically tied to multimodal behavior. Code is publicly available at https://github.com/he-h/vlm-graph-probing.

A central goal for mechanistic interpretability has been to identify the right units of analysis in large language models (LLMs) that causally explain their outputs. While early work focused on individual neurons, evidence that neurons often encode multiple concepts has motivated a shift toward analyzing directions in activation space. A key question is how to find directions that capture interpretable features in an unsupervised manner. Current methods rely on dictionary learning with sparse autoencoders (SAEs), commonly trained over residual stream activations to learn directions from scratch. However, SAEs often struggle in causal evaluations and lack intrinsic interpretability, as their learning is not explicitly tied to the computations of the model. Here, we tackle these limitations by directly decomposing MLP activations with semi-nonnegative matrix factorization (SNMF), such that the learned features are (a) sparse linear combinations of co-activated neurons, and (b) mapped to their activating inputs, making them directly interpretable. Experiments on Llama 3.1, Gemma 2 and GPT-2 show that SNMF derived features outperform SAEs and a strong supervised baseline (difference-in-means) on causal steering, while aligning with human-interpretable concepts. Further analysis reveals that specific neuron combinations are reused across semantically-related features, exposing a hierarchical structure in the MLP's activation space. Together, these results position SNMF as a simple and effective tool for identifying interpretable features and dissecting concept representations in LLMs.

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.

We study the problem of learning a single neuron with respect to the L_2^2-loss in the presence of adversarial label noise. We give an efficient algorithm that, for a broad family of activations including ReLUs, approximates the optimal L_2^2-error within a constant factor. Our algorithm applies under much milder distributional assumptions compared to prior work. The key ingredient enabling our results is a novel connection to local error bounds from optimization theory.

Note: This paper describes an older version of DeepLIFT. See https://arxiv.org/abs/1704.02685 for the newer version. Original abstract follows: The purported "black box" nature of neural networks is a barrier to adoption in applications where interpretability is essential. Here we present DeepLIFT (Learning Important FeaTures), an efficient and effective method for computing importance scores in a neural network. DeepLIFT compares the activation of each neuron to its 'reference activation' and assigns contribution scores according to the difference. We apply DeepLIFT to models trained on natural images and genomic data, and show significant advantages over gradient-based methods.

Activation functions are fundamental elements of deep learning architectures as they significantly influence training dynamics. ReLU, while widely used, is prone to the dying neuron problem, which has been mitigated by variants such as LeakyReLU, PReLU, and ELU that better handle negative neuron outputs. Recently, self-gated activations like GELU and Swish have emerged as state-of-the-art alternatives, leveraging their smoothness to ensure stable gradient flow and prevent neuron inactivity. In this work, we introduce the Gompertz Linear Unit (GoLU), a novel self-gated activation function defined as GoLU(x) = x , Gompertz(x), where Gompertz(x) = e^{-e^{-x}}. The GoLU activation leverages the asymmetry in the Gompertz function to reduce variance in the latent space more effectively compared to GELU and Swish, while preserving robust gradient flow. Extensive experiments across diverse tasks, including Image Classification, Language Modeling, Semantic Segmentation, Object Detection, Instance Segmentation, and Diffusion, highlight GoLU's superior performance relative to state-of-the-art activation functions, establishing GoLU as a robust alternative to existing activation functions.

Large vision-language models (LVLMs) have rapidly advanced across various domains, yet they still lag behind strong text-only large language models (LLMs) on tasks that require multi-step inference and compositional decision-making. Motivated by their shared transformer architectures, we investigate whether the two model families rely on common internal computation for such inference. At the neuron level, we uncover a surprisingly large overlap: more than half of the top-activated units during multi-step inference are shared between representative LLMs and LVLMs, revealing a modality-invariant inference subspace. Through causal probing via activation amplification, we further show that these shared neurons encode consistent and interpretable concept-level effects, demonstrating their functional contribution to inference. Building on this insight, we propose Shared Neuron Low-Rank Fusion (SNRF), a parameter-efficient framework that transfers mature inference circuitry from LLMs to LVLMs. SNRF profiles cross-model activations to identify shared neurons, computes a low-rank approximation of inter-model weight differences, and injects these updates selectively within the shared-neuron subspace. This mechanism strengthens multimodal inference performance with minimal parameter changes and requires no large-scale multimodal fine-tuning. Across diverse mathematics and perception benchmarks, SNRF consistently enhances LVLM inference performance while preserving perceptual capabilities. Our results demonstrate that shared neurons form an interpretable bridge between LLMs and LVLMs, enabling low-cost transfer of inference ability into multimodal models. Our code is available at [https://github.com/chenhangcuisg-code/Do-LLMs-VLMs-Share-Neurons](https://github.com/chenhangcuisg-code/Do-LLMs-VLMs-Share-Neurons).

In this study, we reveal an in-context learning (ICL) capability of multilingual large language models (LLMs): by translating the input to several languages, we provide Parallel Input in Multiple Languages (PiM) to LLMs, which significantly enhances their comprehension abilities. To test this capability, we design extensive experiments encompassing 8 typical datasets, 7 languages and 8 state-of-the-art multilingual LLMs. Experimental results show that (1) incorporating more languages help PiM surpass the conventional ICL further; (2) even combining with the translations that are inferior to baseline performance can also help. Moreover, by examining the activated neurons in LLMs, we discover a counterintuitive but interesting phenomenon. Contrary to the common thought that PiM would activate more neurons than monolingual input to leverage knowledge learned from diverse languages, PiM actually inhibits neurons and promotes more precise neuron activation especially when more languages are added. This phenomenon aligns with the neuroscience insight about synaptic pruning, which removes less used neural connections, strengthens remainders, and then enhances brain intelligence.

