Synaptic Plasticity

Artificial neural networks (ANNs), as core components of modern deep learning (DL) systems, lack the adaptive flexibility and long-term stability exhibited by biological systems. This limitation largely stems from the fact that conventional ANNs rely on uniform, local, and gradient-based parameter updates, while neglecting internal learning principles that are biological mechanisms such as neurotransmitters signalling or neuroplasticity. Consequently, many existing approaches focus on architectural expansion or mathematical fine-tuning techniques such as regularisation or parameter isolation. Inspired by the superior adaptability and plasticity of mammalian brains, we posit that neuromodulation with neurotransmitters constitutes a third axis of learning, complementary to neural activity and synaptic plasticity, and should be explicitly modelled in artificial neural networks. In this positional paper, we argue that incorporating neuromodulatory principles into ANN design represents a promising and underexplored research direction, and we advocate for greater attention to this perspective in the development of adaptive and continual learning systems.

Astrocytes are non-neuronal glial cells that are receiving widespread attention due to their emerging role in neural computation. In this paper, we propose and study dynamical mechanisms by which astrocytes may augment the ability of neural networks to infer context in reinforcement learning (RL) settings. We construct a biologically inspired, two-level dynamical neural-astrocyte network with distinct spatial and temporal organization. We train this model on a hierarchical multi-context task that requires the agent to infer changes in latent task rules based on derived rewards. We find that in this setting, astrocytes enable evidence accumulation of changes in context and subsequent context-specific modulation of neural dynamics. We show that these functions are implemented via two dynamical mechanisms: (i) reward-induced bifurcations that relocate an asymptotically stable attractor into different, context-specific regions of state space, and (ii) the relative shallowness of these attractors, mediated by the entropy of the environment, giving rise to behavioral stickiness. Together, these mechanisms amount to a hybrid automaton, in which uncertainty accumulates until, eventually, the neural dynamics are switched to a new context. This model provides a neuro-dynamic schema, compatible with neural-astrocyte biology and prior empirical observations, for how astrocytes may integrate information from the periphery and drive contextual changes in neural circuits.

This work presents a fully analog memristive synaptic circuit for online spike-timing-dependent plasticity (STDP) learning in spiking neural networks (SNNs). The proposed synapse integrates a local STDP circuit generating gradual timing-dependent conductance updates directly from pre- and post-synaptic spikes. Learning occurs during normal network operation without requiring external digital control or explicit STDP waveform synthesis. Post-layout simulations of the memristive synapse implemented in a 130 nm CMOS technology show spike-timing-dependent conductance adaptation during SNN operation. A 2x2 SNN simulation further illustrates online neuron specialization through unsupervised 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.

Quantization is widely used to reduce the computational and memory demands of neural-network inference. In recurrent networks, however, the quantized state is stored and returned at the next time step, so the rule used to store that state can alter subsequent computations. Here, we introduce recurrent-state write-back to denote this rule and isolate its effect in a compact GRU encoder--decoder for fluorescence lifetime imaging, a molecular imaging modality used in quantitative biological imaging. A central task is estimating two lifetime parameters, the short-lived component {\tau}1 and the long-lived component {\tau}2, from high-noise time-resolved fluorescence signals. Holding the trained model fixed, replacing continuous state propagation with deterministic 4-bit state storage increases estimation errors for {\tau}1 and {\tau}2 by approximately 70x and 300x, respectively. Failure occurs when repeated small updates remain below the write threshold, leaving the stored state nearly fixed while the network continues to propose change. Error feedback, residual memory, and direction memory carry information from these suppressed updates across time and recover accuracy without retraining. Precision sweeps show that increasing state precision can worsen a fixed recurrent solution, while matched training shows that compatibility with the state interface can be learned. To test whether this behavior extends beyond the GRU, we repeat the post-training intervention in an independently trained LSTM, where coarse write-back reproduces the failure, error feedback restores accuracy, and state-specific interventions reveal greater sensitivity of the cell state than the hidden state. Our results establish recurrent-state write-back as a key determinant of low-precision recurrent dynamics and identify the state-storage interface as a central design consideration for quantized recurrent inference.

Cortical neurons fire sparsely -- often fewer than one spike per sensory window -- making rate coding insufficient and temporal coding a necessity. That conduction delays convert firing order into synchrony is long established. What governs which class of temporal feature a neuron detects -- one volley of coincident input, or two in a particular order -- has not been examined. We propose a delay-signature framework in which the axonal conduction delays converging on a dendritic branch constitute a physical key: only input sequences whose spike-time differences the delays compensate arrive synchronously, and coincidence detection, via calcium plateau thresholds, converts that synchrony into an all-or-none output. In simulations of an integrator-neuron model we report three results. First, a single physical scalar -- the dispersion of the delay set -- moves a population from event detection to order-selective sequence detection. The transition is emergent under random delays and connectivity: at narrow dispersion sequence detectors do not exist, and the dispersion at which they overtake event detectors tracks the inter-event interval with a slope statistically indistinguishable from one. This maps a computational distinction onto the anatomical one between myelinated and unmyelinated projections, making myelination a switch on what a neuron computes, not only a regulator of speed. Second, the same dispersion sets the code's limits: it bounds the longest codable interval and fixes an absolute timing tolerance of about a millisecond, with slowing better tolerated than speeding. Third, that millisecond window and horizontal conduction velocity together predict cortical column diameter, and the two areas with direct measurements fall where the relation puts them. One anatomically measurable parameter thus sets what a neuron detects and the limits of what it can represent.

Canonical neural circuit motifs are usually described functionally: divisive normalization rescales population activity by a pooled signal, and winner-take-all competition selects one pattern through recurrent excitation and shared inhibition. We represent them, and their compositions, algebraically as finite transformation systems and analyze the transition monoids generated by their input-conditioned updates, distinguishing structure already present in a generator from structure that appears only through composition, and, on a joint state space, structure inherited from one factor from structure that lives on a joint configuration. Individually aperiodic updates can generate non-aperiodic monoids. In the WTA, every frozen-drive generator collapses to fixed points, yet short input sequences create local cycles of winner-dependent inhibitory gating: globally dissipative dynamics with a reversible action. The strongest result arises in WTA-to-DN composition. The composed monoid then contains a genuinely composite local cycle in which normalization state and the winner's gating state change together, although every primitive generator is aperiodic. Holonomy analysis certifies this as a group component of the Krohn-Rhodes cascade rather than an incidental cycle, and finds most group-carrying image sets on joint configurations, whereas the uncoupled product has none. An exhaustive interface sweep shows that the composite cycle is a property of the coupling rather than of a chosen map. If motifs are building blocks of neural computation, composing them is a form of programming: one chooses primitives and interfaces so that the generated algebra has the intended repertoire. The transition monoid is that repertoire - what a primitive presents to any later construction. Recurrent circuits are compositional transformation systems; their algebra constrains what they can be programmed to compute.

Plasticity loss has emerged as a critical challenge in continual learning that significantly hinders the acquisition of sequential tasks. While optimizing activation designs offers a potential solution, current fixed-form functions suffer from an inherent spectral bias towards low-frequency variations, whereas learnable variants permit unconstrained updates that induce catastrophic forgetting. To address these limitations, we propose a novel learnable wavelet activation that decomposes the activation function into low-frequency and high-frequency components to explicitly counter spectral bias. Furthermore, we employ dynamic wavelet injection to adaptively enhance plasticity for new tasks, alongside a regularization strategy to ensure the stability of previous learned knowledge. Theoretically, we provide rigorous mathematical guarantees for the proposed framework, proving the structural necessity of the hybrid wavelet architecture for efficient $L^2$ approximation and demonstrating that the decoupled learning rate mechanism successfully restores network plasticity for high-frequency information. Additionally, we provide a formal derivation of the loss-driven injection trigger mechanism to precisely guide the injection. Extensive empirical evaluations demonstrate that our approach maintains superior trainability and generalization throughout the learning process and achieves state-of-the-art performance across diverse continual learning benchmarks.

