In this paper, we systematically investigate both the synfire propagation and firing rate propagation in feedforward neuronal network coupled in an all-to-all fashion. In contrast to most earlier work, where only reliable synaptic connections are considered, we mainly examine the effects of unreliable synapses on both types of neural activity propagation in this work. We first study networks composed of purely excitatory neurons. Our results show that both the successful transmission probability and excitatory synaptic strength largely influence the propagation of these two types of neural activities, and better tuning of these synaptic parameters makes the considered network support stable signal propagation. It is also found that noise has significant but different impacts on these two types of propagation. The additive Gaussian white noise has the tendency to reduce the precision of the synfire activity, whereas noise with appropriate intensity can enhance the performance of firing rate propagation. Further simulations indicate that the propagation dynamics of the considered neuronal network is not simply determined by the average amount of received neurotransmitter for each neuron in a time instant, but also largely influenced by the stochastic effect of neurotransmitter release. Second, we compare our results with those obtained in corresponding feedforward neuronal networks connected with reliable synapses but in a random coupling fashion. We confirm that some differences can be observed in these two different feedforward neuronal network models. Finally, we study the signal propagation in feedforward neuronal networks consisting of both excitatory and inhibitory neurons, and demonstrate that inhibition also plays an important role in signal propagation in the considered networks.
The characterisation of neuronal connectivity is one of the most important matters in neuroscience. In this work, we show that a recently proposed informational quantity, the causal mutual information, employed with an appropriate methodology, can be used not only to correctly infer the direction of the underlying physical synapses, but also to identify their excitatory or inhibitory nature, considering easy to handle and measure bivariate time-series. The success of our approach relies on a surprising property found in neuronal networks by which non-adjacent neurons do "understand" each other…
Many experiments have evidenced that electrical and chemical synapses -- hybrid synapses -- coexist in most organisms and brain structures. The role of electrical and chemical synapse connection in diversity of neural activity generation has been investigated separately in networks of varying complexities. Nevertheless, theoretical understanding of hybrid synapses in diverse dynamical states of neural networks for self-organization and robustness still has not been fully studied. Here, we present a model of neural network built with hybrid synapses to investigate the emergence of global and co…
Brain plasticity, also known as neuroplasticity, is a fundamental mechanism of neuronal adaptation in response to changes in the environment or due to brain injury. In this review, we show our results about the effects of synaptic plasticity on neuronal networks composed by Hodgkin-Huxley neurons. We show that the final topology of the evolved network depends crucially on the ratio between the strengths of the inhibitory and excitatory synapses. Excitation of the same order of inhibition revels an evolved network that presents the rich-club phenomenon, well known to exist in the brain. For ini…
A recurrent loop consisting of a single neuron is considered which is influenced by a chemical excitatory delayed synaptic feedback. We show the response of the system is dependent to the duration of the activity of the synapse which is determined by the deactivation time constant of the synapse. We show that loops with slow synapses, those which the effect of the synaptic activation remains for time constants comparable to the period of firing, show more predictable results where the effect of the fast synapses is tightly dependent on the loop delay time. The results are compared to those of …
Graph theoretical analyses of nervous systems usually omit the aspect of connection polarity, due to data insufficiency. The chemical synapse network of Caenorhabditis elegans is a well-reconstructed directed network, but the signs of its connections are yet to be elucidated. Here, we present the gene expression-based sign prediction of the ionotropic chemical synapse connectome of C. elegans (3,638 connections and 20,589 synapses total), incorporating available presynaptic neurotransmitter and postsynaptic receptor gene expression data for three major neurotransmitter systems. We made predict…
Conventionally, information is represented by spike rates in the neural system. Here, we consider the ability of temporally modulated activities in neuronal networks to carry information extra to spike rates. These temporal modulations, commonly known as population spikes, are due to the presence of synaptic depression in a neuronal network model. We discuss its relevance to an experiment on transparent motions in macaque monkeys by Treue et al. in 2000. They found that if the moving directions of objects are too close, the firing rate profile will be very similar to that with one direction. A…
Biocomputing technologies exploit biological communication mechanisms involving cell-cell signal propagation to perform computations. Researchers recently worked toward realising logic gates made by neurons to develop novel devices such as organic neuroprostheses or brain implants made by cells, herein termed living implants. Several challenges arise from this approach, mainly associated with the stochastic nature and noise of neuronal communication. Since astrocytes play a crucial role in the regulation of neurons activity, there is a possibility whereby astrocytes can be engineered to contro…
In this work, we study the dynamic range in a neuronal network modelled by cellular automaton. We consider deterministic and non-deterministic rules to simulate electrical and chemical synapses. Chemical synapses have an intrinsic time-delay and are susceptible to parameter variations guided by learning Hebbian rules of behaviour. Our results show that chemical synapses can abruptly enhance sensibility of the neural network, a manifestation that can become even more predominant if learning rules of evolution are applied to the chemical synapses.