Individual neurons participate in the representation of multiple high-level concepts. To what extent can different interpretability methods successfully disentangle these roles? To help address this question, we introduce RAVEL (Resolving Attribute-Value Entanglements in Language Models), a dataset that enables tightly controlled, quantitative comparisons between a variety of existing interpretability methods. We use the resulting conceptual framework to define the new method of Multi-task Distributed Alignment Search (MDAS), which allows us to find distributed representations satisfying multiple causal criteria. With Llama2-7B as the target language model, MDAS achieves state-of-the-art results on RAVEL, demonstrating the importance of going beyond neuron-level analyses to identify features distributed across activations. We release our benchmark at https://github.com/explanare/ravel.

Large language models (LLMs) achieve strong capabilities by scaling model capacity and training data, yet many real-world deployments rely on smaller models trained or adapted from low-resource data. This gap motivates the need for mechanisms to transfer knowledge from large, high-resource models to smaller, low-resource targets. While model merging provides an effective transfer mechanism, most existing approaches assume architecture-compatible models and therefore cannot directly transfer knowledge from large high-resource LLMs to heterogeneous low-resource targets. In this work, we propose a cross-architecture merging framework based on optimal transport (OT) that aligns activations to infer cross-neuron correspondences between heterogeneous models. The resulting transport plans are then used to guide direct weight-space fusion, enabling effective high-resource to low-resource transfer using only a small set of inputs. Extensive experiments across low-resource languages and specialized domains demonstrate consistent improvements over target models.

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 growing computational demands of large language models (LLMs) make efficient inference and activation strategies increasingly critical. While recent approaches, such as Mixture-of-Experts (MoE), leverage selective activation but require specialized training, training-free sparse activation methods offer broader applicability and superior resource efficiency through their plug-and-play design. However, many existing methods rely solely on hidden state magnitudes to determine activation, resulting in high approximation errors and suboptimal inference accuracy. To address these limitations, we propose WINA (Weight Informed Neuron Activation), a novel, simple, and training-free sparse activation framework that jointly considers hidden state magnitudes and the column-wise ell_2-norms of weight matrices. We show that this leads to a sparsification strategy that obtains optimal approximation error bounds with theoretical guarantees tighter than existing techniques. Empirically, WINA also outperforms state-of-the-art methods (e.g., TEAL) by up to 2.94% in average performance at the same sparsity levels, across a diverse set of LLM architectures and datasets. These results position WINA as a new performance frontier for training-free sparse activation in LLM inference, advancing training-free sparse activation methods and setting a robust baseline for efficient inference. The source code is available at https://github.com/microsoft/wina.

In recent years, text-to-image (T2I) diffusion models have garnered significant attention for their ability to generate high-quality images reflecting text prompts. However, their growing popularity has also led to the emergence of backdoor threats, posing substantial risks. Currently, effective defense strategies against such threats are lacking due to the diversity of backdoor targets in T2I synthesis. In this paper, we propose NaviDet, the first general input-level backdoor detection framework for identifying backdoor inputs across various backdoor targets. Our approach is based on the new observation that trigger tokens tend to induce significant neuron activation variation in the early stage of the diffusion generation process, a phenomenon we term Early-step Activation Variation. Leveraging this insight, NaviDet detects malicious samples by analyzing neuron activation variations caused by input tokens. Through extensive experiments, we demonstrate the effectiveness and efficiency of our method against various T2I backdoor attacks, surpassing existing baselines with significantly lower computational overhead. Furthermore, we rigorously demonstrate that our method remains effective against potential adaptive attacks.

Deep neural networks (DNNs) have demonstrated state-of-the-art results on many pattern recognition tasks, especially vision classification problems. Understanding the inner workings of such computational brains is both fascinating basic science that is interesting in its own right - similar to why we study the human brain - and will enable researchers to further improve DNNs. One path to understanding how a neural network functions internally is to study what each of its neurons has learned to detect. One such method is called activation maximization (AM), which synthesizes an input (e.g. an image) that highly activates a neuron. Here we dramatically improve the qualitative state of the art of activation maximization by harnessing a powerful, learned prior: a deep generator network (DGN). The algorithm (1) generates qualitatively state-of-the-art synthetic images that look almost real, (2) reveals the features learned by each neuron in an interpretable way, (3) generalizes well to new datasets and somewhat well to different network architectures without requiring the prior to be relearned, and (4) can be considered as a high-quality generative method (in this case, by generating novel, creative, interesting, recognizable images).

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.

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.

Protein language models (PLMs) encode rich biological information, yet their internal neuron representations are poorly understood. We introduce the first automated framework for labeling every neuron in a PLM with biologically grounded natural language descriptions. Unlike prior approaches relying on sparse autoencoders or manual annotation, our method scales to hundreds of thousands of neurons, revealing individual neurons are selectively sensitive to diverse biochemical and structural properties. We then develop a novel neuron activation-guided steering method to generate proteins with desired traits, enabling convergence to target biochemical properties like molecular weight and instability index as well as secondary and tertiary structural motifs, including alpha helices and canonical Zinc Fingers. We finally show that analysis of labeled neurons in different model sizes reveals PLM scaling laws and a structured neuron space distribution.

