Neuropeptides, members of a large and evolutionarily ancient family of proteinaceous cell-cell signaling molecules, are widely recognized as extremely potent regulators of brain function and behavior. At the cellular level, neuropeptides are known to act mainly via modulation of ion channel and synapse function, but functional impacts emerging at the level of complex cortical synaptic networks have resisted mechanistic analysis. New findings from single-cell RNA-seq transcriptomics now illuminate intricate patterns of cortical neuropeptide signaling gene expression and new tools now offer powerful molecular access to cortical neuropeptide signaling. Here we highlight some of these new findings and tools, focusing especially on prospects for experimental and theoretical exploration of peptidergic and synaptic networks interactions underlying cortical function and plasticity.
Connectomics has focused primarily on the mapping of synaptic links in the brain; yet it is well established that extrasynaptic volume transmission, especially via monoamines and neuropeptides, is also critical to brain function. Here we present a draft monoamine connectome, along with a partial neuropeptide connectome, for the nematode C. elegans, based on new and published expression data for biosynthetic genes and receptors. Thus, the neuronal connectome can be represented as a multiplex network, with synaptic, gap junction, and neuromodulatory layers representing alternative modes of inter…
Cortical neurons are subject to sustained and irregular synaptic activity which causes important fluctuations of the membrane potential (Vm). We review here different methods to characterize this activity and its impact on spike generation. The simplified, fluctuating point-conductance model of synaptic activity provides the starting point of a variety of methods for the analysis of intracellular Vm recordings. In this model, the synaptic excitatory and inhibitory conductances are described by Gaussian-distributed stochastic variables, or colored conductance noise. The matching of experimental…
Correlated electrical activity in neurons is a prominent characteristic of cortical microcircuits. Despite a growing amount of evidence concerning both spike-count and subthreshold membrane potential pairwise correlations, little is known about how different types of cortical neurons convert correlated inputs into correlated outputs. We studied pyramidal neurons and two classes of GABAergic interneurons of layer 5 in neocortical brain slices obtained from rats of both sexes, and we stimulated them with biophysically realistic correlated inputs, generated using dynamic clamp. We found that the …
Objective: Hepatic encephalopathy (HE) is a potentially reversible brain dysfunction caused by liver failure. Altered synaptic plasticity is supposed to play a major role in the pathophysiology of HE. Here, we used paired associative stimulation with an inter-stimulus interval of 25 ms (PAS25), a transcranial magnetic stimulation (TMS) protocol, to test synaptic plasticity of the motor cortex in patients with manifest HE. Methods: 23 HE-patients and 23 healthy controls were enrolled in the study. Motor evoked potential (MEP) amplitudes were assessed as measure for cortical excitability. Time c…
Cortical populations of neurons develop sparse representations adapted to the statistics of the environment. While existing synaptic plasticity models reproduce some of the observed receptive-field properties, a major obstacle is the sensitivity of Hebbian learning to omnipresent spurious correlations in cortical networks which can overshadow relevant latent input features. Here we develop a theory for synaptic plasticity that is invariant to second-order correlations in the input. Going beyond classical Hebbian learning, we show how Hebbian long-term depression (LTD) cancels the sensitivity t…
When a neuron fires and the resulting action potential travels down its axon toward other neurons' dendrites, the effect on each of those neurons is mediated by the weight of the synapse that separates it from the firing neuron. This weight, in turn, is affected by the postsynaptic neuron's response through a mechanism that is thought to underlie important processes such as learning and memory. Although of difficult quantification, cortical synaptic weights have been found to obey a long-tailed unimodal distribution peaking near the lowest values, thus confirming some of the predictive models …
Two recent experimental observations pose a challenge to many cortical models. First, the activity in the auditory cortex is sparse, and firing rates can be described by a lognormal distribution. Second, the distribution of non-zero synaptic strengths between nearby cortical neurons can also be described by a lognormal distribution. Here we use a simple model of cortical activity to reconcile these observations. The model makes the experimentally testable prediction that synaptic efficacies onto a given cortical neuron are statistically correlated, i.e. it predicts that some neurons receive ma…
In the study of biological networks, one of the major challenges is to understand the relationships between network structure and dynamics. In this paper, we model in vitro cortical neuronal cultures as stochastic dynamical systems and apply a method that reconstructs directed networks from dynamics [Ching and Tam, Phys. Rev. E 95, 010301(R), 2017] to reveal directed effective connectivity, namely the directed links and synaptic weights, of the neuronal cultures from voltage measurements recorded by a multielectrode array. The effective connectivity so obtained reproduces several features of c…