In this work, we study the dynamic range of a neuronal network of excitable neurons with excitatory and inhibitory synapses. We obtain an analytical expression for the critical point as a function of the excitatory and inhibitory synaptic intensities. We also determine an analytical expression that gives the critical point value in which the maximal dynamic range occurs. Depending on the mean connection degree and coupl\-ing weights, the critical points can exhibit ceasing or ceaseless dynamics. However, the dynamic range is equal in both cases. We observe that the external stimulus mask some …
Information transmission is a key element for information processing in the brain. A number of mechanisms have been proposed for transferring volleys of spikes between layers of a feedforward neural circuit. Many of these mechanisms use synchronous activity to provide windows in time when spikes may be transferred more easily from layer to layer. Recently, we have demonstrated that a pulse-gating mechanism can transfer graded information between layers in a feedforward neuronal network. Our transfer mechanism resulted in a time-translationally invariant firing rate and synaptic current wavefor…
Chemical and electrical synapses shape the dynamics of neuronal networks. Numerous theoretical studies have investigated how each of these types of synapses contributes to the generation of neuronal oscillations, but their combined effect is less understood. This limitation is further magnified by the impossibility of traditional neuronal mean-field models ---also known as firing rate models, or firing rate equations--- to account for electrical synapses. Here we introduce a novel firing rate model that exactly describes the mean field dynamics of heterogeneous populations of quadratic integra…
Neural coding is a key problem in neuroscience, which can promote people's understanding of the mechanism that brain processes information. Among the classical theories of neural coding, the population rate coding has been studied widely in many works. Most computational studies considered the neurons and the corresponding presynaptic synapses as pre-determined excitatory or inhibitory types. According to physiological evidence, however, that the real effect of a synapse is inhibitory or excitatory is determined by the type of the activated receptors. The co-release of excitatory and inhibitor…
Avalanches with power-law distributed size parameters have been observed in neuronal networks. This observation might be a manifestation of the self-organized criticality (SOC). Yet, the physiological mechanicsm of this behavior is currently unknown. Describing synaptic noise as transmission failures mainly originating from the probabilistic nature of neurotransmitter release, this study investigates the potential of this noise as a mechanism for driving the functional architecture of the neuronal networks towards SOC. To this end, a simple finite state neuron model, with activity dependent an…
Feedforward networks (FFN) are ubiquitous structures in neural systems and have been studied to understand mechanisms of reliable signal and information transmission. In many FFNs, neurons in one layer have intrinsic properties that are distinct from those in their pre-/postsynaptic layers, but how this affects network-level information processing remains unexplored. Here we show that layer-to-layer heterogeneity arising from lamina-specific cellular properties facilitates signal and information transmission in FFNs. Specifically, we found that signal transformations, made by each layer of neu…
Despite recent interest in reconstructing neuronal networks, complete wiring diagrams on the level of individual synapses remain scarce and the insights into function they can provide remain unclear. Even for Caenorhabditis elegans, whose neuronal network is relatively small and stereotypical from animal to animal, published wiring diagrams are neither accurate nor complete and self-consistent. Using materials from White et al. and new electron micrographs we assemble whole, self-consistent gap junction and chemical synapse networks of hermaphrodite C. elegans. We propose a method to visualize…
Centre for Theoretical Neuroscience, University of Waterloo（滑铁卢大学理论神经科学中心）
The mathematical model underlying the Neural Engineering Framework (NEF) expresses neuronal input as a linear combination of synaptic currents. However, in biology, synapses are not perfect current sources and are thus nonlinear. Detailed synapse models are based on channel conductances instead of currents, which require independent handling of excitatory and inhibitory synapses. This, in particular, significantly affects the influence of inhibitory signals on the neuronal dynamics. In this technical report we first summarize the relevant portions of the NEF and conductance-based synapse model…