Sparse computation offers a compelling solution for the inference of Large Language Models (LLMs) in low-resource scenarios by dynamically skipping the computation of inactive neurons. While traditional approaches focus on ReLU-based LLMs, leveraging zeros in activation values, we broaden the scope of sparse LLMs beyond zero activation values. We introduce a general method that defines neuron activation through neuron output magnitudes and a tailored magnitude threshold, demonstrating that non-ReLU LLMs also exhibit sparse activation. To find the most efficient activation function for sparse computation, we propose a systematic framework to examine the sparsity of LLMs from three aspects: the trade-off between sparsity and performance, the predictivity of sparsity, and the hardware affinity. We conduct thorough experiments on LLMs utilizing different activation functions, including ReLU, SwiGLU, ReGLU, and ReLU^2. The results indicate that models employing ReLU^2 excel across all three evaluation aspects, highlighting its potential as an efficient activation function for sparse LLMs. We will release the code to facilitate future research.

The choice of activation function plays a critical role in neural networks, yet most architectures still rely on fixed, uniform activation functions across all neurons. We introduce SmartMixed, a two-phase training strategy that allows networks to learn optimal per-neuron activation functions while preserving computational efficiency at inference. In the first phase, neurons adaptively select from a pool of candidate activation functions (ReLU, Sigmoid, Tanh, Leaky ReLU, ELU, SELU) using a differentiable hard-mixture mechanism. In the second phase, each neuron's activation function is fixed according to the learned selection, resulting in a computationally efficient network that supports continued training with optimized vectorized operations. We evaluate SmartMixed on the MNIST dataset using feedforward neural networks of varying depths. The analysis shows that neurons in different layers exhibit distinct preferences for activation functions, providing insights into the functional diversity within neural architectures.

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.

This paper introduces PowerInfer, a high-speed Large Language Model (LLM) inference engine on a personal computer (PC) equipped with a single consumer-grade GPU. The key underlying the design of PowerInfer is exploiting the high locality inherent in LLM inference, characterized by a power-law distribution in neuron activation. This distribution indicates that a small subset of neurons, termed hot neurons, are consistently activated across inputs, while the majority, cold neurons, vary based on specific inputs. PowerInfer exploits such an insight to design a GPU-CPU hybrid inference engine: hot-activated neurons are preloaded onto the GPU for fast access, while cold-activated neurons are computed on the CPU, thus significantly reducing GPU memory demands and CPU-GPU data transfers. PowerInfer further integrates adaptive predictors and neuron-aware sparse operators, optimizing the efficiency of neuron activation and computational sparsity. Evaluation shows that PowerInfer attains an average token generation rate of 13.20 tokens/s, with a peak of 29.08 tokens/s, across various LLMs (including OPT-175B) on a single NVIDIA RTX 4090 GPU, only 18% lower than that achieved by a top-tier server-grade A100 GPU. This significantly outperforms llama.cpp by up to 11.69x while retaining model accuracy.

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.

Neural networks have proven to be a highly effective tool for solving complex problems in many areas of life. Recently, their importance and practical usability have further been reinforced with the advent of deep learning. One of the important conditions for the success of neural networks is the choice of an appropriate activation function introducing non-linearity into the model. Many types of these functions have been proposed in the literature in the past, but there is no single comprehensive source containing their exhaustive overview. The absence of this overview, even in our experience, leads to redundancy and the unintentional rediscovery of already existing activation functions. To bridge this gap, our paper presents an extensive survey involving 400 activation functions, which is several times larger in scale than previous surveys. Our comprehensive compilation also references these surveys; however, its main goal is to provide the most comprehensive overview and systematization of previously published activation functions with links to their original sources. The secondary aim is to update the current understanding of this family of functions.

We introduce a robust, error-tolerant adaptive training algorithm for generalized learning paradigms in high-dimensional superposed quantum networks, or adaptive quantum networks. The formalized procedure applies standard backpropagation training across a coherent ensemble of discrete topological configurations of individual neural networks, each of which is formally merged into appropriate linear superposition within a predefined, decoherence-free subspace. Quantum parallelism facilitates simultaneous training and revision of the system within this coherent state space, resulting in accelerated convergence to a stable network attractor under consequent iteration of the implemented backpropagation algorithm. Parallel evolution of linear superposed networks incorporating backpropagation training provides quantitative, numerical indications for optimization of both single-neuron activation functions and optimal reconfiguration of whole-network quantum structure.

In recent years, generative adversarial networks (GANs) and its variants have achieved unprecedented success in image synthesis. They are widely adopted in synthesizing facial images which brings potential security concerns to humans as the fakes spread and fuel the misinformation. However, robust detectors of these AI-synthesized fake faces are still in their infancy and are not ready to fully tackle this emerging challenge. In this work, we propose a novel approach, named FakeSpotter, based on monitoring neuron behaviors to spot AI-synthesized fake faces. The studies on neuron coverage and interactions have successfully shown that they can be served as testing criteria for deep learning systems, especially under the settings of being exposed to adversarial attacks. Here, we conjecture that monitoring neuron behavior can also serve as an asset in detecting fake faces since layer-by-layer neuron activation patterns may capture more subtle features that are important for the fake detector. Experimental results on detecting four types of fake faces synthesized with the state-of-the-art GANs and evading four perturbation attacks show the effectiveness and robustness of our approach.