We consider the theoretical constraints on interactions between coupled cortical columns. Each column comprises a set of neural populations, where each population is modelled as a neural mass. The existence of semi-stable states within a cortical column has been shown to be dependent on the type of interaction between the constituent neuronal subpopulations, i.e., the form of the implicit synaptic convolution kernels. Current-to-current coupling has been shown, in contrast to potential-to-current coupling, to create semi-stable states within a cortical column. In this analytic and numerical st…
Our current understanding of neuroplasticity paints a picture of a complex interconnected system of dependent processes which shape cortical structure so as to produce an efficient information processing system. Indeed, the cooperation of these processes is associated with robust, stable, adaptable networks with characteristic features of activity and synaptic topology. However, combining the actions of these mechanisms in models has proven exceptionally difficult and to date no model has been able to do so without significant hand-tuning. Until such a model exists that can successfully combin…
Cell assemblies manipulation by optogenetics is pivotal to advance neuroscience and neuroengineering. In in vivo applications, photostimulation often broadly addresses a population of cells simultaneously, leading to feed-forward and to reverberating responses in recurrent microcircuits. The former arise from direct activation of targets downstream, and are straightforward to interpret. The latter are consequence of feedback connectivity and may reflect a variety of time-scales and complex dynamical properties. We investigated wide-field photostimulation in cortical networks in vitro, employin…
Spontaneous cortical population activity exhibits a multitude of oscillatory patterns, which often display synchrony during slow-wave sleep or under certain anesthetics and stay asynchronous during quiet wakefulness. The mechanisms behind these cortical states and transitions among them are not completely understood. Here we study spontaneous population activity patterns in random networks of spiking neurons of mixed types modeled by Izhikevich equations. Neurons are coupled by conductance-based synapses subject to synaptic noise. We localize the population activity patterns on the parameter d…
Graphene has the potential to make a very significant impact on society, with important applications in the biomedical field. The possibility to engineer graphene-based medical devices at the neuronal interface is of particular interest, making it imperative to determine the biocompatibility of graphene materials with neuronal cells. Here we conducted a comprehensive analysis of the effects of chronic and acute exposure of rat primary cortical neurons to few-layers pristine graphene (GR) and monolayer graphene oxide (GO) flakes. By combining a range of cell biology, microscopy, electrophysiolo…
The synaptic connectivity of cortical networks features an overrepresentation of certain wiring motifs compared to simple random-network models. This structure is shaped, in part, by synaptic plasticity that promotes or suppresses connections between neurons depending on their spiking activity. Frequently, theoretical studies focus on how feedforward inputs drive plasticity to create this network structure. We study the complementary scenario of self-organized structure in a recurrent network, with spike timing-dependent plasticity driven by spontaneous dynamics. We develop a self-consistent t…
INFN Sezione di Roma（意大利国家核物理研究所罗马分所）；INFN Sezione di Cagliari（意大利国家核物理研究所卡利亚里分所）；Università di Cagliari（卡利亚里大学）
The occurrence of sleep passed through the evolutionary sieve and is widespread in animal species. Sleep is known to be beneficial to cognitive and mnemonic tasks, while chronic sleep deprivation is detrimental. Despite the importance of the phenomenon, a complete understanding of its functions and underlying mechanisms is still lacking. In this paper, we show interesting effects of deep-sleep-like slow oscillation activity on a simplified thalamo-cortical model which is trained to encode, retrieve and classify images of handwritten digits. During slow oscillations, spike-timing-dependent-plas…
We investigate cortical learning from the perspective of mechanism design. First, we show that discretizing standard models of neurons and synaptic plasticity leads to rational agents maximizing simple scoring rules. Second, our main result is that the scoring rules are proper, implying that neurons faithfully encode expected utilities in their synaptic weights and encode high-scoring outcomes in their spikes. Third, with this foundation in hand, we propose a biologically plausible mechanism whereby neurons backpropagate incentives which allows them to optimize their usefulness to the rest of …
