Biologically Plausible Learning

Learning and decision-making in animals are often modeled as Bayesian processes, where sensory evidence is integrated with prior beliefs to guide behavior in the face of uncertainty. But what are the inherent neural dynamics that give rise to this ability, and how could they be replicated in computing systems? This abstract discusses a biologically grounded framework in which noisy neural and synaptic dynamics perform inference and learning via stochastic sampling from an internal energy function, capturing uncertainty over latent states and model parameters through neural and synaptic variability, respectively. This enables approaches such as predictive coding networks to account for epistemic uncertainty via Markov chain Monte Carlo sampling. Drawing a parallel between intrinsic noise in biological systems and electrical noise in emerging probabilistic analogue memory technologies, we highlight how analogue in-memory computing hardware naturally emerges as the solution for massively scalable and energy-efficient probabilistic inference.

Locust visual systems have long served as an important biological paradigm for studying looming perception and collision avoidance. Numerous computational models have successfully reproduced the selective responses of Lobula Giant Movement Detector (LGMD) neurons to approaching objects, thereby emulating the fundamental functionality of the biological system. However, existing models remain limited in biological plausibility and robustness when operating in complex and dynamic visual environments. To address these limitations, we propose a biologically plausible neural network for locust-inspired looming detection. The proposed framework incorporates a spatially isotropic sampling strategy that mimics the ommatidial organization of the locust compound eye, a population-voting mechanism inspired by population coding in biological neural systems, and leaky integrate-and-fire neuronal dynamics to replace conventional sigmoid-based membrane activation. Systematic experiments on synthetic stimuli, laboratory sequences, and real-world driving scenarios demonstrate that the proposed model improves robustness under challenging visual conditions while preserving computational efficiency and enhancing biological fidelity. These results highlight the potential of biologically grounded neural computation for robust and efficient collision perception.

Representational similarity analysis (RSA) is increasingly used to ask which learning rules give convolutional networks brain-like representations. Because biologically plausible rules such as feedback alignment, predictive coding and STDP do not scale, studies that include them train small networks on small images (typically 32x32 CIFAR) and then compare them to brain responses modeled at much higher resolution. We find that a common result in this setting, that untrained or locally trained networks rival or beat backpropagation at early visual cortex, depends strongly on the resolution at which the network is evaluated. The V1 gap between an untrained network and a backpropagation-trained one widens from -0.001 +/- 0.007 at the 32px training resolution to +0.044 +/- 0.006 at 224px, growing monotonically across six resolutions (n=5 seeds). It holds in human fMRI and, directionally, in single-seed macaque electrophysiology, along the training trajectory, and for an ImageNet ResNet-50 and a Swin-Tiny transformer trained at 224px. Four candidate mechanisms are tested and none accounts for it: train/eval resolution matching, low-level Gabor and pixel structure, the normalization state of the untrained baseline, and convergence of the pooled descriptor toward a global brightness statistic; three are excluded by interventions holding the convolutional weights bit-identical. A fifth experiment locates the effect: capping image detail at the training resolution while letting the pooled positions grow 12-fold removes about 90% of it, so the dependence is carried by image detail rather than by pooling. Separately, a single scalar luminance value per image reaches rho = 0.075 against the V1 RDM, essentially matching the untrained network's 0.076, which bounds what this style of comparison can resolve. The one learning effect that holds across resolution is backprop above untrained, at LOC.

Biologically plausible learning models aim to explain how neural circuits can implement effective learning under the constraints of real neurons. Although significant progress has been made, a major remaining challenge is that existing models often allow neurons or synapses to represent mixed-sign values, both positive and negative, in violation of a basic aspect of cortical circuitry -- Dale's constraint: biological neurons are either excitatory or inhibitory, but not both, and synapses cannot change sign. In this work, we address this discrepancy by introducing a biologically motivated neural architecture in which both neural activations and learning signals are represented by non-negative activity, and synapses have fixed sign, while still supporting backpropagation-like learning. Our approach uses two complementary interacting non-negative channels to represent positive and negative contributions, inspired by evidence of on-off representations in the brain. These channels are implemented through a simple neural circuit motif, which is repeated throughout the network in both bottom-up and top-down pathways. Combined with a local Hebbian learning rule, the resulting model propagates learning signals and updates weights using only local interactions between neurons. We show theoretically that our learning scheme can exactly recover the backpropagation update despite relying solely on non-negative error signals. Empirically, beyond satisfying stronger biological constraints, the on-off architecture learns efficient representations, yielding substantial gains over comparable vanilla networks on the Tiny ImageNet benchmark. These results demonstrate that effective learning can emerge from biologically plausible mechanisms without requiring mixed-sign signals, providing a step toward more realistic models of neural computation.