Molecule-based devices are envisioned to complement silicon devices by providing new functions or already existing functions at a simpler process level and at a lower cost by virtue of their self-organization capabilities, moreover, they are not bound to von Neuman architecture and this may open the way to other architectural paradigms. Here we demonstrate a device made of conjugated molecules and metal nanoparticles (NPs) which behaves as a spiking synapse suitable for integration in neural network architectures. We demonstrate that this device exhibits the main behavior of a biological synap…
Cross-correlations in the activity in neural networks are commonly used to characterize their dynamical states and their anatomical and functional organizations. Yet, how these latter network features affect the spatiotemporal structure of the correlations in recurrent networks is not fully understood. Here, we develop a general theory for the emergence of correlated neuronal activity from the dynamics in strongly recurrent networks consisting of several populations of binary neurons. We apply this theory to the case in which the connectivity depends on the anatomical or functional distance be…
Brain plasticity refers to brain's ability to change neuronal connections, as a result of environmental stimuli, new experiences, or damage. In this work, we study the effects of the synaptic delay on both the coupling strengths and synchronisation in a neuronal network with synaptic plasticity. We build a network of Hodgkin-Huxley neurons, where the plasticity is given by the Hebbian rules. We verify that without time delay the excitatory synapses became stronger from the high frequency to low frequency neurons and the inhibitory synapses increases in the opposite way, when the delay is incre…
Recent experimental studies strongly suggest the influence of glial purinergic transmission in the modulation of synaptic dynamics. By releasing adenosine triphosphate (ATP), which accumulates as adenosine, astrocytes tonically suppressed synaptic transmission. The delayed multi-step feedback of the glial adenosine with the neuron suggest the existence of hysteresis phenomena, which are investigated in the present study from the theoretical point of view. The model suggests that a memory operator, tripartite synaptic plasticity, governs the mysterious delayed feedback inhibition caused by the …
We study the dynamic range of a cellular automaton model for a neuronal network with electrical and chemical synapses. The neural network is separated into two layers, where one layer corresponds to inhibitory, and the other corresponds to excitatory neurons. We randomly distribute electrical synapses in the network, in order to analyse the effects on the dynamic range. We verify that electrical synapses have a complementary effect on the enhancement of the dynamic range. The enhancement depends on the proportion of electrical synapses, and also depends on the layer where they are distributed.
Chimera states are spatiotemporal patterns in which coherence and incoherence coexist. We observe the coexistence of synchronous (coherent) and desynchronous (incoherent) domains in a neuronal network. The network is composed of coupled adaptive exponential integrate-and-fire neurons that are connected by means of chemical synapses. In our neuronal network, the chimera states exhibit spatial structures both with spikes and bursts activities. Furthermore, those desynchronised domains not only have either spike or burst activity, but we show that the structures switch between spikes and bursts a…
Early brain connectivity development consists of multiple stages: birth of neurons, their migration and the subsequent growth of axons and dendrites. Each stage occurs within a certain period of time depending on types of neurons and cortical layers. Forming synapses between neurons either by growing axons starting at similar times for all neurons (much-overlapped time windows) or at different time points (less-overlapped) may affect the topological and spatial properties of neuronal networks. Here, we explore the extreme cases of axon formation especially concerning short-distance connectivit…