Complex coherent dynamics is present in a wide variety of neural systems. A typical example is the voltage transitions between up and down states observed in cortical areas in the brain. In this work, we study this phenomenon via a biologically motivated stochastic model of up and down transitions. The model is constituted by a simple bistable rate model, where the synaptic current is modulated by short-term synaptic processes which introduce stochasticity and temporal correlations. A complete analysis of our model, both with mean-field approaches and numerical simulations, shows the appearanc…
In this report trial-to-trial variations in the synchronized responses of neural networks are offered as evidence for excitation-inhibition ratio being a dynamic variable over time scales of minutes. Synchronized network responses to stimuli were studied in ex-vivo large scale cortical networks. We show that sub-second measures of the individual synchronous response, namely -- its latency and decay duration, are related to minutes-scale network response dynamics. Network responsiveness is reflected as residency in, or shifting amongst, areas of the latency-decay plane. The different sensitivit…
We study the spike statistics of neurons in a network with dynamically balanced excitation and inhibition. Our model, intended to represent a generic cortical column, comprises randomly connected excitatory and inhibitory leaky integrate-and-fire neurons, driven by excitatory input from an external population. The high connectivity permits a mean-field description in which synaptic currents can be treated as Gaussian noise, the mean and autocorrelation function of which are calculated self-consistently from the firing statistics of single model neurons. Within this description, we find that th…
Excitatory synaptic connections in the adult neocortex consist of multiple synaptic contacts, almost exclusively formed on dendritic spines. Changes of dendritic spine shape and volume, a correlate of synaptic strength, can be tracked in vivo for weeks. Here, we present a combined model of spike-timing dependent dendritic spine plasticity and turnover that explains the steady state multi-contact configuration of synapses in adult neocortical networks. In this model, many presynaptic neurons compete to make strong synaptic connections onto postsynaptic neurons, while the synaptic contacts compr…
We present an effective model for timing-dependent synaptic plasticity (STDP) in terms of two interacting traces, corresponding to the fraction of activated NMDA receptors and the Ca2+ concentration in the dendritic spine of the postsynaptic neuron. This model intends to bridge the worlds of existing simplistic phenomenological rules and highly detailed models, constituting thus a practical tool for the study of the interplay between neural activity and synaptic plasticity in extended spiking neural networks. For isolated pairs of pre- and postsynaptic spikes the standard pairwise STDP rule is…
Department of Physiology University of Bern（伯尔尼大学生理学系）；Montreal Institute for Learning Algorithms Université de Montréal（蒙特利尔大学蒙特利尔学习算法研究所）
Animal behaviour depends on learning to associate sensory stimuli with the desired motor command. Understanding how the brain orchestrates the necessary synaptic modifications across different brain areas has remained a longstanding puzzle. Here, we introduce a multi-area neuronal network model in which synaptic plasticity continuously adapts the network towards a global desired output. In this model synaptic learning is driven by a local dendritic prediction error that arises from a failure to predict the top-down input given the bottom-up activities. Such errors occur at apical dendrites of …
Cortical synapse organization supports a range of dynamic states on multiple spatial and temporal scales, from synchronous slow wave activity (SWA), characteristic of deep sleep or anesthesia, to fluctuating, asynchronous activity during wakefulness (AW). Such dynamic diversity poses a challenge for producing efficient large-scale simulations that embody realistic metaphors of short- and long-range synaptic connectivity. In fact, during SWA and AW different spatial extents of the cortical tissue are active in a given timespan and at different firing rates, which implies a wide variety of loads…
Fast cortical rhythms with stochastic and intermittent neural discharges have been observed in electric recordings of brain activity. Recently, Brunel et al. developed a framework to describe this kind of fast sparse synchronization in both random and globally-coupled networks of suprathreshold spiking neurons. However, in a real cortical circuit, synaptic connections are known to have complex topology which is neither regular nor random. Hence, in order to extend the works of Brunel et al. to realistic neural networks, we study the effect of network architecture on these fast sparsely synchro…
Sparse coding algorithms trained on natural images can accurately predict the features that excite visual cortical neurons, but it is not known whether such codes can be learned using biologically realistic plasticity rules. We have developed a biophysically motivated spiking network, relying solely on synaptically local information, that can predict the full diversity of V1 simple cell receptive field shapes when trained on natural images. This represents the first demonstration that sparse coding principles, operating within the constraints imposed by cortical architecture, can successfully …