The pruning of network connections is key to brain function but, despite its importance, there exist few biologically-plausible pruning rules with demonstrated good performance. In this work we evaluate noise-prune, a recently introduced unsupervised local pruning rule for recurrent networks that uses noisy fluctuations to determine the importance of connections. Noise-prune has previously only been empirically tested on random networks without a specific computational function. We show that noise-prune preserves task-performance in task-trained recurrent neural networks, greatly outperforming a strategy that only uses the magnitude of connections and performing on par with or exceeding a non-local strategy that uses second-order information. Rather than deterministically removing connections that fall below a certain threshold importance, noise-prune samples connections to preserve based on their importance and strengthens retained connections to preserve average synaptic strength. We show that this sampling and rescaling is essential to good performance, but that the optimal empirical degree of rescaling is lower than that predicted by the original theoretical argument. Our work thus validates noise-prune as a biologically-plausible pruning rule for functional recurrent network architectures and characterizes its optimal parameter settings.

Biological neural circuits obey Dale's principle: each neuron's synapses are uniformly excitatory or inhibitory. Artificial networks that respect this constraint must coordinate separate excitatory and inhibitory populations, fundamentally changing how credit is assigned during learning. Several biologically plausible learning rules avoid backpropagation's weight transport requirement, but it has been difficult to achieve strong performance under Dale's principle beyond MNIST. Error Diffusion (ED) was originally proposed in a dual-stream excitatory/inhibitory architecture, where learning is driven by routing global error signals to all layers without transporting transposed forward weights or relying on random feedback matrices. Whether such a rule can scale under Dale's principle across both supervised classification and reinforcement learning remains unknown. Here, we introduce modulo error routing to extend Error Diffusion beyond binary classification, and show that a dual-stream excitatory/inhibitory architecture trained with this method achieves 96.7% on MNIST and establishes a 61.7% baseline on CIFAR-10, demonstrating that representation learning is possible even when strictly enforcing Dale's principle. For the classification setting, we introduce three domain-specific innovations: layer-specific sigmoid widths, batch-centered class error signals, and asymmetric initialization, and ablation analysis reveals that their relative importance reverses between MNIST and CIFAR-10, exposing task-dependent credit-assignment bottlenecks invisible to single-benchmark evaluation. In reinforcement learning, we integrate ED with Proximal Policy Optimization (PPO) and evaluate it on continuous-control tasks in Google Brax and on Craftax, an open-ended exploration task. We show that ED-PPO achieves competitive performance relative to Direct Feedback Alignment, a backpropagation-free baseline.

Learning in biological multilayer neuronal networks offers insights that extend beyond the classical weighted-sum neuron model commonly used in artificial neural networks. This article presents an accessible guide to a mechanistic neuronal network model that more accurately captures aspects of biological computation while enabling a simple yet powerful mechanism for learning in multilayer neural networks. The proposed approach supports efficient online streamed learning and provides a practical alternative to backpropagation. We demonstrate its potential in an image classification task, achieving competitive classification performance. The approach's simplicity, biological grounding, and broad applicability highlight a promising path toward algorithms that unify mechanistic neuron models and machine learning.

We introduce Score Broadcast and Decorrelation (SBD), a principled framework for broadcast-based credit assignment for general families of differentiable losses. Error broadcast is a biologically plausible alternative to backpropagation that sends output information to hidden layers without weight transport. The Error Broadcast and Decorrelation (EBD) framework, recently introduced for the mean-squared-error (MSE) setting, grounded this mechanism in the stochastic orthogonality of optimal estimators, under which the optimal residual is orthogonal to functions of the input. We generalize that foundation by introducing an orthogonality principle between the output score (the gradient of loss with respect to the final-layer output) and hidden-layer activations, which holds whenever the optimal score has conditional mean zero. This single principle unifies broadcast-based credit assignment across the standard differentiable-loss families, including cross-entropy, Bregman divergences, proper scoring rules, and exponential-family negative log-likelihoods. The framework supplies a theoretical grounding for the three-factor learning rule under general losses, with the neuromodulatory factor derived as the broadcast loss score. We derive the cross-entropy case explicitly, characterize the admissible loss class, and introduce a score vector expansion technique that enriches the broadcast signal while preserving the orthogonality framework. Experiments on CIFAR-10 and Tiny ImageNet show that SBD substantially improves over existing broadcast approaches, with score vector expansion delivering further gains. Overall, this work identifies the loss score as the signal to broadcast, supplies the orthogonality theory and theoretical grounding for the three-factor learning rule from neuroscience, and shows how score vector expansion enriches the decorrelation directions of the resulting objective.

Random, untrained neural networks consistently match or exceed trained networks in representational similarity to early visual cortex. This puzzling finding challenges the assumption that learning improves brain alignment. We investigate it by tracking representational similarity analysis (RSA) alignment to human fMRI data across training for four learning rules: backpropagation (BP), feedback alignment (FA), predictive coding (PC), and spike-timing-dependent plasticity (STDP). Using 720 object images from the THINGS database and fMRI data from three subjects across six visual ROIs, we measure Spearman correlations between model and brain representational dissimilarity matrices at eight training checkpoints (epochs 0-40). We find that (1) a single epoch of training reduces V1 alignment by 25-90%, depending on the learning rule; (2) backpropagation reduces V1 alignment most severely (delta r = -0.080), while predictive coding and STDP preserve substantially more (delta r ~ -0.04); and (3) a weaker, opposite tendency appears in object-selective cortex (LOC), where BP shows the largest increase in alignment during training, although the absolute change is small. These results suggest that untrained architectures capture low-level visual statistics through inductive biases alone, and that global error signals (BP) reshape early representations more aggressively than local learning rules (PC, STDP), which better preserve brain-like structure.