In recent years, neural networks have become the default choice for image classification and many other learning tasks, even though they are vulnerable to so-called adversarial attacks. To increase their robustness against these attacks, there have emerged numerous detection mechanisms that aim to automatically determine if an input is adversarial. However, state-of-the-art detection mechanisms either rely on being tuned for each type of attack, or they do not generalize across different attack types. To alleviate these issues, we propose a novel technique for adversarial-image detection, RAID, that trains a secondary classifier to identify differences in neuron activation values between benign and adversarial inputs. Our technique is both more reliable and more effective than the state of the art when evaluated against six popular attacks. Moreover, a straightforward extension of RAID increases its robustness against detection-aware adversaries without affecting its effectiveness.

Large language models (LLMs) demonstrate remarkable performance across a spectrum of languages. In this work, we delve into the question: How do LLMs handle multilingualism? We introduce a framework that depicts LLMs' processing of multilingual inputs: In the first several layers, LLMs understand the question, converting multilingual inputs into English to facilitate the task-solving phase. In the intermediate layers, LLMs engage in problem-solving by thinking in English and incorporating multilingual knowledge to obtain factual content, leveraging the self-attention and feed-forward structures, respectively. In the last several layers, LLMs generate responses that align with the original language of the query. In addition, we investigate the existence of language-specific neurons when processing a certain language. To detect neurons activated by the input language, even without labels, we innovatively design a Parallel Language specific Neuron Detection (PLND) method that effectively measures the significance of neurons when handling multilingual inputs. By comprehensive ablation analysis through deactivating neurons of different layers and structures, we verify the framework that we propose. Additionally, we demonstrate that we can utilize such a framework to effectively enhance the multilingual ability with much less training effort.

Sequence-to-sequence (seq2seq) learning is a popular fashion for large-scale pretraining language models. However, the prior seq2seq pretraining models generally focus on reconstructive objectives on the decoder side and neglect the effect of encoder-side supervision, which we argue may lead to sub-optimal performance. To verify our hypothesis, we first empirically study the functionalities of the encoder and decoder in seq2seq pretrained language models, and find that the encoder takes an important but under-exploitation role than the decoder regarding the downstream performance and neuron activation. Therefore, we propose an encoding-enhanced seq2seq pretraining strategy, namely E2S2, which improves the seq2seq models via integrating more efficient self-supervised information into the encoders. Specifically, E2S2 adopts two self-supervised objectives on the encoder side from two aspects: 1) locally denoising the corrupted sentence (denoising objective); and 2) globally learning better sentence representations (contrastive objective). With the help of both objectives, the encoder can effectively distinguish the noise tokens and capture high-level (i.e. syntactic and semantic) knowledge, thus strengthening the ability of seq2seq model to accurately achieve the conditional generation. On a large diversity of downstream natural language understanding and generation tasks, E2S2 dominantly improves the performance of its powerful backbone models, e.g. BART and T5. For example, upon BART backbone, we achieve +1.1% averaged gain on the general language understanding evaluation (GLUE) benchmark and +1.75% F_0.5 score improvement on CoNLL2014 dataset. We also provide in-depth analyses to show the improvement stems from better linguistic representation. We hope that our work will foster future self-supervision research on seq2seq language model pretraining.

While deep learning models have achieved state-of-the-art accuracies for many prediction tasks, understanding these models remains a challenge. Despite the recent interest in developing visual tools to help users interpret deep learning models, the complexity and wide variety of models deployed in industry, and the large-scale datasets that they used, pose unique design challenges that are inadequately addressed by existing work. Through participatory design sessions with over 15 researchers and engineers at Facebook, we have developed, deployed, and iteratively improved ActiVis, an interactive visualization system for interpreting large-scale deep learning models and results. By tightly integrating multiple coordinated views, such as a computation graph overview of the model architecture, and a neuron activation view for pattern discovery and comparison, users can explore complex deep neural network models at both the instance- and subset-level. ActiVis has been deployed on Facebook's machine learning platform. We present case studies with Facebook researchers and engineers, and usage scenarios of how ActiVis may work with different models.

The outbreak of COVID-19 has shocked the entire world with its fairly rapid spread and has challenged different sectors. One of the most effective ways to limit its spread is the early and accurate diagnosing infected patients. Medical imaging, such as X-ray and Computed Tomography (CT), combined with the potential of Artificial Intelligence (AI), plays an essential role in supporting medical personnel in the diagnosis process. Thus, in this article five different deep learning models (ResNet18, ResNet34, InceptionV3, InceptionResNetV2 and DenseNet161) and their ensemble, using majority voting have been used to classify COVID-19, pneumoni{\ae} and healthy subjects using chest X-ray images. Multilabel classification was performed to predict multiple pathologies for each patient, if present. Firstly, the interpretability of each of the networks was thoroughly studied using local interpretability methods - occlusion, saliency, input X gradient, guided backpropagation, integrated gradients, and DeepLIFT, and using a global technique - neuron activation profiles. The mean Micro-F1 score of the models for COVID-19 classifications ranges from 0.66 to 0.875, and is 0.89 for the ensemble of the network models. The qualitative results showed that the ResNets were the most interpretable models. This research demonstrates the importance of using interpretability methods to compare different models before making a decision regarding the best performing model.

