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Canard solutions in neural mass models: consequences on critical regimes. 神经质量模型中的鸭式解:临界状态的后果。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-09-16 DOI: 10.1186/s13408-021-00109-z
Elif Köksal Ersöz, Fabrice Wendling

Mathematical models at multiple temporal and spatial scales can unveil the fundamental mechanisms of critical transitions in brain activities. Neural mass models (NMMs) consider the average temporal dynamics of interconnected neuronal subpopulations without explicitly representing the underlying cellular activity. The mesoscopic level offered by the neural mass formulation has been used to model electroencephalographic (EEG) recordings and to investigate various cerebral mechanisms, such as the generation of physiological and pathological brain activities. In this work, we consider a NMM widely accepted in the context of epilepsy, which includes four interacting neuronal subpopulations with different synaptic kinetics. Due to the resulting three-time-scale structure, the model yields complex oscillations of relaxation and bursting types. By applying the principles of geometric singular perturbation theory, we unveil the existence of the canard solutions and detail how they organize the complex oscillations and excitability properties of the model. In particular, we show that boundaries between pathological epileptic discharges and physiological background activity are determined by the canard solutions. Finally we report the existence of canard-mediated small-amplitude frequency-specific oscillations in simulated local field potentials for decreased inhibition conditions. Interestingly, such oscillations are actually observed in intracerebral EEG signals recorded in epileptic patients during pre-ictal periods, close to seizure onsets.

在多个时间和空间尺度上的数学模型可以揭示大脑活动关键转变的基本机制。神经质量模型(nmm)考虑相互连接的神经元亚群的平均时间动态,而没有明确表示潜在的细胞活动。神经团的形成所提供的介观水平已被用于模拟脑电图(EEG)记录,并研究各种大脑机制,如生理和病理脑活动的产生。在这项工作中,我们考虑了在癫痫背景下广泛接受的NMM,其中包括四个相互作用的神经元亚群,它们具有不同的突触动力学。由于所产生的三时间尺度结构,该模型产生了松弛型和破裂型的复杂振荡。利用几何奇异摄动理论的原理,揭示了鸭尔德解的存在性,并详细说明了鸭尔德解如何组织模型的复振荡和可激性。特别是,我们表明病理性癫痫放电和生理背景活动之间的界限是由鸭式溶液决定的。最后,我们报告了鸭翼介导的小幅度频率特异性振荡在抑制条件下降的模拟局部场电位中存在。有趣的是,这种振荡实际上是在癫痫患者在癫痫发作前的脑电图信号中观察到的。
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引用次数: 2
Rendering neuronal state equations compatible with the principle of stationary action. 使神经元状态方程符合静止作用原理
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-08-12 DOI: 10.1186/s13408-021-00108-0
Erik D Fagerholm, W M C Foulkes, Karl J Friston, Rosalyn J Moran, Robert Leech

The principle of stationary action is a cornerstone of modern physics, providing a powerful framework for investigating dynamical systems found in classical mechanics through to quantum field theory. However, computational neuroscience, despite its heavy reliance on concepts in physics, is anomalous in this regard as its main equations of motion are not compatible with a Lagrangian formulation and hence with the principle of stationary action. Taking the Dynamic Causal Modelling (DCM) neuronal state equation as an instructive archetype of the first-order linear differential equations commonly found in computational neuroscience, we show that it is possible to make certain modifications to this equation to render it compatible with the principle of stationary action. Specifically, we show that a Lagrangian formulation of the DCM neuronal state equation is facilitated using a complex dependent variable, an oscillatory solution, and a Hermitian intrinsic connectivity matrix. We first demonstrate proof of principle by using Bayesian model inversion to show that both the original and modified models can be correctly identified via in silico data generated directly from their respective equations of motion. We then provide motivation for adopting the modified models in neuroscience by using three different types of publicly available in vivo neuroimaging datasets, together with open source MATLAB code, to show that the modified (oscillatory) model provides a more parsimonious explanation for some of these empirical timeseries. It is our hope that this work will, in combination with existing techniques, allow people to explore the symmetries and associated conservation laws within neural systems - and to exploit the computational expediency facilitated by direct variational techniques.

静止作用原理是现代物理学的基石,它为研究从经典力学到量子场论的动力学系统提供了一个强大的框架。然而,尽管计算神经科学在很大程度上依赖物理学中的概念,但它在这方面却很反常,因为它的主要运动方程与拉格朗日公式不兼容,因此也与静止作用原理不兼容。以动态因果建模(DCM)神经元状态方程作为计算神经科学中常见的一阶线性微分方程的原型,我们证明可以对该方程进行某些修改,使其符合静止作用原理。具体来说,我们证明了使用复杂因变量、振荡解和赫米特内在连接矩阵可以方便地对 DCM 神经元状态方程进行拉格朗日表述。我们首先使用贝叶斯模型反演来证明原理,表明通过直接从各自运动方程生成的硅学数据,可以正确识别原始模型和修正模型。然后,我们利用三种不同类型的公开活体神经成像数据集和开放源代码 MATLAB,证明修正(振荡)模型为其中一些经验时间序列提供了更合理的解释,从而为在神经科学中采用修正模型提供了动力。我们希望这项工作能与现有技术相结合,让人们探索神经系统的对称性和相关守恒定律,并利用直接变分技术带来的计算便利。
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引用次数: 0
Pattern formation in a 2-population homogenized neuronal network model. 2种群均质神经网络模型的模式形成。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-06-26 DOI: 10.1186/s13408-021-00107-1
Karina Kolodina, John Wyller, Anna Oleynik, Mads Peter Sørensen