Network architecture forms a critical constraint on neuronal function. Here we examine the role of structural autapses, when a neuron synapses onto itself, in driving network-wide bursting behavior. Using a simple spiking model of neuronal activity, we study how autaptic connections affect activity patterns, and evaluate if neuronal degree or controllability are significant factors that affect changes in bursting from these autaptic connections. We observed that adding increasing numbers of autaptic connections to excitatory neurons increased the number of spiking events in the network and the…
The $1/f$-like decay observed in the power spectrum of electro-physiological signals, along with scale-free statistics of the so-called neuronal avalanches, constitute evidences of criticality in neuronal systems. Recent in vitro studies have shown that avalanche dynamics at criticality corresponds to some specific balance of excitation and inhibition, thus suggesting that this is a basic feature of the critical state of neuronal networks. In particular, a lack of inhibition significantly alters the temporal structure of the spontaneous avalanche activity and leads to an anomalous abundance of…
Reconstructing neuronal circuits at the level of synapses is a central problem in neuroscience and becoming a focus of the emerging field of connectomics. To date, electron microscopy (EM) is the most proven technique for identifying and quantifying synaptic connections. As advances in EM make acquiring larger datasets possible, subsequent manual synapse identification ({\em i.e.}, proofreading) for deciphering a connectome becomes a major time bottleneck. Here we introduce a large-scale, high-throughput, and semi-automated methodology to efficiently identify synapses. We successfully applied …
Neurons are spatially extended structures that receive and process inputs on their dendrites. It is generally accepted that neuronal computations arise from the active integration of synaptic inputs along a dendrite between the input location and the location of spike generation in the axon initial segment. However, many application such as simulations of brain networks, use point-neurons --neurons without a morphological component-- as computational units to keep the conceptual complexity and computational costs low. Inevitably, these applications thus omit a fundamental property of neuronal …
Neuroprosthesis, as one type of precision medicine device, is aiming for manipulating neuronal signals of the brain in a closed-loop fashion, together with receiving stimulus from the environment and controlling some part of our brain/body. In terms of vision, incoming information can be processed by the brain in millisecond interval. The retina computes visual scenes and then sends its output as neuronal spikes to the cortex for further computation. Therefore, the neuronal signal of interest for retinal neuroprosthesis is spike. Closed-loop computation in neuroprosthesis includes two stages: …
When handling real-world data modeled by a complex network dynamical system, the number of the parameters is always even much more than the size of the data. Therefore, in many cases, it is impossible to estimate these parameters and however, the exact value of each parameter is frequently less interesting than the distribution of the parameters may contain important information towards understanding the system and data. In this paper, we propose this question arising by employing a data assimilation approach to estimate the distribution of the parameters in the leakage-integrate-fire (LIF) ne…
Several studies with brain signals suggested that bottom-up and top-down influences are exerted through distinct frequency bands among visual cortical areas. It has been recently shown that theta and gamma rhythms subserve feedforward, whereas the feedback influence is dominated by the alpha-beta rhythm in primates. A few theoretical models for reproducing these effects have been proposed so far. Here we show that a simple but biophysically plausible two-network motif composed of spiking-neuron models and chemical synapses can exhibit feedforward and feedback influences through distinct freque…
We study the dynamics of the structure of a formal neural network wherein the strengths of the synapses are governed by spike-timing-dependent plasticity (STDP). For properly chosen input signals, there exists a steady state with a residual network. We compare the motif profile of such a network with that of a real neural network of \emph{C. elegans} and identify robust qualitative similarities. In particular, our extensive numerical simulations show that this STDP-driven resulting network is robust under variations of the model parameters.