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.

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.

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.

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.

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.

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

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.

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.

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.

In-context learning is a powerful emergent ability in transformer models. Prior work in mechanistic interpretability has identified a circuit element that may be critical for in-context learning -- the induction head (IH), which performs a match-and-copy operation. During training of large transformers on natural language data, IHs emerge around the same time as a notable phase change in the loss. Despite the robust evidence for IHs and this interesting coincidence with the phase change, relatively little is known about the diversity and emergence dynamics of IHs. Why is there more than one IH, and how are they dependent on each other? Why do IHs appear all of a sudden, and what are the subcircuits that enable them to emerge? We answer these questions by studying IH emergence dynamics in a controlled setting by training on synthetic data. In doing so, we develop and share a novel optogenetics-inspired causal framework for modifying activations throughout training. Using this framework, we delineate the diverse and additive nature of IHs. By clamping subsets of activations throughout training, we then identify three underlying subcircuits that interact to drive IH formation, yielding the phase change. Furthermore, these subcircuits shed light on data-dependent properties of formation, such as phase change timing, already showing the promise of this more in-depth understanding of subcircuits that need to "go right" for an induction head.

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.

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.

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.

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.

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.

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.

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.

In this work we identify the dormant neuron phenomenon in deep reinforcement learning, where an agent's network suffers from an increasing number of inactive neurons, thereby affecting network expressivity. We demonstrate the presence of this phenomenon across a variety of algorithms and environments, and highlight its effect on learning. To address this issue, we propose a simple and effective method (ReDo) that Recycles Dormant neurons throughout training. Our experiments demonstrate that ReDo maintains the expressive power of networks by reducing the number of dormant neurons and results in improved performance.

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.

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.

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.

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.

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}.

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.

Interpretability researchers have attempted to understand MLP neurons of language models based on both the contexts in which they activate and their output weight vectors. They have paid little attention to a complementary aspect: the interactions between input and output. For example, when neurons detect a direction in the input, they might add much the same direction to the residual stream ("enrichment neurons") or reduce its presence ("depletion neurons"). We address this aspect by examining the cosine similarity between input and output weights of a neuron. We apply our method to 12 models and find that enrichment neurons dominate in early-middle layers whereas later layers tend more towards depletion. To explain this finding, we argue that enrichment neurons are largely responsible for enriching concept representations, one of the first steps of factual recall. Our input-output perspective is a complement to activation-dependent analyses and to approaches that treat input and output separately.

The truthfulness of existing explanation methods in authentically elucidating the underlying model's decision-making process has been questioned. Existing methods have deviated from faithfully representing the model, thus susceptible to adversarial attacks. To address this, we propose a novel eXplainable AI (XAI) method called SRD (Sharing Ratio Decomposition), which sincerely reflects the model's inference process, resulting in significantly enhanced robustness in our explanations. Different from the conventional emphasis on the neuronal level, we adopt a vector perspective to consider the intricate nonlinear interactions between filters. We also introduce an interesting observation termed Activation-Pattern-Only Prediction (APOP), letting us emphasize the importance of inactive neurons and redefine relevance encapsulating all relevant information including both active and inactive neurons. Our method, SRD, allows for the recursive decomposition of a Pointwise Feature Vector (PFV), providing a high-resolution Effective Receptive Field (ERF) at any layer.

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.

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.

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.

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.

Parameter-efficient fine-tuning (PEFT) methods such as adapt large pretrained models by adding small weight-space updates. While effective, weight deltas are hard to interpret mechanistically, and they do not directly expose which internal computations are reused versus bypassed for a new task. We explore an alternative view inspired by neuromodulation: adaptation as a change in mode -- selecting and rescaling existing computations -- rather than rewriting the underlying weights. We propose , a simple activation-space PEFT technique that freezes base weights and learns per-neuron thresholds and gains. During training, a smooth gate decides whether a neuron's activation participates; at inference the gate can be hardened to yield explicit conditional computation and neuron-level attributions. As a proof of concept, we study ``mode specialization'' on MNIST (0^circ) versus rotated MNIST (45^circ). We pretrain a small MLP on a 50/50 mixture (foundation), freeze its weights, and then specialize to the rotated mode using . Across seeds, improves rotated accuracy over the frozen baseline while using only a few hundred trainable parameters per layer, and exhibits partial activation sparsity (a minority of units strongly active). Compared to , trades some accuracy for substantially fewer trainable parameters and a more interpretable ``which-neurons-fire'' mechanism. We discuss limitations, including reduced expressivity when the frozen base lacks features needed for the target mode.

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.

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.

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.

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.

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.

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.

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.

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 NEURONA, a neuro-symbolic framework for fMRI decoding and concept grounding in neural activity. Leveraging image- and video-based fMRI question-answering datasets, NEURONA learns to decode interacting concepts from visual stimuli based on patterns of fMRI responses, integrating symbolic reasoning and compositional execution with fMRI grounding across brain regions. We demonstrate that incorporating structural priors (e.g., compositional predicate-argument dependencies between concepts) into the decoding process significantly improves both decoding accuracy over precise queries, and notably, generalization to unseen queries at test time. With NEURONA, we highlight neuro-symbolic frameworks as promising tools for understanding neural activity.