Coarse-graining microscopic models of biological neural networks to obtain mesoscopic models of neural activities is an essential step towards multi-scale models of the brain. Here, we extend a recent theory for mesoscopic population dynamics with static synapses to the case of dynamic synapses exhibiting short-term plasticity (STP). Under the assumption that spike arrivals at synapses have Poisson statistics, we derive analytically stochastic mean-field dynamics for the effective synaptic coupling between finite-size populations undergoing Tsodyks-Markram STP. The novel mean-field equations a…
Institute of Neuroinformatics（神经信息学研究所）；University of Zürich（苏黎世大学）；ETH Zürich（苏黎世联邦理工学院）；University of Bern（伯尔尼大学）；University of Bristol（布里斯托大学）；Mila（米拉研究所）；Université de Montréal（蒙特利尔大学）
Deep learning has seen remarkable developments over the last years, many of them inspired by neuroscience. However, the main learning mechanism behind these advances - error backpropagation - appears to be at odds with neurobiology. Here, we introduce a multilayer neuronal network model with simplified dendritic compartments in which error-driven synaptic plasticity adapts the network towards a global desired output. In contrast to previous work our model does not require separate phases and synaptic learning is driven by local dendritic prediction errors continuously in time. Such errors orig…
The dynamics of local cortical networks are irregular, but correlated. Dynamic excitatory--inhibitory balance is a plausible mechanism that generates such irregular activity, but it remains unclear how balance is achieved and maintained in plastic neural networks. In particular, it is not fully understood how plasticity induced changes in the network affect balance, and in turn, how correlated, balanced activity impacts learning. How does the dynamics of balanced networks change under different plasticity rules? How does correlated spiking activity in recurrent networks change the evolution of…
An autoassociative network of Potts units, coupled via tensor connections, has been proposed and analysed as an effective model of an extensive cortical network with distinct short- and long-range synaptic connections, but it has not been clarified in what sense it can be regarded as an effective model. We draw here the correspondence between the two, which indicates the need to introduce a local feedback term in the reduced model, i.e., in the Potts network. An effective model allows the study of phase transitions. As an example, we study the storage capacity of the Potts network with this ad…
The spectacular successes of recurrent neural network models where key parameters are adjusted via backpropagation-based gradient descent have inspired much thought as to how biological neuronal networks might solve the corresponding synaptic credit assignment problem. There is so far little agreement, however, as to how biological networks could implement the necessary backpropagation through time, given widely recognized constraints of biological synaptic network signaling architectures. Here, we propose that extra-synaptic diffusion of local neuromodulators such as neuropeptides may afford …
The representation of the natural-density, heterogeneous connectivity of neuronal network models at relevant spatial scales remains a challenge for Computational Neuroscience and Neuromorphic Computing. In particular, the memory demands imposed by the vast number of synapses in brain-scale network simulations constitutes a major obstacle. Limiting the number resolution of synaptic weights appears to be a natural strategy to reduce memory and compute load. In this study, we investigate the effects of a limited synaptic-weight resolution on the dynamics of recurrent spiking neuronal networks res…
When brain signals are recorded in an electroencephalogram or some similar large-scale record of brain activity, oscillatory patterns are typically observed that are thought to reflect the aggregate electrical activity of the underlying neuronal ensemble. Although it now seems that such patterns participate in feedback loops both temporally with the neurons' spikes and spatially with other brain regions, the mechanisms that might explain the existence of such loops have remained essentially unknown. Here we present a theoretical study of these issues on a cortical model we introduced earlier […
Cortical pyramidal neurons receive inputs from multiple distinct neural populations and integrate these inputs in separate dendritic compartments. We explore the possibility that cortical microcircuits implement Canonical Correlation Analysis (CCA), an unsupervised learning method that projects the inputs onto a common subspace so as to maximize the correlations between the projections. To this end, we seek a multi-channel CCA algorithm that can be implemented in a biologically plausible neural network. For biological plausibility, we require that the network operates in the online setting and…
Darwinian evolution tends to produce energy-efficient outcomes. On the other hand, energy limits computation, be it neural and probabilistic or digital and logical. Taking a particular energy-efficient viewpoint, we define neural computation and make use of an energy-constrained, computational function. This function can be optimized over a variable that is proportional to the number of synapses per neuron. This function also implies a specific distinction between ATP-consuming processes, especially computation \textit{per se} vs the communication processes including action potentials and tran…