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.

The ability to robustly maintain and update continuous variables is a hallmark of working memory. While classical continuous attractor networks suffer from severe fine-tuning fragility, standard artificial recurrent neural networks (RNNs) like GRUs and LSTMs typically fail to stably learn continuous manifolds, instead shattering the state space into discretized point attractors. To bridge this gap, we draw inspiration from divisive normalization, a canonical neural computation widely observed across cortical circuits, and propose the Recurrent Divisive Normalization Network (RDNN), a minimal and algebraically isolated model of dynamic division. Through dynamical systems analysis on canonical working memory tasks, we demonstrate that this biophysical constraint allows the network to converge to robust, high-fidelity slow manifolds. Furthermore, we analyze the gradient dynamics of divisive normalization during Backpropagation Through Time (BPTT), showing that it introduces an activity-dependent local gradient scaling. This scaling dampens parameter updates in highly active regimes, which empirically aligns with a significant self-compression of the network's effective rank, confining the recurrent dynamics to a tight, low-dimensional subspace while avoiding the optimization pathologies associated with explicit low-rank factorization. Finally, ablations demonstrate that while subtractive inhibition can maintain static memories, divisive normalization is mathematically essential to prevent manifold shattering under time-varying inputs. Our findings identify divisive normalization not merely as a biological artifact, but as a critical computational mechanism for learning high-fidelity continuous representations.

Brains and large language models (LLMs) are fundamentally different memory systems, but they can be compared through shared functional questions: where memory-related information is represented, how partial cues recover broader associations, how new information is written or updated, and how memory-related states can be perturbed. In biological systems, these questions span synapses, neuronal ensembles, hippocampal-cortical interactions, and plasticity; in LLMs, they span weights, activations, context windows, retrieval systems, and external stores. The comparison is therefore functional and experimental rather than anatomical. Human studies reveal sparse concept responses, temporal binding, rapid association formation, episode-specific coding, and recall-related reactivation, but selective intervention remains limited. Rodent studies provide more selective causal access to learning-related ensembles, whereas human and macaque interventions usually affect broader circuits. LLMs lack lived episodic memory, yet they permit unusually direct and repeatable manipulation of internal states and stored information. We argue that this asymmetry creates a new opportunity. LLMs are not ahead in memory itself, but in experimental access. Their tools may help turn broad questions about retrieval, updating, persistence, reversibility, and unintended effects into sharper biological hypotheses. The productive bridge is to transfer experimental logic, not anatomical parts.

Direct feedback alignment (DFA) trains hidden layers with fixed random projections of the output error, avoiding the transposed-weight backward pass of backpropagation (BP). We study a failure mode of DFA training that is distinct from feedback quality: the local weight update is calculated by an outer product, so anisotropy can enter through either its presynaptic-activity factor or its local-error factor. Our analyses with controlled synthetic regimes isolate the first failure mode and show an approximately 40-percentage-point activity-conditioning gain when high-variance directions contain task-irrelevant nuisance. Three clean confirmations isolate a different regime: error conditioning improves raw DFA by 1.77--7.53 percentage points, and combining independently selected activity and error factors adds 0.40--0.90 points over activity conditioning. The signs hold for tanh/one-vs-rest MNIST and preregistered Fashion-MNIST, and replicate on eight fresh seeds in a ReLU/softmax MNIST model. This factorization yields a symmetric block-local family of normalized DFA (nDFA): activity nDFA right-preconditions by an inverse activity second moment, error nDFA left-preconditions by an inverse local-error second moment, and K-nDFA applies both factors with separately tuned damping. A linearized post-alignment calculation gives an exact input-side spectral identity and a Kronecker-factor motivation for the two-sided rule, whereas norm matching rules out a scalar step-size explanation. The error factor is fragile when under-damped, BatchNorm is a strong activity-side alternative, and convnet gains remain partial. We therefore frame conditioned DFA as a factor-level study of when local outer-product rules fail, not as a general replacement for BP or a solution to all-layer convolutional credit assignment.

Continuous-time spiking neural networks (SNNs) provide an event-driven framework for temporal computation, computational neuroscience, and neuromorphic hardware. However, training deep continuous-time SNNs is severely constrained by the memory required for exact spike-time computation, which evaluates and retains candidate firing times over intervals determined by presynaptic spike ordering. Here we introduce a memory-efficient training framework based on differentiable spike-time discretization (DSTD) for leaky integrate-and-fire neurons with general membrane and synaptic time constants. DSTD maps irregular presynaptic spikes onto differentiable weighted events at fixed time points, replacing the input-dependent candidate dimension with $M$ fixed time intervals while accurately approximating continuous-time membrane-potential dynamics. This reduces candidate-related activation memory from $O(N_{\mathrm{out}}N_{\mathrm{in}})$ to $O(N_{\mathrm{out}}M)$ in the case of time-to-first-spike (TTFS) coding, where $N_{\mathrm{in}}$ and $N_{\mathrm{out}}$ denote the numbers of presynaptic and postsynaptic neurons, respectively. We further introduce synfire-chain-inspired temporal regularization that organizes layer-wise firing windows, mitigates dead-neuron failures, and enables pipeline-like processing. In dense LIF layers, DSTD reduced peak memory consumption by up to approximately 100-fold and training time by up to approximately 20-fold compared with exact spike-time computation. Together, these methods allowed us to train 9-layer convolutional SNNs on CIFAR-10 and 20-layer convolutional SNNs on Fashion-MNIST on a single GPU.

Interaction with AI agents has become one of the most frequent activities of everyday digital life. Whether conversing with an assistant, working with a coding copilot, or generating images, the interaction follows a common iterative loop: a request is issued, a result returned, appraised, and the request revised. We observe that this loop is a high-frequency stream of contact events -- moments at which a result meets a person and a conditioned response may fire before deliberate appraisal -- making everyday agent interaction an unrecognised neuroplastic training environment. When a result disappoints, reactive patterns of impatience, perfectionism, frustration, and self-criticism are repeatedly evoked, and under activity-dependent synaptic plasticity each uninterrupted cycle deepens the underlying pathway through long-term potentiation. Ordinary agent use may thus quietly strengthen the very patterns it provokes. We propose that the same training environment can be engaged to the opposite effect. Treating conditioned reactive patterns as physical neurone paths -- activated through a pre-cognitive feeling tone that opens a brief regulatory gap -- we develop a framework in which, at that gap, in place of the reactive re-prompt, a person performs behind-the-scenes observation: watching the neural process operate so the cascade does not complete and long-term depression weakens the path rather than potentiation strengthening it. We characterise this practice through three layers of observation and two modes of application: a user-guided mode requiring no change to existing tools, and an agent-assisted mode in which an ordinary agent is lightly configured to support observation at the gap. We illustrate the framework through generative image prompting, showing how a single frustrating session is behaviourally nearly identical whether or not it is observed, yet neurologically opposite.

Growing Neural Cellular Automata (GNCA) are capable of robust self-maintenance and self-repair, yet the internal dynamical mechanisms that support these capabilities remain poorly understood. Here, we investigate the role of internal fluctuations--temporal micro-variability of hidden channel states--in a trained GNCA model, challenging the assumption that such variability is merely residual stochastic noise. Through systematic analysis spanning update-rate sweeps, spatial correlation measurements, dimensionality reduction of collective state trajectories, localized damage experiments, transfer entropy vector field estimation, and partial information decomposition, we show that internal fluctuations are spatially structured, dynamically coupled to an attracting collective state, and associated with distributed small-magnitude updates that contribute to damage recovery. Damage induces a global deviation in latent state space followed by gradual re-convergence, and suppressing distributed small-magnitude updates associated with baseline fluctuation dynamics outside a permissive radius that encompasses the majority of the cells significantly impairs recovery. Transfer entropy analysis characterizes a spatially differentiated repair response: corrective inward flow near the damage site coexists with outward perturbation propagation at greater distances. Partial information decomposition further suggests a regime shift from synergy-dominant resting computation to redundancy-increased coordination during recovery. These findings indicate that GNCA self-repair emerges from high-dimensional nonlinear collective dynamics in which internal fluctuations serve as a functional component supporting information flow, coordination, and return toward an attracting recurrent state.