We study the time delay in the synaptic conductance for suppression of spike synchronisation in a random network of Hodgkin Huxley neurons coupled by means of chemical synapses. In the first part, we examine in detail how the time delay acts over the network during the synchronised and desynchronised neuronal activities. We observe a relation between the neuronal dynamics and the syaptic conductance distributions. We find parameter values in which the time delay has high effectiveness in promoting the suppression of spike synchronisation. In the second part, we analyse how the delayed neuronal…
We investigate the effect of electric synapses (gap junctions) on collective neuronal dynamics and spike statistics in a conductance-based Integrate-and-Fire neural network, driven by a Brownian noise, where conductances depend upon spike history. We compute explicitly the time evolution operator and show that, given the spike-history of the network and the membrane potentials at a given time, the further dynamical evolution can be written in a closed form. We show that spike train statistics is described by a Gibbs distribution whose potential can be approximated with an explicit formula, whe…
We investigate the synaptic noise as a novel mechanism for creating critical avalanches in the activity of neural networks. We model neurons and chemical synapses by dynamical maps with a uniform noise term in the synaptic coupling. An advantage of utilizing maps is that the dynamical properties (action potential profile, excitability properties, post synaptic potential summation etc.) are not imposed to the system, but occur naturally by solving the system equations. We discuss the relevant neuronal and synaptic properties to achieve the critical state. We verify that networks of excitatory b…
Experimental data have revealed that neuronal connection efficacy exhibits two forms of short-term plasticity, namely, short-term depression (STD) and short-term facilitation (STF). They have time constants residing between fast neural signaling and rapid learning, and may serve as substrates for neural systems manipulating temporal information on relevant time scales. The present study investigates the impact of STD and STF on the dynamics of continuous attractor neural networks (CANNs) and their potential roles in neural information processing. We find that STD endows the network with slow-d…
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…
Understanding the basic operational logics of the nervous system is essential to advancing neuroscientific research. However, theoretical efforts to tackle this fundamental problem are lacking, despite the abundant empirical data about the brain that has been collected in the past few decades. To address this shortcoming, this document introduces a hypothetical framework for the functional nature of primitive neural networks. It analyzes the idea that the activity of neurons and synapses can symbolically reenact the dynamic changes in the world and thus enable an adaptive system of behavior. M…
Various neurophysiological and cognitive functions are based on transferring information between spiking neurons via a complex system of synaptic connections. In particular, the capacity of presynaptic inputs to influence the postsynaptic outputs---the efficacy of the synapses---plays a principal role in all aspects of hippocampal neurophysiology. However, a direct link between the information processed at the level of individual synapses and the animal's ability to form memories at the organismal level has not yet been fully understood. Here, we investigate the effect of synaptic transmission…
Late long-term potentiation (L-LTP) appears essential for the formation of long-term memory, with memories at least partly encoded by patterns of strengthened synapses. How memories are preserved for months or years, despite molecular turnover, is not well understood. Ongoing recurrent neuronal activity, during memory recall or during sleep, has been hypothesized to preferentially potentiate strong synapses, preserving memories. This hypothesis has not been evaluated in the context of a mathematical model representing biochemical pathways important for L-LTP. I incorporated ongoing activity in…
The connectome is a wiring diagram mapping all the neural connections in the brain. At the cellular level, it provides a map of the neurons and synapses within a part or all of the brain of an organism. In recent years, significant advances have been made in the study of the connectome via network science and graph theory. This analysis is fundamental to understand neurotransmission (fast synaptic transmission) networks. However, neurons use other forms of communication as neuromodulation that, instead of conveying excitation or inhibition, change neuronal and synaptic properties. This additio…
Spike-timing dependent plasticity (STDP) is an organizing principle of biological neural networks. While synchronous firing of neurons is considered to be an important functional block in the brain, how STDP shapes neural networks possibly toward synchrony is not entirely clear. We examine relations between STDP and synchronous firing in spontaneously firing neural populations. Using coupled heterogeneous phase oscillators placed on initial networks, we show numerically that STDP prunes some synapses and promotes formation of a feedforward network. Eventually a pacemaker, which is the neuron w…
In neuroscience, learning and memory are usually associated to long-term changes of neuronal connectivity. In this context, synaptic plasticity refers to the set of mechanisms driving the dynamics of neuronal connections, called {\em synapses} and represented by a scalar value, the synaptic weight. Spike-Timing Dependent Plasticity (STDP) is a biologically-based model representing the time evolution of the synaptic weight as a functional of the past spiking activity of adjacent neurons. If numerous models of neuronal cells have been proposed in the mathematical literature, few of them include …