We study pattern formation in a 2-population homogenized neural field model of the Hopfield type in one spatial dimension with periodic microstructure. The connectivity functions are periodically modulated in both the synaptic footprint and in the spatial scale. It is shown that the nonlocal synaptic interactions promote a finite band width instability. The stability method relies on a sequence of wave-number dependent invariants of [Formula: see text]-stability matrices representing the sequence of Fourier-transformed linearized evolution equations for the perturbation imposed on the homogeneous background. The generic picture of the instability structure consists of a finite set of well-separated gain bands. In the shallow firing rate regime the nonlinear development of the instability is determined by means of the translational invariant model with connectivity kernels replaced with the corresponding period averaged connectivity functions. In the steep firing rate regime the pattern formation process depends sensitively on the spatial localization of the connectivity kernels: For strongly localized kernels this process is determined by the translational invariant model with period averaged connectivity kernels, whereas in the complementary regime of weak and moderate localization requires the homogenized model as a starting point for the analysis. We follow the development of the instability numerically into the nonlinear regime for both steep and shallow firing rate functions when the connectivity kernels are modeled by means of an exponentially decaying function. We also study the pattern forming process numerically as a function of the heterogeneity parameters in four different regimes ranging from the weakly modulated case to the strongly heterogeneous case. For the weakly modulated regime, we observe that stable spatial oscillations are formed in the steep firing rate regime, whereas we get spatiotemporal oscillations in the shallow regime of the firing rate functions.

本文研究了具有周期微观结构的Hopfield型2种群均质神经场模型在一维空间上的模式形成。连接功能在突触足迹和空间尺度上都是周期性调节的。结果表明,非局域突触相互作用促进了有限带宽的不稳定性。稳定性方法依赖于[公式:见文本]稳定性矩阵的波数不变量序列,稳定性矩阵表示施加在齐次背景上的扰动的傅立叶变换线性化演化方程序列。不稳定结构的一般图像由一组分离良好的增益带组成。在浅发射速率下,用相应周期平均连通性函数代替连通性核的平移不变模型确定了不稳定性的非线性发展。在急剧燃烧速率下,模式形成过程敏感地依赖于连通性核的空间局部化:对于强局部化核,该过程由具有周期平均连通性核的平移不变模型决定,而在弱和中等局部化互补状态下,需要均质化模型作为分析的起点。当连通性核用指数衰减函数建模时,我们从数值上跟踪陡射率函数和浅射率函数的不稳定性发展到非线性状态。我们还用数值方法研究了从弱调制到强非均质情况下四种不同情况下的非均质参数的函数模式形成过程。对于弱调制区,我们观察到在陡峭发射速率区形成了稳定的空间振荡,而在发射速率函数的浅区则形成了时空振荡。
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引用次数: 1
Auditory streaming emerges from fast excitation and slow delayed inhibition. 听觉流产生于快速兴奋和缓慢延迟抑制。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-05-03 DOI: 10.1186/s13408-021-00106-2
Andrea Ferrario, James Rankin

In the auditory streaming paradigm, alternating sequences of pure tones can be perceived as a single galloping rhythm (integration) or as two sequences with separated low and high tones (segregation). Although studied for decades, the neural mechanisms underlining this perceptual grouping of sound remains a mystery. With the aim of identifying a plausible minimal neural circuit that captures this phenomenon, we propose a firing rate model with two periodically forced neural populations coupled by fast direct excitation and slow delayed inhibition. By analyzing the model in a non-smooth, slow-fast regime we analytically prove the existence of a rich repertoire of dynamical states and of their parameter dependent transitions. We impose plausible parameter restrictions and link all states with perceptual interpretations. Regions of stimulus parameters occupied by states linked with each percept match those found in behavioural experiments. Our model suggests that slow inhibition masks the perception of subsequent tones during segregation (forward masking), whereas fast excitation enables integration for large pitch differences between the two tones.