On analog neuromorphic hardware, intrinsic device noise is normally an accuracy tax. We ask whether it can instead consolidate memories. We cast per-synapse consolidation as a Doob h-transform: condition each weight's stochastic dynamics on never crossing a memory-critical barrier around its consolidated value. The conditioned diffusion gains an extra drift sigma^2 d/dw log h, a restoring force amplified by the noise variance itself that diverges at the barrier. We are explicit about novelty: the anchored drift -s(w-mu) our rule also contains is not ours (the limit of OUA, MESU, and EWC), and we surrender it. We claim only the conjunction of (a) the Doob barrier-conditioning as a synaptic rule, to our knowledge unclaimed (every h-transform use we found is generative modeling, none synaptic), and (b) a falsifiable prediction: increasing intrinsic noise non-monotonically improves sequential-task retention, an inverted-U that anchored-drift methods cannot produce. We pre-registered this as a go/no-go gate; it passes. On single-head Split-MNIST (8 seeds) the rule lifts retention 10.9 points at an interior optimum (paired Wilcoxon p=0.004), while matched OU/EWC/MESU anchors are monotone. Ablating the conditioning removes the effect; the optimum tracks the barrier; the inverted-U survives a second task stream and the realization where noise enters the forward pass. We then measure the intrinsic noise on real BrainScaleS-2 silicon (additive, trial-to-trial independent, tunable via on-chip averaging) and run the rule on the chip with its noise in the training loop: barrier-conditioning retains a prior task 15.6 points better than the matched control at matched average accuracy, a stability-plasticity shift, not a net-accuracy win (single seed; retention measured, energy modelled). Intrinsic analog noise thus becomes a consolidation dividend a digital accelerator must spend energy to generate.

Energy-efficient neuromorphic computing at the edge requires simulation tools that can capture the non-ideal behavior of mixed-signal spiking neural network (SNN) hardware while supporting system-level design exploration. This work presents an open-source hardware-aware simulation framework for mixed-signal SNNs that enables comparative analysis across neuron, synapse and architecture choices. The framework supports multiple neuron models, including Leaky Integrate-and-Fire (LIF), Hodgkin-Huxley (HH) and Axon-Hillock (AH), together with non-volatile analog synapses based on floating-gate transistors and ReRAM devices. By incorporating device-level nonlinearities directly into PyTorch-based training and inference, the tool enables optimization of physical synaptic parameters rather than idealized abstract weights. The framework is evaluated on standard neuromorphic benchmarks, including N-MNIST, DVS Gesture and Spiking Heidelberg Digits (SHD). For each model dataset configuration, it reports classification accuracy together with hardware-oriented metrics such as silicon area, power consumption and quantization sensitivity. These capabilities enable cross-layer design space exploration and help identify neuron-synapse configurations that best satisfy application-specific constraints on accuracy, energy efficiency, area and hardware fidelity.

Which properties of a partially trained network are causally portable to a different, independently trained network? Single-trajectory interventions show necessity within one run, not portability across runs. We introduce cross-trajectory chimera interventions: given two runs from different seeds, we split each weight vector into a norm and a unit direction, recombine one run's norm with the other's direction, and continue training. On two modular-arithmetic tasks that grok, the components dissociate. Direction carries a transferable, donor-specific circuit identity: implanting a donor's direction at the recipient's norm drives the run to the donor's circuit in 40/40 cases, while an angle-matched random control yields no shift. The transfer is threshold-like, and its location is predicted by the recipient's norm, separating perfectly by norm class over all 20 pairs (joint permutation probability 1.9e-4). Norm carries only a modest, distributed delay effect and no identity signal. An adaptive bisection procedure localizes the threshold to +/-1/64. Direction indexes which solution a trajectory approaches; norm governs how susceptible that identity is to being overwritten.

In-context learning (ICL) operates via implicit gradient descent embedded in the forward pass of modern AI architectures -- Transformers, Mamba, state-space models, and MLPs. Capturing this capability in biologically plausible Spiking Neural Networks (SNNs) has remained an open challenge: existing SNNs fail the Garg-2022 benchmark at non-trivial task dimensions. We trace this failure to a structural assumption: prior SNN designs route adaptation through inference-time synaptic plasticity, viewing the dendritic compartment as a passive conduit for error or teacher signals. We challenge this assumption. The subthreshold dynamics of a single dendritic compartment already implement a complete online learning algorithm. By treating the compartment as the computational substrate rather than a passive conduit, we propose DendriCL -- a single-layer compartmental spiking architecture whose apical recurrence is structurally identical to leaky online Widrow-Hoff LMS. This dynamics-only update collapses the architectural depth required for general-purpose ICL to a single layer. DendriCL is uniquely seed-stable at super-dimensional Garg-2022 ICL -- where dense Transformers exhibit grokking-style instability and fail past moderate task dimension -- and a linear probe recovers the reference online-LMS trajectory directly from the apical membrane at R^2 = 0.93, showing the algorithm is structurally embedded in the dynamics rather than implicitly discovered during training. Taken together, ICL requires neither attention, depth, nor inference-time plasticity: a single compartment with online-LMS dynamics is sufficient.

Recent breakthroughs in synaptic-resolution network connectomics have revealed that brain circuits feature fine-scale structural connectivity, such as pairs of correlated synaptic couplings known as second-order motifs. Large-scale recordings of neuronal activity in networks containing nonlinear neurons reveal macroscopic heterogeneous population dynamics throughout the brain. These findings rekindle the inquiry into this intriguing question: Can microscale synaptic structures contribute to macroscopic heterogeneous dynamics and computations in ways that canonical brain circuit models cannot? To answer this question, we create random RNNs with various cell types, nonlinear non-negative neural responses, and arbitrary marginal and second-order correlated synaptic statistics. We derive mean-field low-rank equations for P-population networks in which the pre- and postsynaptic neuronal population identities determine the synaptic and motif strengths. Our framework requires 2P latent dynamic variables with P variables describing mean population activity and P variables capturing within-population variability. Theoretical and simulational results demonstrate that chain motifs induce correlations in synaptic variability, enabling microscopic fluctuations to be integrated and influence mesoscopic mean population dynamics. We apply this framework to reverse engineer network connectivity that recapitulates the heterogeneous activity across the population in the mouse primary visual cortex. By bridging the gap between synaptic organization and nonlinear heterogeneous population dynamics, our results offer a principled approach and testable predictions regarding the relationship between fine-scale connectivity, heterogeneous dynamics, and functional computations.

Deploying spiking neural networks (SNNs) on neuromorphic hardware demands aggressive synaptic pruning while preserving temporal computation integrity. Existing strategies either neglect neuronal criticality or rely on convex relaxations of the inherently combinatorial pruning problem whose fractional masks, upon binarisation, destroy accuracy at moderate-to-high sparsity. We present Criticality-Constrained Quadratic Pruning (CQP), a native PyTorch pipeline that fuses weight magnitude with surrogate-gradient criticality into an analytically exact importance metric, eliminating the rounding artefacts endemic to solver-based approaches. We formally characterise a continuous-relaxation trap wherein OSQP-solver fractional masks overshoot the intended sparsity by up to 12 percentage points (pp), precipitating a 44 pp accuracy collapse. We identify and remediate a zombie-weight failure mode in which Adam's first-moment tensors resurrect pruned synapses, violating the binary sparsity guarantee. An iterative schedule - prune, fine-tune with gradient masking, recompute criticality, and repeat - eliminates gradient staleness at high sparsity. A KL-divergence temporal analysis identifies a redundant simulation timestep, enabling a free 10% theoretical energy reduction without weight modification. On MNIST (60,000 training examples), CQP yields 95.6% accuracy at 90% sparsity versus 93.4% for magnitude pruning (+2.2 pp). A criticality-threshold sweep reveals an empirical criticality cliff: accuracy falls from 87.0% to 14.4% as the threshold reaches tau = 0.9, constituting a quantitative SNN-level analogue of the Critical Brain Hypothesis. Combined weight sparsification and temporal truncation yield a compound 73% reduction in per-inference energy at 70% sparsity, confirming the practical value of the proposed pipeline for neuromorphic deployment.