In large language models (LLMs), certain neurons can store distinct pieces of knowledge learned during pretraining. While knowledge typically appears as a combination of relations and entities, it remains unclear whether some neurons focus on a relation itself -- independent of any entity. We hypothesize such neurons detect a relation in the input text and guide generation involving such a relation. To investigate this, we study the Llama-2 family on a chosen set of relations with a statistics-based method. Our experiments demonstrate the existence of relation-specific neurons. We measure the effect of selectively deactivating candidate neurons specific to relation r on the LLM's ability to handle (1) facts whose relation is r and (2) facts whose relation is a different relation r' neq r. With respect to their capacity for encoding relation information, we give evidence for the following three properties of relation-specific neurons. (i) Neuron cumulativity. The neurons for r present a cumulative effect so that deactivating a larger portion of them results in the degradation of more facts in r. (ii) Neuron versatility. Neurons can be shared across multiple closely related as well as less related relations. Some relation neurons transfer across languages. (iii) Neuron interference. Deactivating neurons specific to one relation can improve LLM generation performance for facts of other relations. We will make our code publicly available at https://github.com/cisnlp/relation-specific-neurons.

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.

In this paper, we identify a sparse reward subsystem within the hidden states of Large Language Models (LLMs), drawing an analogy to the biological reward subsystem in the human brain. We demonstrate that this subsystem contains value neurons that represent the model's internal expectation of state value, and through intervention experiments, we establish the importance of these neurons for reasoning. Our experiments reveal that these value neurons are robust across diverse datasets, model scales, and architectures; furthermore, they exhibit significant transferability across different datasets and models fine-tuned from the same base model. By examining cases where value predictions and actual rewards diverge, we identify dopamine neurons within the reward subsystem which encode reward prediction errors (RPE). These neurons exhibit high activation when the reward is higher than expected and low activation when the reward is lower than expected.

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.

Exposing meaningful and interpretable neural interactions is critical to understanding neural circuits. Inferred neural interactions from neural signals primarily reflect functional interactions. In a long experiment, subject animals may experience different stages defined by the experiment, stimuli, or behavioral states, and hence functional interactions can change over time. To model dynamically changing functional interactions, prior work employs state-switching generalized linear models with hidden Markov models (i.e., HMM-GLMs). However, we argue they lack biological plausibility, as functional interactions are shaped and confined by the underlying anatomical connectome. Here, we propose a novel prior-informed state-switching GLM. We introduce both a Gaussian prior and a one-hot prior over the GLM in each state. The priors are learnable. We will show that the learned prior should capture the state-constant interaction, shedding light on the underlying anatomical connectome and revealing more likely physical neuron interactions. The state-dependent interaction modeled by each GLM offers traceability to capture functional variations across multiple brain states. Our methods effectively recover true interaction structures in simulated data, achieve the highest predictive likelihood with real neural datasets, and render interaction structures and hidden states more interpretable when applied to real neural data.

Deep Learning has revolutionized various fields, including Computer Vision, Natural Language Processing, as well as Biomedical research. Within the field of neuroscience, specifically in electrophysiological neuroimaging, researchers are starting to explore leveraging deep learning to make predictions on their data without extensive feature engineering. The availability of large-scale datasets is a crucial aspect of allowing the experimentation of Deep Learning models. We are publishing the first large-scale clinical EEG dataset that simplifies data access and management for Deep Learning. This dataset contains eyes-closed EEG data prepared from a collection of 1,574 juvenile participants from the Healthy Brain Network. We demonstrate a use case integrating this framework, and discuss why providing such neuroinformatics infrastructure to the community is critical for future scientific discoveries.

Uncovering the fundamental neural correlates of biological intelligence, developing mathematical models, and conducting computational simulations are critical for advancing new paradigms in artificial intelligence (AI). In this study, we implemented a comprehensive visual decision-making model that spans from visual input to behavioral output, using a neural dynamics modeling approach. Drawing inspiration from the key components of the dorsal visual pathway in primates, our model not only aligns closely with human behavior but also reflects neural activities in primates, and achieving accuracy comparable to convolutional neural networks (CNNs). Moreover, magnetic resonance imaging (MRI) identified key neuroimaging features such as structural connections and functional connectivity that are associated with performance in perceptual decision-making tasks. A neuroimaging-informed fine-tuning approach was introduced and applied to the model, leading to performance improvements that paralleled the behavioral variations observed among subjects. Compared to classical deep learning models, our model more accurately replicates the behavioral performance of biological intelligence, relying on the structural characteristics of biological neural networks rather than extensive training data, and demonstrating enhanced resilience to perturbation.

We interpret the function of individual neurons in CLIP by automatically describing them using text. Analyzing the direct effects (i.e. the flow from a neuron through the residual stream to the output) or the indirect effects (overall contribution) fails to capture the neurons' function in CLIP. Therefore, we present the "second-order lens", analyzing the effect flowing from a neuron through the later attention heads, directly to the output. We find that these effects are highly selective: for each neuron, the effect is significant for <2% of the images. Moreover, each effect can be approximated by a single direction in the text-image space of CLIP. We describe neurons by decomposing these directions into sparse sets of text representations. The sets reveal polysemantic behavior - each neuron corresponds to multiple, often unrelated, concepts (e.g. ships and cars). Exploiting this neuron polysemy, we mass-produce "semantic" adversarial examples by generating images with concepts spuriously correlated to the incorrect class. Additionally, we use the second-order effects for zero-shot segmentation and attribute discovery in images. Our results indicate that a scalable understanding of neurons can be used for model deception and for introducing new model capabilities.