The free energy principle (FEP) is a mathematical framework that describes how biological systems self-organize and survive in their environment. This principle provides insights on multiple scales, from high-level behavioral and cognitive functions such as attention or foraging, down to the dynamics of specialized cortical microcircuits, suggesting that the FEP manifests on several levels of brain function. Here, we apply the FEP to one of the smallest functional units of the brain: single excitatory synaptic connections. By focusing on an experimentally well understood biological system we a…
It is currently accepted that cortical maps are dynamic constructions that are altered in response to external input. Experience-dependent structural changes in cortical microcurcuts lead to changes of activity, i.e. to changes in information encoded. Specific patterns of external stimulation can lead to creation of new synaptic connections between neurons. The calcium influxes controlled by neuronal activity regulate the processes of neurotrophic factors released by neurons, growth cones movement and synapse differentiation in developing neural systems. We propose a model for description and …
Synaptic delays modulate population phase and amplitude responses in oscillatory excitatory-inhibitory networks
Institute for Advanced Studies in Basic Sciences (IASBS)（基础科学高等研究院）；Institute for Research in Fundamental Sciences (IPM)（基本科学研究院）；Pasargad Institute for Advanced Innovative Solutions (PIAIS)（帕萨尔加德先进创新解决方案研究所）；The Zapata-Briceño Institute of Neuroscience（萨帕塔-布里斯尼奥神经科学研究所）
Short-Term Synaptic Plasticity (STSP) strongly affects the neural dynamics of cortical networks. The Tsodyks and Markram (TM) model for STSP accurately accounts for a wide range of physiological responses at different types of cortical synapses. Here, we report for the first time a route to chaotic behavior via a Shilnikov homoclinic bifurcation that dynamically organises some of the responses in the TM model. In particular, the presence of such a homoclinic bifurcation strongly affects the shape of the trajectories in the phase space and induces highly irregular transient dynamics; indeed, in…
Cortical networks can maintain memories for decades despite the short lifetime of synaptic strength. Can a neural network store long-lasting memories in unstable synapses? Here, we study the effects of random noise on the stability of memory stored in synapses of an attractor neural network. The model includes ongoing spike timing dependent plasticity (STDP). We show that certain classes of STDP rules can lead to the stabilization of memory patterns stored in the network. The stabilization results from rehearsals induced by noise. We show that unstructured neural noise, after passing through t…
All higher order central nervous systems exhibit spontaneous neural activity, though the purpose and mechanistic origin of such activity remains poorly understood. We explore the ignition and spread of collective spontaneous electrophysiological burst activity in networks of cultured cortical neurons growing on microelectrode arrays using information theory and first-spike-in-burst analysis methods. We show the presence of burst leader neurons, which form a mono-synaptically connected primary circuit, and initiate a majority of network bursts. Leader/follower firing delay times form temporally…
Background: The roles of neuromodulation in a neural network, such as in a cortical microcolumn, are still incompletely understood. Neuromodulation influences neural processing by presynaptic and postsynaptic regulation of synaptic efficacy. Neuromodulation also affects ion channels and intrinsic excitability. Methods: Synaptic efficacy modulation is an effective way to rapidly alter network density and topology. We alter network topology and density to measure the effect on spike synchronization. We also operate with differently parameterized neuron models which alter the neurons intrinsic ex…
The cortex learns to make associations between stimuli and spiking activity which supports behaviour. It does this by adjusting synaptic weights. The complexity of these transformations implies that synapses have to change without access to the full error information, a problem typically referred to as "credit-assignment". However, it remains unknown how the cortex solves this problem. We propose that a combination of plasticity rules, 1) Hebbian, 2) acetylcholine-dependent and 3) noradrenaline-dependent excitatory plasticity, together with 4) inhibitory plasticity restoring E/I balance, effec…
IBM Research – Zurich（IBM苏黎世研究院）；Huawei – Zurich Research Center（华为苏黎世研究中心）
Biological neurons and their in-silico emulations for neuromorphic artificial intelligence (AI) use extraordinarily energy-efficient mechanisms, such as spike-based communication and local synaptic plasticity. It remains unclear whether these neuronal mechanisms only offer efficiency or also underlie the superiority of biological intelligence. Here, we prove rigorously that, indeed, the Bayes-optimal prediction and inference of randomly but continuously transforming environments, a common natural setting, relies on short-term spike-timing-dependent plasticity, a hallmark of biological synapses…