Short-term changes in efficacy have been postulated to enhance the ability of synapses to transmit information between neurons, and within neuronal networks. Even at the level of connections between single neurons, direct confirmation of this simple conjecture has proven elusive. By combining paired-cell recordings, realistic synaptic modelling and information theory, we provide evidence that short-term plasticity can not only improve, but also reduce information transfer between neurons. We focus on a concrete example in rat neocortex, but our results may generalise to other systems. When inf…
Due to the point-like nature of neuronal spiking, efficient neural network simulators often employ event-based simulation schemes for synapses. Yet many types of synaptic plasticity rely on the membrane potential of the postsynaptic cell as a third factor in addition to pre- and postsynaptic spike times. Synapses therefore require continuous information to update their strength which a priori necessitates a continuous update in a time-driven manner. The latter hinders scaling of simulations to realistic cortical network sizes and relevant time scales for learning. Here, we derive two efficient…
An essential step toward understanding neural circuits is linking their structure and their dynamics. In general, this relationship can be almost arbitrarily complex. Recent theoretical work has, however, begun to identify some broad principles underlying collective spiking activity in neural circuits. The first is that local features of network connectivity can be surprisingly effective in predicting global statistics of activity across a network. The second is that, for the important case of large networks with excitatory-inhibitory balance, correlated spiking persists or vanishes depending …
We present an array of leaky integrate-and-fire (LIF) neuron circuits designed for the second-generation BrainScaleS mixed-signal 65-nm CMOS neuromorphic hardware. The neuronal array is embedded in the analog network core of a scaled-down prototype HICANN-DLS chip. Designed as continuous-time circuits, the neurons are highly tunable and reconfigurable elements with accelerated dynamics. Each neuron integrates input current from a multitude of incoming synapses and evokes a digital spike event output. The circuit offers a wide tuning range for synaptic and membrane time constants, as well as fo…
We study synaptically coupled neuronal networks to identify the role of coupling delays in network's synchronized behaviors. We consider a network of excitable, relaxation oscillator neurons where two distinct populations, one excitatory and one inhibitory, are coupled and interact with each other. The excitatory population is uncoupled, while the inhibitory population is tightly coupled. A geometric singular perturbation analysis yields existence and stability conditions for synchronization states under different firing patterns between the two populations, along with formulas for the periods…
Reduced models of neuronal activity such as Integrate-and-Fire models allow a description of neuronal dynamics in simple, intuitive terms and are easy to simulate numerically. We present a method to fit an Integrate-and-Fire-type model of neuronal activity, namely a modified version of the Spike Response Model, to a detailed Hodgkin-Huxley-type neuron model driven by stochastic spike arrival. In the Hogkin-Huxley model, spike arrival at the synapse is modeled by a change of synaptic conductance. For such conductance spike input, more than 70% of the postsynaptic action potentials can be predic…
Synaptic connections in neuronal circuits are modulated by pre- and post-synaptic spiking activity. Heuristic models of this process of synaptic plasticity can provide excellent fits to results from in-vitro experiments in which pre- and post-synaptic spiking is varied in a controlled fashion. However, the plasticity rules inferred from fitting such data are inevitably unstable, in that given constant pre- and post-synaptic activity the synapse will either fully potentiate or depress. This instability can be held in check by adding additional mechanisms, such as homeostasis. Here we consider a…
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…
We propose a neural network model of multi-neuron interacting system that simulates neurons to interact each other through the surroundings of neuronal cell bodies. We physically model the neuronal cell surroundings, include the dendrites, the axons and the synapses as well as the surrounding glial cells, as a collection of all kinds of oscillating modes arisen from the electric circuital environment of neuronal action potentials. By analyzing the dynamics of this neural model through the master equation approach of open quantum systems, we investigate the collective behavior of neurons. After…
Line attractors in neuronal networks have been suggested to be the basis of many brain functions, such as working memory, oculomotor control, head movement, locomotion, and sensory processing. In this paper, we make the connection between line attractors and pulse-gating in feedforward neuronal networks. In this context, because of their neutral stability along a one-dimensional manifold, line attractors are associated with a time-translational invariance that allows graded information to be propagated from one neuronal population to the next. To understand how pulse-gating manifests itself in…
The impact of inhibitory and excitatory synapses in delay-coupled Hodgkin--Huxley neurons that are driven by noise is studied. If both synaptic types are used for coupling, appropriately tuned delays in the inhibition feedback induce multiple firing coherence resonances at sufficiently strong coupling strengths, thus giving rise to tongues of coherency in the corresponding delay-strength parameter plane. If only inhibitory synapses are used, however, appropriately tuned delays also give rise to multiresonant responses, yet the successive delays warranting an optimal coherence of excitations ob…