Multi-Agent Reinforcement Learning (MARL) value factorization methods can suffer from a loss of plasticity, gradually failing to adapt when transferring to new task instances. We trace this issue to stagnant neurons, units whose gradient updates become negligibly small relative to their weights, thereby hindering learning. While existing plasticity injection methods exist, they prove ineffective for such neurons. To address this, we propose Knowledge-retentive Neuron-level PlastIcity Focusing InjEction (KNIFE), a novel method that directly targets stagnant neurons. KNIFE replaces each stagnant neuron with a composite unit comprising three specialized components: a frozen knowledge neuron to preserve acquired knowledge, a re-initialized active neuron to restore learning capacity, and a compensation neuron to ensure the combined output matches the original, thus maintaining previous learned cooperation knowledge. Extensive experiments on SMACv2, predator-prey, and matrix games demonstrate that KNIFE significantly outperforms state-of-the-art plasticity injection methods.

Backpropagation-trained dense neural networks are powerful function approximators, but they couple learning across many parameters and can overwrite previous associations when tasks conflict. This paper describes Local Pheromone Network, a small research prototype for sparse, local, manually updated neural networks. In Local Pheromone Network, each output unit reads only a fixed local neighborhood of input units subject to geometric distance and molecular-tag compatibility. Each synapse stores a weight, a short-term pheromone trace, a long-term pheromone trace, and an optional consolidation state. Training does not call automatic differentiation. Instead, every layer performs a pheromone-weighted Hebbian-style update on a budgeted subset of local synapses selected from local error and co-activity. The update budget adapts online: it shrinks when loss improves and expands toward recently active neighborhoods when loss worsens. Optional mechanisms add structural plasticity, local replay, output masks for partitioned learning, and a target-free local contrastive step. We present the implementation, learning rule, and preliminary experiments on synthetic regression, partitioned memory, conflicting memory, consolidated conflict, structural plasticity, replay, and a synthetic long-context hybrid memory task. The prototype learns local linear rules, preserves partitioned memories through tags and masks, reduces forgetting under consolidation, and uses replay under conflict.

Building on resistive communication, this paper presents a physics-based design of an on-chip neural network with multi-level memristive synapses supporting a dense spectrum of conductance states. Derived from ionic transport physics, we develop a state-variable model and quantify storable sub-levels under thermal noise, drift, and quantized conductance. We assemble these devices into a 1T1R crossbar fabric, derive the linear algebra of analog vector-matrix multiplication (VMM) under wire resistance, and design a differential synapse for signed weights. A multilayer pipeline executes inference, backpropagation, and weight updates physically in the analog domain. We derive the in-situ outer-product learning rule, its discretization onto the conductance lattice, and the resulting quantization noise. We provide energy, area, capacity, and inter-tile models, showing this substrate is ideally suited for large language models (LLMs). Our design eliminates weight movement, surpassing binary ReRAM and traditional CMOS. We detail the material stack (HfO_2-based), the FEOL/BEOL CMOS foundry-integration flow, a self-contained SPICE model, the complete memristive-FPGA neuromorphic system, and an in-memory self-attention engine with current-mode translinear softmax. Finally, a ternary BitNet datapath shows projected per-token efficiency orders of magnitude better than advanced CPUs or GPUs. The result is a self-contained hardware-native blueprint for a high-density, analog, in-memory neural processor.

Spiking Neural Networks (SNNs) offer an energy-efficient paradigm for edge AI, making them attractive for hardware acceleration. However, deploying dense SNNs on FPGAs is constrained by limited on-chip memory for synaptic weight storage. To address this bottleneck, we propose VQ4SNN, a hardware-aware architecture that reduces memory requirements through Vector Quantization (VQ)-based weight sharing. To the best of our knowledge, this is the first application of VQ to pipelined spatial-dataflow SNN accelerators on FPGAs. VQ4SNN replaces conventional weight storage with a two-level memory organization consisting of compact pointers and a shared codebook of quantized weight vectors. The proposed design integrates FPGA-aware memory mapping with analytical VQ parameter selection, enabling efficient deployment on such accelerators while preserving inference accuracy. The experimental results show a reduction of 52-61% in the total number of BRAMs compared to the state-of-the-art uncompressed FPGA SNNs without increasing overall logic utilization.

Spiking Neural Networks (SNNs) model biological neural dynamics more faithfully than classical artificial networks, but their stochastic, event-driven computation -- rooted in ion-channel noise and unreliable synaptic vesicle release -- demands probabilistic models for which deterministic abstractions are mathematically inadequate. Formal verification of such models via probabilistic model checking faces a fundamental barrier: the state space explosion problem, where the Discrete-Time Markov Chain (DTMC) encoding grows exponentially with the number of neurons. General-purpose quotient model abstractions [1] can in principle mitigate this growth by partitioning membrane potentials into equivalence classes, but a naïve application to SNNs discards synaptic weight information, limiting the properties that can be verified. This paper introduces a weight-discretized quotient model abstraction that maps continuous synaptic weights to a compact integer range while preserving the relative contribution of each synapse, and presents CogSpike, a unified workbench that integrates SNN design, simulation, and PRISM-based formal verification within a single isomorphic tool chain. The discretization is accompanied by formal correctness guarantees: a two-sided fidelity theorem confines any firing disagreement to a bounded gray zone around threshold, and an Asymptotic Silence theorem gives the exact limit guarantee that unforced neurons fall permanently silent. A topology-dependent scaling analysis shows that the state space reduction compounds exponentially -- approximately $17\times$ per neuron for discretization parameter $W = 3$ -- enabling verification of networks that are otherwise intractable, as confirmed empirically across seven canonical topologies.

Spiking Neural Networks (SNNs) offer a brain-inspired path toward highly efficient computation, but their practical deployment is constrained by the challenge of managing and executing their massive parallelism on physical hardware. This problem mirrors the historical challenge in processor design of moving beyond serial execution, a barrier broken by superscalar architectures that dispatch multiple instructions to parallel functional units. Drawing inspiration from this paradigm, we introduce a hardware-software co-design framework that treats synaptic events as parallelizable micro-operations. We present SupraSNN, a superscalar-inspired architecture that achieves high synapse-level parallelism by physically decoupling synaptic and neuronal computations. Within this architecture, a Multi-Cast Tree routes spike data to multiple parallel Synapse Processing Units serve as the computational pipelines, while a Merge Tree consolidates distributed results for processing by a unified Neuron Unit--deliberately centralizing complex neuron state dynamics to mitigate hardware overhead and resource duplication. The efficacy of this architecture is enabled by a sophisticated partitioning and scheduling framework that first maps the SNN onto hardware respecting memory constraints, then heuristic scheduling determines the synaptic execution order, maximizing throughput and resource utilization. Implementing a feedforward SNN trained on MNIST (93.44% accuracy), SupraSNN achieves 149 $μs$ inference latency and 0.025 mJ per image (0.276 nJ per synapse) on the Xilinx Zynq XC7Z020 FPGA--delivering 47.6% lower latency and 5.6$\times$ better energy efficiency than prior FPGA-based SNN accelerators. Beyond vision tasks, a recurrent SNN on the Spiking Heidelberg Dataset (71.82% accuracy) achieves 1.41 ms latency and 0.77 mJ per sample on XC7Z030.

