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1. Calcium-modulated supervised spike-timing-dependent plasticity for readout training and sparsification of the liquid state machine

   Jin, Y; Li, P (May 2017, Proceedings of ... International Joint Conference on Neural Networks)

   The Liquid State Machine (LSM) is a promising model of recurrent spiking neural networks. It consists of a fixed recurrent network, or the reservoir, which projects to a readout layer through plastic readout synapses. The classification performance is highly dependent on the training of readout synapses which tend to be very dense and contribute significantly to the overall network complexity. We present a unifying biologically inspired calcium-modulated supervised spike-timing dependent plasticity (STDP) approach to training and sparsification of readout synapses, where supervised temporal learning is modulated by the post-synaptic firing level characterized by the post-synaptic calcium concentration. The proposed approach prevents synaptic weight saturation, boosts learning performance, and sparsifies the connectivity between the reservoir and readout layer. Using the recognition rate of spoken English letters adopted from the TI46 speech corpus as a measure of performance, we demonstrate that the proposed approach outperforms a baseline supervised STDP mechanism by up to 25%, and a competitive non-STDP spike-dependent training algorithm by up to 2.7%. Furthermore, it can prune out up to 30% of readout synapses without causing significant performance degradation. 

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2. Heterosynaptic plasticity in memristive and memcapacitive lipid bilayers: A snapshot review

   https://doi.org/10.1557/s43580-024-00800-9  

   Bolmatov, Dima; Katsaras, John; Collier, C Patrick (February 2024, MRS Advances)

   Synaptic plasticity refers to activity-dependent synaptic strengthening or weakening between neurons. It is usually associated with homosynaptic plasticity, which refers to a synaptic junction controlled by interactions between specific neurons. Heterosynaptic plasticity, on the other hand, lacks this specificity. It involves much larger populations of synapses and neurons and can be associated with changes in synaptic strength due to nonlocal alterations in the ambient electrochemical environment. This paper presents specific examples demonstrating how variations in the ambient electrochemical environment of lipid membranes can impact the nonlinear dynamical behaviors of memristive and memcapacitive systems in droplet interface bilayers (DIBs). Examples include the use of pH as a modulatory factor that alters the voltage-dependent memristive behavior of alamethicin ion channels in DIB lipid bilayers, and the discovery of long-term potentiation (LTP) in a lipid bilayer-only system after application of electrical stimulation protocols. 

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3. Long-term potentiation expands information content of hippocampal dentate gyrus synapses

   https://doi.org/pnas.1716189115  

   Cailey Bromer, Thomas M. (February 2018, Proceedings of the National Academy of Sciences of the United States of America)

   An approach combining signal detection theory and precise 3D reconstructions from serial section electron microscopy (3DEM) was used to investigate synaptic plasticity and information storage capacity at medial perforant path synapses in adult hippocampal dentate gyrus in vivo. Induction of long-term potentiation (LTP) markedly increased the frequencies of both small and large spines measured 30 minutes later. This bidirectional expansion resulted in heterosynaptic counterbalancing of total synaptic area per unit length of granule cell dendrite. Control hemispheres exhibited 6.5 distinct spine sizes for 2.7 bits of storage capacity while LTP resulted in 12.9 distinct spine sizes (3.7 bits). In contrast, control hippocampal CA1 synapses exhibited 4.7 bits with much greater synaptic precision than either control or potentiated dentate gyrus synapses. Thus, synaptic plasticity altered total capacity, yet hippocampal subregions differed dramatically in their synaptic information storage capacity, reflecting their diverse functions and activation histories. 

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4. Stability through plasticity: Finding robust memories through representational drift

   https://doi.org/10.1073/pnas.2500077122  

   Natrajan, Maanasa; Fitzgerald, James E (November 2025, Proceedings of the National Academy of Sciences)

   Memories are believed to be stored in synapses and retrieved by reactivating neural ensembles. Learning alters synaptic weights, which can interfere with previously stored memories that share the same synapses, creating a trade-off between plasticity and stability. Interestingly, neural representations change even in stable environments, without apparent learning or forgetting—a phenomenon known as representational drift. Theoretical studies have suggested that multiple neural representations can correspond to a memory, with postlearning exploration of these representation solutions driving drift. However, it remains unclear whether representations explored through drift differ from those learned or offer unique advantages. Here, we show that representational drift uncovers noise-robust representations that are otherwise difficult to learn. We first define the nonlinear solution space manifold of synaptic weights for fixed input–output mappings, which allows us to disentangle drift from learning and forgetting and simulate drift as diffusion within this manifold. Solutions explored by drift have many inactive and saturated neurons, making them robust to weight perturbations due to noise or continual learning. Such solutions are prevalent and entropically favored by drift, but their lack of gradients makes them difficult to learn and nonconducive to future learning. To overcome this, we introduce an allocation procedure that selectively shifts representations for new stimuli into a learning-conducive regime. By combining allocation with drift, we resolve the trade-off between learnability and robustness. 

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5. NeoHebbian synapses to accelerate online training of neuromorphic hardware

   https://doi.org/10.1038/s41598-026-35641-z  

   Pande, S; Bezugam, S S; Bhattacharya, T; Wlazlak, E; Chakravorty, A; Chakrabarti, B; Strukov, D (December 2026, Scientific Reports)

   Abstract Neuromorphic systems that employ advanced synaptic learning rules, such as the three-factor learning rule, require synaptic devices of increased complexity. Herein, a novel neoHebbian artificial synapse utilizing ReRAM devices has been proposed and experimentally validated to meet this demand. This synapse features two distinct state variables: a neuron coupling weight and an “eligibility trace” that dictates synaptic weight updates. The coupling weight is encoded in the ReRAM conductance, while the “eligibility trace” is encoded in the local temperature of the ReRAM and is modulated by applying voltage pulses to a physically co-located resistive heating element. The utility of the proposed synapse has been investigated using two representative tasks: first, temporal signal classification using Recurrent Spiking Neural Networks (RSNNs) employing the e-prop algorithm, and second, Reinforcement Learning (RL) for path planning tasks in feedforward networks using a modified version of the same learning rule. System-level simulations, accounting for various device and system-level non-idealities, confirm that these synapses offer a robust solution for the fast, compact, and energy-efficient implementation of advanced learning rules in neuromorphic hardware. 

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