Why do brains have inhibitory connections? Why do deep networks have negative weights? We propose an answer from the perspective of representation capacity. We believe representing functions is the primary role of both (i) the brain in natural intelligence, and (ii) deep networks in artificial intelligence. Our answer to why there are inhibitory/negative weights is: to learn more functions. We prove that, in the absence of negative weights, neural networks with non-decreasing activation functions are not universal approximators. While this may be an intuitive result to some, to the best of our knowledge, there is no formal theory, in either machine learning or neuroscience, that demonstrates why negative weights are crucial in the context of representation capacity. Further, we provide insights on the geometric properties of the representation space that non-negative deep networks cannot represent. We expect these insights will yield a deeper understanding of more sophisticated inductive priors imposed on the distribution of weights that lead to more efficient biological and machine learning.

Activation sparsity improves compute efficiency and resource utilization in sparsity-aware neural network accelerators. As the predominant operation in DNNs is multiply-accumulate (MAC) of activations with weights to compute inner products, skipping operations where (at least) one of the two operands is zero can make inference more efficient in terms of latency and power. Spatial sparsification of activations is a popular topic in DNN literature and several methods have already been established to bias a DNN for it. On the other hand, temporal sparsity is an inherent feature of bio-inspired spiking neural networks (SNNs), which neuromorphic processing exploits for hardware efficiency. Introducing and exploiting spatio-temporal sparsity, is a topic much less explored in DNN literature, but in perfect resonance with the trend in DNN, to shift from static signal processing to more streaming signal processing. Towards this goal, in this paper we introduce a new DNN layer (called Delta Activation Layer), whose sole purpose is to promote temporal sparsity of activations during training. A Delta Activation Layer casts temporal sparsity into spatial activation sparsity to be exploited when performing sparse tensor multiplications in hardware. By employing delta inference and ``the usual'' spatial sparsification heuristics during training, the resulting model learns to exploit not only spatial but also temporal activation sparsity (for a given input data distribution). One may use the Delta Activation Layer either during vanilla training or during a refinement phase. We have implemented Delta Activation Layer as an extension of the standard Tensoflow-Keras library, and applied it to train deep neural networks on the Human Action Recognition (UCF101) dataset. We report an almost 3x improvement of activation sparsity, with recoverable loss of model accuracy after longer training.

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.

The training process of neural networks usually optimize weights and bias parameters of linear transformations, while nonlinear activation functions are pre-specified and fixed. This work develops a systematic approach to constructing matrix activation functions whose entries are generalized from ReLU. The activation is based on matrix-vector multiplications using only scalar multiplications and comparisons. The proposed activation functions depend on parameters that are trained along with the weights and bias vectors. Neural networks based on this approach are simple and efficient and are shown to be robust in numerical experiments.

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.

Biological nervous systems are created in a fundamentally different way than current artificial neural networks. Despite its impressive results in a variety of different domains, deep learning often requires considerable engineering effort to design high-performing neural architectures. By contrast, biological nervous systems are grown through a dynamic self-organizing process. In this paper, we take initial steps toward neural networks that grow through a developmental process that mirrors key properties of embryonic development in biological organisms. The growth process is guided by another neural network, which we call a Neural Developmental Program (NDP) and which operates through local communication alone. We investigate the role of neural growth on different machine learning benchmarks and different optimization methods (evolutionary training, online RL, offline RL, and supervised learning). Additionally, we highlight future research directions and opportunities enabled by having self-organization driving the growth of neural networks.

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.

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.

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.

Quantifying similarity between neural representations -- e.g. hidden layer activation vectors -- is a perennial problem in deep learning and neuroscience research. Existing methods compare deterministic responses (e.g. artificial networks that lack stochastic layers) or averaged responses (e.g., trial-averaged firing rates in biological data). However, these measures of _deterministic_ representational similarity ignore the scale and geometric structure of noise, both of which play important roles in neural computation. To rectify this, we generalize previously proposed shape metrics (Williams et al. 2021) to quantify differences in _stochastic_ representations. These new distances satisfy the triangle inequality, and thus can be used as a rigorous basis for many supervised and unsupervised analyses. Leveraging this novel framework, we find that the stochastic geometries of neurobiological representations of oriented visual gratings and naturalistic scenes respectively resemble untrained and trained deep network representations. Further, we are able to more accurately predict certain network attributes (e.g. training hyperparameters) from its position in stochastic (versus deterministic) shape space.

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.

Massive activations are scalar values in transformer hidden states that achieve values orders of magnitude larger than typical activations and have been shown to be critical for model functionality. While prior work has characterized these phenomena in fully trained models, the temporal dynamics of their emergence during training remain poorly understood. We present the first comprehensive analysis of massive activation development throughout transformer training, using the Pythia model family as our testbed. Through systematic analysis of various model sizes across multiple training checkpoints, we demonstrate that massive activation emergence follows predictable mathematical patterns that can be accurately modeled using an exponentially-modulated logarithmic function with five key parameters. We develop a machine learning framework to predict these mathematical parameters from architectural specifications alone, achieving high accuracy for steady-state behavior and moderate accuracy for emergence timing and magnitude. These findings enable architects to predict and potentially control key aspects of massive activation emergence through design choices, with significant implications for model stability, training cycle length, interpretability, and optimization. Our findings demonstrate that the emergence of massive activations is governed by model design and can be anticipated, and potentially controlled, before training begins.