Behavioral control over stressful stimuli induces resilience to future conditions when control is lacking. The medial prefrontal cortex(mPFC) is a critically important brain region required for plasticity of stress resilience. We found that control over stress induces plasticity of the intrinsic voltage-gated conductances of pyramidal neurons in the PFC. To gain insight into the underlying biophysical mechanisms of this plasticity we used the conductance- based neural simulation software tool, NEURON, to model the increase in membrane excitability associated with resilience to stress. A ball a…
Sparse coding networks, which utilize unsupervised learning to maximize coding efficiency, have successfully reproduced response properties found in primary visual cortex \cite{AN:OlshausenField96}. However, conventional sparse coding models require that the coding circuit can fully sample the sensory data in a one-to-one fashion, a requirement not supported by experimental data from the thalamo-cortical projection. To relieve these strict wiring requirements, we propose a sparse coding network constructed by introducing synaptic learning in the framework of compressed sensing. We demonstrate …
General results from statistical learning theory suggest to understand not only brain computations, but also brain plasticity as probabilistic inference. But a model for that has been missing. We propose that inherently stochastic features of synaptic plasticity and spine motility enable cortical networks of neurons to carry out probabilistic inference by sampling from a posterior distribution of network configurations. This model provides a viable alternative to existing models that propose convergence of parameters to maximum likelihood values. It explains how priors on weight distributions …
Transcranial Electrical Stimulation (TCES) and Deep Brain Stimulation (DBS) are two different applications of electrical current to the brain used in different areas of medicine. Both have a similar frequency dependence of their efficiency, with the most pronounced effects around 100Hz. We apply superthreshold electrical stimulation, specifically depolarizing DC current, interrupted at different frequencies, to a simple model of a population of cortical neurons which uses phenomenological descriptions of neurons by Izhikevich and synaptic connections on a similar level of sophistication. With …
Dendritic spines, the carriers of long-term memory, occupy a small fraction of cortical space, and yet they are the major consumers of brain metabolic energy. What fraction of this energy goes for synaptic plasticity, correlated with learning and memory? It is estimated here based on neurophysiological and proteomic data for rat brain that, depending on the level of protein phosphorylation, the energy cost of synaptic plasticity constitutes a small fraction of the energy used for fast excitatory synaptic transmission, typically $4.0-11.2 \%$. Next, this study analyzes a metabolic cost of a new…
In network models of spiking neurons, the joint impact of network structure and synaptic parameters on activity propagation is still an open problem. Here we use an information-theoretical approach to investigate activity propagation in spiking networks with hierarchical modular topology. We observe that optimized pairwise information propagation emerges due to the increase of either (i) the global synaptic strength parameter or (ii) the number of modules in the network, while the network size remains constant. At the population level, information propagation of activity among adjacent modules…
Most cortical inhibitory cell types exclusively express one of three genes, parvalbumin, somatostatin and 5HT3a. The visual responses of cortical neurons are affected not only by local cues, but also by visual context. As the inhibitory neuron types have distinctive synaptic sources and targets over different spatial extents and from different areas, we conjecture that they possess distinct roles in contextual processing. We use modeling to relate structural information to function in primary visual cortex (V1) of the mouse, and investigate their role in contextual visual processing. Our findi…
Alzheimer's disease (AD) is a common form of dementia observed in the elderly due to neurodegenerative disorder and dysfunction. This arises from alterations in synaptic functioning of neurons leading to cognitive impairment and memory loss. Recent experimental studies indicate that the amyloid beta (A beta) protein, in its dimer and oligomer forms, affects the synaptic activity of neurons in the early stages of AD. However, the precise mechanism underlying A beta induced synaptic depression is still not clearly understood. In this paper, we introduce an electrical model that provides a possib…
Cortical sensory neurons are known to be highly variable, in the sense that responses evoked by identical stimuli often change dramatically from trial to trial. The origin of this variability is uncertain, but it is usually interpreted as detrimental noise that reduces the computational accuracy of neural circuits. Here we investigate the possibility that such response variability might, in fact, be beneficial, because it may partially compensate for a decrease in accuracy due to stochastic changes in the synaptic strengths of a network. We study the interplay between two kinds of noise, respo…