Recent developments in brain recording are driving a demand for machine learning tools capable of decoding the latent structure of large populations of neurons. In this paper, we provide a comprehensive survey that outlines the trajectory of Latent Variable Models (LVMs) from early state-space models to more recent deep generative models. We organize the literature into three closely related domains: (1) Single-Region Latent Dynamics, which includes models such as linear dynamical systems to more complex dynamics represented by Recurrent Neural Networks (RNNs) and Neural Ordinary Differential Equations (ODEs); (2) Multi-Region Communication, which employs probabilistic as well as subspace methods to study how information is transferred across different brain areas considering synaptic propagation delays and network connectivity; and (3) Behavior-Aligned Modeling, which seeks to disentangle neural activity related to task performance from other internal states via supervised or contrastive learning. This survey also includes large-scale neural foundation models, such as Transformers and diffusion models, that rely on large-scale pre-training for optimal performance across subjects. Finally, we conclude and discuss benchmarks, evaluation criteria, and open challenges, such as the ability to identify causal links or directionality of communication, to facilitate future research for bridging interpretable brain dynamics with reliable neural decoding.

Reservoir computing exploits the fixed dynamics of a recurrent network for temporal processing, requiring only a trained linear readout. Biological neural connectomes, shaped by millions of years of evolution, may encode computational structure beyond what random reservoirs provide, yet whether that structure can be further enhanced by principled optimisation remains an open question. We address it by applying four gradient-free, bio-inspired optimisers (Particle Swarm Optimisation, Differential Evolution, Grey Wolf Optimiser, and Whale Optimisation Algorithm) to the edge weights of connectome-based echo-state networks across six species spanning six orders of magnitude in neural complexity: C. elegans (279 neurons), Drosophila (49 nodes), mouse (112), rat (73), macaque (29 regions, continuous FLNe synaptic strengths), and human structural MRI connectivity (83 parcels). Each connectome is evaluated on four canonical reservoir computing benchmarks: Memory Capacity (MC), Lorenz attractor prediction, NARMA-10 system identification, and Mackey-Glass chaotic time-series prediction. All four optimisers consistently outperform unoptimised biological baselines across every task and species when initialised from biological weights. WOA achieves the largest gains on every task: up to a 17x MC improvement (C. elegans: 1.39 to 23.91) and up to 89% NRMSE reduction (Mackey-Glass, human), corresponding to an average 214% improvement across all species and tasks. Crucially, random initialisation on the same topology reliably underperforms biology, establishing biological weight values as an essential inductive bias that topology alone cannot recover. These results position bio-inspired, biologically-initialised optimisation as a principled and broadly effective strategy for connectome reservoir computing across the animal kingdom.

The prefrontal cortex (PFC) maintains goal information for action planning, but how recurrent circuits preserve it in an action-usable form over behavioral timescales remains unclear. Here we ask whether short-term synaptic plasticity (STP) can stabilize goal information as action-usable, goal-conditioned dynamics. We incorporated STP into a PFC-inspired reservoir computing model with basal-ganglia-inspired temporal-difference readout learning, and evaluated paired models with and without STP across 100 independently generated networks in a multistep goal-directed action-selection task with delayed execution. Goal identity was highly decodable during the delay even without STP, so STP was not required to form a linearly readable goal representation. Under state noise, however, success without STP fell from 75.8% to 49.5%, whereas the model with STP remained essentially unchanged (91.8% without noise versus 89.2% under noise; paired Cohen's dz=1.31). Time-resolved decoding, state-space separability, and action-value-difference analyses showed that STP preserved goal information as action-relevant goal-conditioned dynamics available at later action opportunities. Gain-matched and STP-state perturbation controls argued against a simple fixed recurrent-scaling explanation and supported online, history-dependent synaptic modulation. Effective-connectivity analyses showed delay-period goal-specific patterning that increased toward the later part of the trial with STP, where it should be read as goal- and task-state-conditioned patterning; effective connectivity without STP was time-invariant. A grid search identified a facilitation-dominant range of STP time constants associated with high success rates. These results suggest that STP supports robust goal-conditioned dynamics through dynamic modulation of goal-dependent effective recurrent connectivity.

Spiking neural networks (SNNs) provide event-driven and low-power computation inspired by biological neural systems, but current implementations rely on von Neumann graphics processing units (GPUs) and central processing units (CPUs) platforms, where memory and computation bottlenecks limit energy efficiency. To address this challenge, this paper proposes an analog memristor-based spiking neural network (SNN) accelerator that integrates in-memory synaptic computation with analog integrate-and-fire (IF) neurons, eliminating multi-transistor CMOS synapse circuits and enabling asynchronous event-driven operation at the 45nm technology node. Additionally, a digital SNN accelerator is designed and optimized at the 5 nm technology node for comparison. The proposed architecture is evaluated using a predator-prey tracking task that emulates pursuit behavior. In this task, the analog SNN accelerator's inference closely matches the ideal software inference with a mean squared error (MSE) of 0.004. HSPICE simulation results show that the proposed analog SNN accelerator achieves 12.7 times lower energy consumption and 1.26 times lower delay compared to the digital baseline, demonstrating the potential of memristor-based neuromorphic circuits for energy-efficient real-time edge intelligence.

A hallmark of intelligence is the ability to adapt in non-stationary environments, yet deep Reinforcement Learning (RL) agents often struggle in such settings. Prior studies introduce non-stationarity through abrupt shifts in features or dynamics, whereas real-world environments often evolve gradually through continual drift. This distinction has important implications for the "stability-plasticity dilemma" in RL, as abrupt task changes may demand more plasticity than naturalistic settings. To address this, we modify existing 3D Miniworld and MuJoCo environments to incorporate naturalistic, continual non-stationarity, and use them to examine how stability and adaptation affect performance under continuous environmental change. We find that methods favoring stability, such as synaptic consolidation, outperform approaches focused on plasticity, such as parameters resetting. Motivated by this result, and prior evidence that Successor Features (SFs) reduce interference, we investigate whether SFs are better consolidation targets than Q-values. Across both environments, applying neuro-inspired synaptic consolidation to SFs yields superior performance on continually changing settings. Moreover, consolidation is most effective when SFs are stabilized across multiple timescales, which capture complementary aspects of gradual environmental change. Together, these results suggest that stability is more critical in continual learning when changes are gradual, and that multi-timescale consolidation of predictive representations is an effective approach.

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.

Post-Traumatic Stress Disorder (PTSD) is fundamentally a neuroplastic problem traumatic contact events encode over-reactive neural pathways through Hebbian long-term potentiation, producing hair-triggered amygdala-HPA stress cascades that fire before conscious awareness can intercept them. Existing therapeutic approaches, prolonged exposure, EMDR, cognitive behavioural therapy, operate predominantly downstream of the reactive cascade, teaching patients to tolerate or reframe distress after it has arisen. While clinically valuable, these suppression-based approaches do not produce the upstream pathway dissolution that constitutes lasting structural neural reorganisation. This paper proposes MindGap, a privacy-preserving on-device conversational AI framework that delivers structured neuroplastic rehabilitation for PTSD through the practice of dependent origination, a Buddhist psychological framework that identifies the precise moment between the pre-cognitive affective signal and the reactive elaboration that follows as the site of therapeutic intervention. MindGap guides patients through three progressive layers of observation at this feeling tone gap: noticing the bare affective signal before reactive elaboration, recognising it as self-arising rather than caused by the stimulus, and recognising the conditioned implicit belief beneath the feeling. Each layer corresponds to progressively deeper prefrontal regulatory engagement and progressively deeper long-term depression-mediated weakening of the reactive pathway, producing genuine upstream dissolution rather than downstream suppression. Running entirely on-device with no data egress, MindGap delivers daily calibrated exposure sessions through a fine-tuned lightweight large language model, making it deployable in sensitive clinical and military contexts where cloud-based solutions are not permitted.