Large Reasoning Models (LRMs) often suffer from overthinking, a phenomenon in which redundant reasoning steps are generated after a correct solution has already been reached. Existing early reasoning exit methods primarily rely on output-level heuristics or trained probing models to skip redundant reasoning steps, thereby mitigating overthinking. However, these approaches typically require additional rollout computation or externally labeled datasets. In this paper, we propose NEAT, a Neuron-based Early reAsoning exiT framework that monitors neuron-level activation dynamics to enable training-free early exits, without introducing additional test-time computation. NEAT identifies exit-associated neurons and tracks their activation patterns during reasoning to dynamically trigger early exit or suppress reflection, thereby reducing unnecessary reasoning while preserving solution quality. Experiments on four reasoning benchmarks across six models with different scales and architectures show that, for each model, NEAT achieves an average token reduction of 22\% to 28\% when averaged over the four benchmarks, while maintaining accuracy.

We propose the APTx Neuron, a novel, unified neural computation unit that integrates non-linear activation and linear transformation into a single trainable expression. The APTx Neuron is derived from the APTx activation function, thereby eliminating the need for separate activation layers and making the architecture both computationally efficient and elegant. The proposed neuron follows the functional form y = sum_{i=1}^{n} ((alpha_i + tanh(beta_i x_i)) cdot gamma_i x_i) + delta, where all parameters alpha_i, beta_i, gamma_i, and delta are trainable. We validate our APTx Neuron-based architecture on the MNIST dataset, achieving up to 96.69% test accuracy within 11 epochs using approximately 332K trainable parameters. The results highlight the superior expressiveness and computational efficiency of the APTx Neuron compared to traditional neurons, pointing toward a new paradigm in unified neuron design and the architectures built upon it.

Deep Neural Networks (DNNs) are extensively employed in safety-critical applications where ensuring hardware reliability is a primary concern. To enhance the reliability of DNNs against hardware faults, activation restriction techniques significantly mitigate the fault effects at the DNN structure level, irrespective of accelerator architectures. State-of-the-art methods offer either neuron-wise or layer-wise clipping activation functions. They attempt to determine optimal clipping thresholds using heuristic and learning-based approaches. Layer-wise clipped activation functions cannot preserve DNNs resilience at high bit error rates. On the other hand, neuron-wise clipping activation functions introduce considerable memory overhead due to the addition of parameters, which increases their vulnerability to faults. Moreover, the heuristic-based optimization approach demands numerous fault injections during the search process, resulting in time-consuming threshold identification. On the other hand, learning-based techniques that train thresholds for entire layers concurrently often yield sub-optimal results. In this work, first, we demonstrate that it is not essential to incorporate neuron-wise activation functions throughout all layers in DNNs. Then, we propose a hybrid clipped activation function that integrates neuron-wise and layer-wise methods that apply neuron-wise clipping only in the last layer of DNNs. Additionally, to attain optimal thresholds in the clipping activation function, we introduce ProAct, a progressive training methodology. This approach iteratively trains the thresholds on a layer-by-layer basis, aiming to obtain optimal threshold values in each layer separately.

Activation functions are core components of all deep learning architectures. Currently, the most popular activation functions are smooth ReLU variants like GELU and SiLU. These are self-gated activation functions where the range of the gating function is between zero and one. In this paper, we explore the viability of using arctan as a gating mechanism. A self-gated activation function that uses arctan as its gating function has a monotonically increasing first derivative. To make this activation function competitive, it is necessary to introduce a trainable parameter for every MLP block to expand the range of the gating function beyond zero and one. We find that this technique also improves existing self-gated activation functions. We conduct an empirical evaluation of Expanded ArcTan Linear Unit (xATLU), Expanded GELU (xGELU), and Expanded SiLU (xSiLU) and show that they outperform existing activation functions within a transformer architecture. Additionally, expanded gating ranges show promising results in improving first-order Gated Linear Units (GLU).

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.

In theoretical neuroscience, recent work leverages deep learning tools to explore how some network attributes critically influence its learning dynamics. Notably, initial weight distributions with small (resp. large) variance may yield a rich (resp. lazy) regime, where significant (resp. minor) changes to network states and representation are observed over the course of learning. However, in biology, neural circuit connectivity could exhibit a low-rank structure and therefore differs markedly from the random initializations generally used for these studies. As such, here we investigate how the structure of the initial weights -- in particular their effective rank -- influences the network learning regime. Through both empirical and theoretical analyses, we discover that high-rank initializations typically yield smaller network changes indicative of lazier learning, a finding we also confirm with experimentally-driven initial connectivity in recurrent neural networks. Conversely, low-rank initialization biases learning towards richer learning. Importantly, however, as an exception to this rule, we find lazier learning can still occur with a low-rank initialization that aligns with task and data statistics. Our research highlights the pivotal role of initial weight structures in shaping learning regimes, with implications for metabolic costs of plasticity and risks of catastrophic forgetting.