The brain learns abstract representations of high-dimensional sensory input, but the plasticity rules that enable such learning are unknown. We study biologically plausible algorithms on the Random Hierarchy Model (RHM), an artificial dataset designed to investigate how deep neural networks learn the intrinsic hierarchical structure of high-dimensional data. We focus on two types of local learning rules that avoid both a long convergence time and the use of a symmetric error network. The first type uses direct feedback signals to approximate error propagation from the output layer. The second type uses layerwise self-supervised contrastive or non-contrastive loss functions that do not explicitly approximate errors at the output layer. We show that all rules of the first type fail to solve the tasks of the RHM and trace this failure back to input-specific nonlinearities (`masking') that are implemented in full backpropagation and are essential for learning complex tasks. However, algorithms of the second type are able to learn the hierarchical hidden structure of the RHM tasks and are as data-efficient as supervised backpropagation training, while being compatible with known rules of synaptic plasticity in cortex.

The feedback alignment (FA) algorithm offers a biologically plausible alternative to backpropagation (BP) for training neural networks yet notably fails to scale to convolutional architectures. Modifications have been proposed to address this limitation, but at questionable cost to biological plausibility. In this paper, we evaluate five learning algorithms including modified FA and standard BP, applied to the same convolutional architecture with the CIFAR-10 dataset. We provide a tripartite comparative analysis focusing on biological plausibility, interpretability, and computational complexity. Our results indicate that modified FA algorithms converge on internal representations that are structurally similar to those produced by backpropagation. In particular, it appears the functional success of modified FA algorithms may be rooted in their ability to mimic the representational geometry of backpropagation, converging on similar representations despite relying on fundamentally different weight update mechanisms.

Can Neural Assemblies -- groups of neurons that fire together and strengthen through co-activation -- learn the direction of causal influence between variables? While established as a computationally general substrate for classification, parsing, and planning, neural assemblies have not yet been shown to internalize causal directionality. We demonstrate that the inherent operations of neural assemblies -- projection, local plasticity control, and sparse winner selection -- are sufficient for directional learning. We introduce DIRECT (DIRectional Edge Coupling/Training), a mechanism that co-activates source and target assemblies under an adaptive gain schedule to internalize directed relations. Unlike backpropagation-based methods, DIRECT relies solely on local plasticity, making the resulting causal claims auditable at the mechanism level. Our findings are verified through a dual-readout validation strategy: (i) synaptic-strength asymmetry, measuring the emergent weight gap between forward and reverse links, and (ii) functional propagation overlap, quantifying the reliability of directional signal flow. Across multiple domains, the framework achieves perfect structural recovery under a supervised, known-structure setting. These results establish neural assemblies as an auditable bridge between biologically plausible dynamics and formal causal models, offering an "explainable by design" framework where causal claims are traceable to specific neural winners and synaptic asymmetries.

Synaptic plasticity is metabolically expensive, yet animals continuously update their internal models without exhausting energy reserves. However, when artificial neural networks are trained, the network parameters are typically updated on every sample that is presented, even if the sample was classified correctly. Inspired by the human negativity bias and error-related negativity, we propose 'memorized mistake-gated learning' -- a biologically plausible plasticity rule where synaptic updates are strictly gated by current and past classification errors. This reduces the number of updates the network needs to make by $50\%\sim80\%$. Mistake gating is particularly well suited in two cases: 1) For incremental learning where new knowledge is acquired on a background of pre-existing knowledge, 2) For online learning scenarios when data needs to be stored for later replay, as mistake-gating reduces storage buffer requirements. The algorithm can be implemented in a few lines of code, adds no hyper-parameters, and comes at negligible computational overhead. Learning on mistakes is an energy efficient and biologically relevant modification to commonly used learning rules that is well suited for continual learning.

Behavior can be described as a temporal sequence of actions driven by neural activity. To learn complex sequential patterns in neural networks, memories of past activities need to persist on significantly longer timescales than the relaxation times of single-neuron activity. While recurrent networks can produce such long transients, training these networks is a challenge. Learning via error propagation confers models such as FORCE, RTRL or BPTT a significant functional advantage, but at the expense of biological plausibility. While reservoir computing circumvents this issue by learning only the readout weights, it does not scale well with problem complexity. We propose that two prominent structural features of cortical networks can alleviate these issues: the presence of a certain network scaffold at the onset of learning and the existence of dendritic compartments for enhancing neuronal information storage and computation. Our resulting model for Efficient Learning of Sequences (ELiSe) builds on these features to acquire and replay complex non-Markovian spatio-temporal patterns using only local, always-on and phase-free synaptic plasticity. We showcase the capabilities of ELiSe in a mock-up of birdsong learning, and demonstrate its flexibility with respect to parametrization, as well as its robustness to external disturbances.