Synapses are densely packed submicron structures that dynamically reorganize during learning and memory formation. Longitudinal \textit{in vivo} imaging of fluorescently tagged synaptic receptors offers a promising opportunity to study large-scale synaptic dynamics and how these processes are disrupted in neurological disease. However, in vivo imaging with 2-photon microscopy uses low laser power and therefore suffers from low signal-to-noise ratio (SNR) and high shot noise, nonlinear tissue motion between days, nonstationary fluctuations in synaptic fluorescence, and significant blur induced by the microscope point spread function (PSF). Together, these factors make it challenging to detect and track synapses, especially in regions with high synaptic density. This paper presents a novel template-based framework for modeling synapses as varying luminance point sources that move under a nonlinear tissue deformation. Taking a unified Bayesian approach, we apply this model to microscopy data by deriving a posterior that incorporates a diffeomorphic mapping for domain warping, a Gaussian point spread function for the imaging process, and a Poisson observation model for raw photon counts. The Bayesian solution simultaneously: (1) Constructs a probabilistic template of synapse locations, (2) denoises and deconvolves the image data, (3) infers fluorescence intensities, (4) performs diffeomorphic image registration to correct for tissue motion, and (5) provides confidence regions for these parameter estimates. We demonstrate the framework on both a 2D+t simulated dataset and a 3D+t longitudinal \textit{in vivo} microscopy dataset of fluorescent synapses imaged in a mouse over two weeks.

Short-term plasticity (STP) is fundamental to temporal information processing in biological neural systems but remains difficult to realize efficiently in neuromorphic hardware. Memristive electrochemical random-access memory (ECRAM) devices naturally exhibit non-equilibrium ionic dynamics that produce transient conductance modulation; however, these behaviors are typically treated as undesirable variability or tolerated as side effects in memory-centric computing paradigms. In this work, we instead transform these volatile dynamics from a tolerated device artifact into a computational resource through a cross-layer device-circuit-system co-design framework. We introduce a delay-feedback leaky integrate-and-fire (LIF) neuron architecture co-designed with ECRAM synapses that exploits activity-dependent conductance modulation with negligible additional circuit overhead. The architecture integrates ECRAM-based synapses with a tunable delay-feedback spike-generation path, enabling transient device dynamics to directly modulate neuron excitability and synaptic efficacy. We used experimentally characterized ECRAM devices exhibiting transient conductance modulation (1.5 KOhms per spike) to develop a compact behavioral model suitable for circuit-level simulation. Circuit simulations demonstrate two key STP behaviors -- synaptic facilitation and intrinsic excitability modulation -- while consuming 2 pJ per spike, and the same device-driven mechanisms extend across multiple neuron topologies. Network-level analysis further demonstrates frequency-selective spike processing, allowing individual synapses to act as tunable temporal filters within spiking neural networks. This work demonstrates that non-equilibrium ECRAM dynamics can serve as a native hardware substrate for STP and temporal computation in neuromorphic circuits.

LLMs are trained once, then deployed into a world that never stops changing. External memory compensates for this, but most systems manage it explicitly rather than letting it adapt on its own. Biological memory works differently: coupled multi-timescale dynamics make new associations immediately usable, strengthen what repetition confirms, and let the rest fade. We argue that external memory should follow a similar principle. In Memini, this view takes the form of an associative memory that organizes knowledge as a directed graph. Each edge carries two coupled internal variables, one fast and one slow, following the Benna-Fusi model of synaptic consolidation. From this coupling, episodic sensitivity, gradual consolidation, and selective forgetting are expected to emerge as facets of a single mechanism, reframing external memory as a learning substrate that reorganizes through its own dynamics. This workshop article describes an early-stage conceptual design without experimental evaluation.

Learning in neural systems arises from synaptic changes that reshape the representations underlying behavior. While low-rank recurrent neural networks (RNNs) have emerged as a powerful framework for linking connectivity to function, a theoretical understanding of their learning process remains elusive. Here, we extend the low-rank framework from activity to learning by deriving gradient-descent dynamics directly in a reduced overlap space. We formulate a closed-form, low-dimensional system of ODEs that governs learning in this space, exact for linear RNNs and asymptotically exact for nonlinear RNNs in the large-$N$ Gaussian limit. Central to our analysis is a distinction between two classes of overlaps: loss-visible overlaps, which fully determine network activity, output, and loss, and loss-invisible overlaps, which do not affect function but are required to describe learning. We illustrate the consequences of this decomposition through two phenomena. First, we show that learning can serve as a perturbation that exposes differences in connectivity between functionally equivalent networks. Second, we show that loss-invisible overlaps can act as memory variables that encode training history, and characterize the conditions under which this occurs. Finally, we present several testable predictions for biological learning experiments derived from our theory.

Continual learning systems face a fundamental tension between plasticity -- acquiring new knowledge -- and stability -- retaining prior knowledge. We introduce MPCS (Multi-Plasticity Continual System), a neuroplastic architecture that integrates eleven complementary mechanisms: task-driven neurogenesis, Fourier-encoded inputs, EWC regularization, meta-replay, mixed consolidation, hybrid gating, synapse pruning/regeneration, Hebbian updates, task similarity routing, adaptive growth control, and continuous neuron importance tracking. We evaluate MPCS on MEP-BENCH, a multi-track benchmark spanning 31 tasks across regression, classification, logic, and mixed domains, using a three-dimensional Pareto criterion over task performance (Perf), representation diversity (RD), and gradient conflict rate (GCR). Across 15 ablation configurations (3 seeds x 4 tracks x 2000 epochs), MPCS achieves a Normalized Efficiency Score of 94.2, placing it on the Pareto frontier among 9 of 14 gate-passing systems. Key findings: (i) Fourier encoding is the single most critical component (removal drops Perf by 30.7 pp and fails the MEP gate on 14% of tasks); (ii) global EWC degrades performance (NES = -4.2); topology-local EWC reduces this penalty (NES 90.5->91.8) but does not eliminate it; removing EWC entirely yields MPCS_EFFICIENT, the highest-Perf system -- establishing a monotone relationship in the high task-similarity regime (s_bar ~= 0.95): global EWC < topology EWC < no EWC; (iii) the Pareto status assessment is predictive: removing the two Pareto-dominated components (EWC + Hebbian) jointly yields MPCS_EFFICIENT, which improves Perf by 0.6 pp at 4.7x lower compute cost (127 vs. 602 min), validating the Pareto frontier as an actionable model-compression guide.

Spiking neural networks (SNNs) are promising for edge sensing due to their event-driven computation and temporal filtering capability. However, standard leaky integrate-and-fire (LIF) neurons communicate only through binary spikes, which severely limit representational capacity. Existing multi-level spiking neurons improve information transmission, but often rely on uniform quantization that mismatches membrane-potential distributions or introduces costly synaptic multiplications. In this paper, we propose ShiftLIF, a multi-level spiking neuron that maps membrane potentials to a logarithmically spaced power-of-two spike set. This design provides finer representation in the small-amplitude regime, where membrane potentials are densely concentrated, while enabling multiplier-free synaptic computation through bit-shift and accumulation operations. As a result, ShiftLIF improves spike-level expressiveness without sacrificing the hardware-friendly nature of standard SNN computation. We evaluate ShiftLIF on 10 datasets spanning wireless, acoustic, motion, and visual sensing tasks. Results show that ShiftLIF consistently matches or exceeds the accuracy of existing multi-level spiking neurons while maintaining synaptic energy consumption close to standard binary LIF. These results indicate that ShiftLIF provides a favorable accuracy-efficiency trade-off for cross-modal edge sensing.

In the pursuit of autonomous learning systems, the foundational assumption of stationarity, the premise that data distributions and model behaviors remain constant, is fundamentally untenable. Historically, the research community has addressed non-stationary environments almost exclusively under the scope of concept drift, focusing primarily on temporal shifts in streams. However, as learning systems become increasingly autonomous and complex, merely adapting to temporal non-stationarity is no longer sufficient. Evolving beyond this traditional perspective, we propose a novel, three-dimensional taxonomy that systematizes the field based on the operational state of the system. First, time stream drift distinguishes between stochastic arbitrary patterns and structural rhythmic dynamics. Second, data stream drift disentangles shifts in feature representations, identified as representation drift, from changes in underlying semantics, recognized as semantic drift. Third, model stream drift characterizes the internal endogenous divergence of learning systems through the lenses of sequential plasticity, decentralized heterogeneity, and policy instability. Based on this framework, we systematically review 193 representative studies and identify key open challenges. By bridging the fragmented paradigms of drift adaptation, continual learning, and temporal generalization, this survey outlines a roadmap for building self-evolving intelligent systems capable of learning autonomously through continuous change.

Spiking Neural Networks (SNNs) are widely regarded as an energy-efficient paradigm for modeling and processing temporal and event-driven information. Incorporating delays in SNNs has been proven to be an effective mechanism for improving spike alignment in event-driven tasks. However, existing delay learning approaches predominantly assign static delays to individual synapses, resulting in a large number of delay parameters and limited adaptability to input-dependent activity dynamics. To this end, we propose a Congestion-Aware Dynamic Axonal Delay (CADAD) mechanism, which decomposes the delay into a channel-wise static base delay for temporal structuring and a global, activity-conditioned shift that dynamically regulates the state update rate under varying spike intensities. The delay parameters are learned using differentiable linear interpolation and discretized at inference time, preserving the benefits of dynamic delay modulation while incurring only minimal additional cost. Experiments on speech benchmarks, including the Spiking Heidelberg Dataset, Spiking Speech Commands, and Google Speech Commands, demonstrate that introducing congestion-aware delays into synaptic signal transmission effectively improves accuracy on temporal tasks, notably achieving 93.75% accuracy on SHD, 80.69% accuracy on SSC, and 95.58% on GSC-35, while reducing the parameter count by approximately 50% compared to state-of-the-art delay-based methods with the same architecture.

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.

In a continual learning setting, we require a model to be plastic enough to learn a new task and stable enough to not disturb previously learned capabilities. We argue that this dilemma has an architectural root. A finite network has limited representational and plastic resources, yet the required capacity depends on properties of the future task stream that are unknown: how many tasks will be encountered, and how much they overlap in feature space. Regularization-based methods preserve past knowledge within fixed-capacity architectures and therefore implicitly rely on an oracle architecture sized for this unknown future. When tasks are only weakly related, fixed architectures progressively run out of plastic resources; when tasks are few or strongly overlapping, models are often over-provisioned. Inspired by neurogenesis in biology, we propose NORACL to address the stability-plasticity dilemma by tackling the oracle architecture problem through neuronal growth. Starting from a compact network, NORACL grows only when needed by monitoring two complementary signals for representational and plasticity saturation. We evaluate NORACL against oracle-sized static baselines across varying task counts and geometries. Across all settings, NORACL achieves final average accuracies that are better than or on par with oracle-provisioned static baselines while using fewer parameters. Additionally, NORACL yields architectures with interpretable growth, i.e. dissimilar tasks predominantly expand feature-extraction layers, whereas tasks which rely on common features shift growth toward later feature-combination layers. Our analysis further explains why fixed-capacity networks lose plasticity as tasks accumulate, whereas NORACL creates fresh capacity for new tasks through growth. Together, these results show that adaptive neurogenesis pushes the stability-plasticity Pareto frontier of continual learning.

Optimization algorithms are fundamental to modern deep learning, yet most widely used methods rely on update rules based primarily on local gradient statistics. We introduce NeuroPlastic, a plasticity-modulated optimizer that augments gradient-based updates with an adaptive multi-signal modulation mechanism inspired by multi-factor synaptic plasticity, a concept from neurobiology. NeuroPlastic dynamically scales gradient updates using interacting components that capture gradient, activity-like, and memory-like statistics, forming a lightweight modulation layer compatible with standard deep learning training pipelines. Across image classification benchmarks, NeuroPlastic consistently improves over a controlled gradient-only ablation, with more pronounced gains on the Fashion-MNIST benchmark and in reduced-data regimes. In transfer experiments on CIFAR-10 with ResNet-18, the method remains stable and competitive without retuning. These results suggest that multi-signal plasticity-inspired modulation can provide a useful extension to conventional gradient-driven optimization, particularly when learning signals are limited or noisy, and offer a promising direction for gradient-based methods in deep learning.

Lateral predictive coding (LPC) is a simple theoretical framework to appreciate feature detection in biological neural circuits. Recent theoretical work [Huang et al., Phys.Rev.E 112, 034304 (2025)] has successfully constructed optimal LPC networks capable of extracting non-Gaussian hidden input features by imposing the tradeoff between energetic cost and information robustness, but the resulting dynamical systems of recurrent interactions can be very slow in responding to external inputs. We investigate response-time reduction in the present paper. We find that the characteristic response time of the LPC system can be minimized to closely approaching the lower-bound value without compromising the mean predictive error (energetic cost) and the information robustness of signal transmission. We further demonstrate that optimal LPC networks taking a modular structural organization with extensively reduced number of lateral interactions are equally excellent as all-to-all completely connected networks, in terms of feature detection performance, response time, energetic cost and information robustness.

We introduce the Tan-HWG framework (Hebbian-Wasserstein-Geometry), a geometric theory of Hebbian plasticity in which memory states are modeled as probability measures evolving through Wasserstein minimizing movements. Hebbian learning rules are formalized as Hebbian energies satisfying a sequential stability condition, ensuring well-posed fiberwise JKO updates, optimal-transport realizations, and an energy descent inequality. This variational structure induces a fundamental separation between internal and observable dynamics. Internal memory states evolve along Wasserstein geodesics in a latent curved space, while observable quantities, such as effective synaptic weights, arise through geometric projection maps into external spaces. Simplicial projections recover classical affine schemes (including exponential moving averages and mirror descent), while revealing synaptic competition and pruning as geometric consequences of mass redistribution. Hilbertian projections provide a geometric account of phase alignment and multi-scale coherence. Classical neural networks appear as flat projections of this curved dynamics, while the framework naturally accommodates richer distributional representations, including structural weights and embedding memories, and their spectral extensions in complex internal spaces. Under mild Lipschitz regularity assumptions, including a quasi-stationary "sleep-mode" regime, we establish the existence of continuous-time limit curves. This yields a variational formulation of memory consolidation as a perturbed Wasserstein gradient flow. The framework thus provides a unified geometric foundation for synaptic plasticity, representation dynamics, and context-dependent computation.

Spiking neural networks (SNNs) are rapidly gaining momentum as an alternative to conventional artificial neural networks in resource constrained edge systems. In this work, we continue a recent research line on recurrent SNNs where axonal delays are learned at runtime along with the other network parameters. The first proposed approach, dubbed DelRec, demonstrated the benefit of recurrent delay learning in SNNs. Here, we extend it by advocating the use of convolutional recurrent connections in conjunction with the DelRec delay learning mechanism. According to our tests on an audio classification task, this leads to a streamlined architecture with smaller memory footprint (around 99% savings in terms of number of recurrent parameters) and a much faster (52x) inference time, while retaining DelRec's accuracy. Our code is available at: https://github.com/luciozebendo/delrec_snn/tree/conv_delays

Experimental evidence indicates that intrinsic temporal dynamics operating across multiple time scales are closely associated with the emergence of periodic spatial activity of increasing complexity. However, how information encoded in grid-like firing patterns for path integration is processed across these intrinsic time scales remains unclear. To address this question, we introduce adaptive time scales through a leak term in recurrent neural networks (RNNs), forming leaky RNNs discretized from the continuous attractors of firing rate models. Our results demonstrate that leaky RNNs substantially enhance the emergence of well-defined and highly regular hexagonal firing patterns. Compared with vanilla RNNs lacking a leak term, the trained leaky RNNs produce more accurate position estimates while generating reliable grid-cell-like representations. Furthermore, under identical noise conditions, leaky RNNs consistently exhibit more stable dynamics and better-defined grid structures. The learned dynamics also give rise to stable torus attractors with a clear central hole, supporting robust and regular grid-like activity. Overall, the dynamic leak acts as a low-pass filtering mechanism that protects recurrent neural circuitry from noise, stabilizes network dynamics, and improves path-integration accuracy in recurrent neural networks.

This work simulates the developmental process of cortical neurogenesis, initiating from a single stem cell and governed by gene regulatory rules derived from mouse single-cell transcriptomic data. The developmental process spontaneously generates a heterogeneous population of 5,000 cells, yet yields only 85 mature neurons - merely 1.7% of the total population. These 85 neurons form a densely interconnected core of 200,400 synapses, corresponding to an average degree of 4,715 per neuron. At iteration zero, this minimal circuit performs at chance level on MNIST. However, after a single epoch of standard training, accuracy surges to over 90% - a gain exceeding 80 percentage points - with typical runs falling in the 89-94% range depending on developmental stochasticity. The identical circuit, without any architectural modification or data augmentation, achieves 40.53% on CIFAR-10 after one epoch. These findings demonstrate that developmental rules sculpt a domain-general topological substrate exceptionally amenable to rapid learning, suggesting that biological developmental processes inherently encode powerful structural priors for efficient computation.

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.

Large Language Models (LLMs) generalize across tasks through reusable representations and flexible reasoning, yet remain brittle in real deployment when faced with evolving tasks and continual distribution shift. While test-time adaptation addresses this by updating models with unsupervised objectives on test data, prevailing methods are fundamentally limited by their neglect of source knowledge preservation and adaptation signal reliability. Inspired by how Drosophila orchestrates memory update by balancing retroactive and proactive interference via Rac1 and MAPK pathways, we design Synapse Consolidation (SyCo) with two core components: a Rac1-inspired plasticity confiner and a MAPK-inspired update controller. The former dynamically confines plasticity to a tail-gradient subspace that is less critical for source knowledge, enabling rapid specialization while preserving source representations. The latter uses a tiered controller to suppress noisy updates and consolidate useful adaptations under non-stationary streams. To further model real deployments with multiple sources and continually emerging tasks, we introduce Multi-source Open-set Adaptation (MOA) setting, where a model is trained on multiple labeled source tasks and then adapts on open, non-stationary unlabeled test streams mixing seen and unseen tasks with partial overlap in label and intent space. Across 18 NLP datasets under the MOA setting, SyCo consistently outperforms strong baselines, achieving 78.31% on unseen-task adaptation and 85.37% versus unseen-data shifts, setting a new state-of-the-art.

Training recurrent neuronal networks consisting of excitatory (E) and inhibitory (I) units with additive noise for working memory computation slows and diversifies inhibitory timescales, leading to improved task performance that is attributed to emergent marginally stable equilibria [PNAS 122 (2025) e2316745122]. Yet the link between trained network characteristics and their roles in shaping desirable dynamical landscapes remains unexplored. Here, we investigate the Jacobian matrices describing the dynamics near these equilibria and show that they are sparse, non-Hermitian rectangular-block matrices modified by heterogeneous synaptic decay timescales and activation-function gains. We specify a random matrix ensemble that faithfully captures the spectra of trained Jacobian matrices, arising from the inhibitory core - excitatory periphery network motif (pruned E weights, broadly distributed I weights) observed post-training. An analytic theory of this ensemble is developed using statistical field theory methods: a Hermitized resolvent representation of the spectral density is processed with a supersymmetry-based treatment in the style of Fyodorov and Mirlin. In this manner, an analytic description of the spectral edge is obtained, relating statistical parameters of the Jacobians (sparsity, weight variances, E/I ratio, and the distributions of timescales and gains) to near-critical features of the equilibria essential for robust working memory computation.

Working memory requires the brain to maintain information from the recent past to guide ongoing behavior. Neurons can contribute to this capacity by slowly integrating their inputs over time, creating persistent activity that outlasts the original stimulus. However, when these slowly integrating neurons are organized hierarchically, they introduce cumulative delays that create a fundamental challenge for learning: teaching signals that indicate whether behavior was correct or incorrect arrive out-of-sync with the neural activity they are meant to instruct. Here, we demonstrate that neurons enhanced with an adaptive current can compensate for these delays by responding to external stimuli prospectively -- effectively predicting future inputs to synchronize with them. First, we show that such prospective neurons enable teaching signal synchronization across a range of learning algorithms that propagate error signals through hierarchical networks. Second, we demonstrate that this successfully guides learning in slowly integrating neurons, enabling the formation and retrieval of memories over extended timescales. We support our findings with a mathematical analysis of the prospective coding mechanism and learning experiments on motor control tasks. Together, our results reveal how neural adaptation could solve a critical timing problem and enable efficient learning in dynamic environments.

Recurrent neural networks are canonical models of biological memory. In these models, memories are represented by distributed patterns of neural activity that are stored in the recurrent connections between neurons, such that they become attractors of the network's dynamics. During memory recall, network dynamics thus converge toward one of these memory patterns when started from a noisy or partial cue. Therefore, memory performance critically hinges on the dynamical stability of the stored patterns. However, previous theoretical approaches only studied dynamical stability under highly restrictive conditions that do not readily apply to biological neural circuits. Here, we develop a theory of the local stability of discrete fixed points in a broad class of networks with graded neural activities and in the presence of noise. Using methods from random matrix theory, we analyze the bulk and outliers of the eigenvalue spectra of the Jacobians that characterize network dynamics around fixed points. We show that either all fixed points are stable or all of them are unstable, depending on whether their number is below a ``critical load for stability'', which is distinct from the classical critical capacity that measures the maximal number of achievable fixed points regardless of their stability. We further analyze the dependence of this critical load for stability on experimentally measurable quantities characterizing the statistics of memory patterns and the activation functions of neurons. Our analysis highlights the computational benefits of sparse-like patterns and threshold-linear activation functions and offers testable predictions for neural circuits supporting memory.

Continuous adaptive learning, the ability to adapt to the environment and keep improving performance, is a hallmark of natural intelligence. Biological organisms excel in acquiring, transferring, and retaining knowledge while adapting to volatile environments, making them a source of inspiration for artificial neural networks (ANNs). This study explores how neuromodulation, a building block of learning in biological systems, can help address catastrophic forgetting and enhance the robustness of ANNs in continual learning. Driven by neuromodulators including dopamine (DA), acetylcholine (ACh), serotonin (5-HT) and noradrenaline (NA), neuromodulatory processes in the brain operate at multiple scales, facilitating dynamic responses to environmental changes through mechanisms ranging from local synaptic plasticity to global network-wide adaptability. Importantly, the relationship between neuromodulators and their interplay in modulating sensory and cognitive processes is more complex than previously expected, demonstrating a "many-to-many" neuromodulator-to-task mapping. To inspire neuromodulation-aware learning rules, we highlight (i) how multi-neuromodulatory interactions enrich single-neuromodulator-driven learning, (ii) the impact of neuromodulators across multiple spatio-temporal scales, and correspondingly, (iii) strategies for approximating and integrating neuromodulated learning processes in ANNs, and (iv) an architectural-general formulation of multi-neuromodulatory dynamics. We also present a conceptual study to showcase how neuromodulation-inspired mechanisms, such as DA-driven reward processing and NA-based cognitive flexibility, can enhance ANN performance in a Go/No-Go task. Though multi-scale neuromodulation, we aim to bridge the gap between biological and artificial learning, paving the way for ANNs with greater flexibility, robustness, and adaptability.