在听觉流范式中,纯音的交替序列可以被理解为一个单一的飞驰节奏(整合)或两个低音调和高音调分离的序列(分离)。尽管研究了几十年,强调这种声音感知分组的神经机制仍然是一个谜。为了确定捕获这种现象的合理的最小神经回路,我们提出了一个具有两个周期性强迫神经群的放电率模型,该模型由快速直接激励和缓慢延迟抑制耦合。通过分析非光滑、慢快状态下的模型,我们解析地证明了丰富的动态状态库及其参数相关跃迁的存在性。我们施加合理的参数限制,并将所有状态与感知解释联系起来。与每个感知相关联的状态所占据的刺激参数区域与行为实验中发现的区域相匹配。我们的模型表明,在分离过程中,缓慢的抑制掩盖了对后续音调的感知(前向掩蔽),而快速的激发能够整合两个音调之间的大音高差异。
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引用次数: 3
A model of on/off transitions in neurons of the deep cerebellar nuclei: deciphering the underlying ionic mechanisms. 小脑深部核神经元的开/关转换模型:解读潜在的离子机制。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-04-01 DOI: 10.1186/s13408-021-00105-3
Hugues Berry, Stéphane Genet

The neurons of the deep cerebellar nuclei (DCNn) represent the main functional link between the cerebellar cortex and the rest of the central nervous system. Therefore, understanding the electrophysiological properties of DCNn is of fundamental importance to understand the overall functioning of the cerebellum. Experimental data suggest that DCNn can reversibly switch between two states: the firing of spikes (F state) and a stable depolarized state (SD state). We introduce a new biophysical model of the DCNn membrane electro-responsiveness to investigate how the interplay between the documented conductances identified in DCNn give rise to these states. In the model, the F state emerges as an isola of limit cycles, i.e. a closed loop of periodic solutions disconnected from the branch of SD fixed points. This bifurcation structure endows the model with the ability to reproduce the [Formula: see text] transition triggered by hyperpolarizing current pulses. The model also reproduces the [Formula: see text] transition induced by blocking Ca currents and ascribes this transition to the blocking of the high-threshold Ca current. The model suggests that intracellular current injections can trigger fully reversible [Formula: see text] transitions. Investigation of low-dimension reduced models suggests that the voltage-dependent Na current is prominent for these dynamical features. Finally, simulations of the model suggest that physiological synaptic inputs may trigger [Formula: see text] transitions. These transitions could explain the puzzling observation of positively correlated activities of connected Purkinje cells and DCNn despite the former inhibit the latter.

小脑深部核(DCNn)的神经元代表了小脑皮层和中枢神经系统其余部分之间的主要功能联系。因此,了解DCNn的电生理特性对于了解小脑的整体功能至关重要。实验数据表明,DCNn可以可逆地在两种状态之间切换:尖峰放电(F态)和稳定的去极化状态(SD态)。我们引入了一种新的DCNn膜电响应的生物物理模型,以研究DCNn中记录的电导之间的相互作用如何引起这些状态。在模型中,F态表现为一个极限环孤立体,即与SD不动点分支断开的周期解闭环。这种分岔结构使模型能够重现由超极化电流脉冲触发的[公式:见文本]过渡。该模型还再现了由阻断Ca电流引起的转变,并将这种转变归因于阻断高阈值Ca电流。该模型表明,细胞内电流注入可以触发完全可逆的[公式:见文本]转变。对低维降维模型的研究表明,电压依赖的钠电流是这些动态特征的突出体现。最后,该模型的模拟表明,生理突触输入可能触发[公式:见文本]转换。这些转变可以解释连接的浦肯野细胞和DCNn的正相关活动的令人困惑的观察,尽管前者抑制后者。
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引用次数: 2
Estimating Fisher discriminant error in a linear integrator model of neural population activity. 估计神经群活动线性积分器模型中的费雪判别误差
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-02-19 DOI: 10.1186/s13408-021-00104-4
Matias Calderini, Jean-Philippe Thivierge

Decoding approaches provide a useful means of estimating the information contained in neuronal circuits. In this work, we analyze the expected classification error of a decoder based on Fisher linear discriminant analysis. We provide expressions that relate decoding error to the specific parameters of a population model that performs linear integration of sensory input. Results show conditions that lead to beneficial and detrimental effects of noise correlation on decoding. Further, the proposed framework sheds light on the contribution of neuronal noise, highlighting cases where, counter-intuitively, increased noise may lead to improved decoding performance. Finally, we examined the impact of dynamical parameters, including neuronal leak and integration time constant, on decoding. Overall, this work presents a fruitful approach to the study of decoding using a comprehensive theoretical framework that merges dynamical parameters with estimates of readout error.

解码方法为估计神经元回路中包含的信息提供了一种有用的手段。在这项工作中,我们分析了基于费雪线性判别分析的解码器的预期分类误差。我们提供了将解码误差与对感觉输入进行线性整合的群体模型的特定参数联系起来的表达式。结果显示了噪声相关性对解码产生有利和不利影响的条件。此外,所提出的框架还揭示了神经元噪声的贡献,强调了在与直觉相反的情况下,噪声的增加可能会导致解码性能的提高。最后,我们研究了动态参数(包括神经元泄漏和整合时间常数)对解码的影响。总之,这项研究提出了一种富有成效的解码研究方法,它采用了一个综合的理论框架,将动态参数与读出误差估计相结合。
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引用次数: 0
M-current induced Bogdanov-Takens bifurcation and switching of neuron excitability class. m电流诱导神经元兴奋性类的Bogdanov-Takens分岔与切换。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-02-15 DOI: 10.1186/s13408-021-00103-5
Isam Al-Darabsah, Sue Ann Campbell

In this work, we consider a general conductance-based neuron model with the inclusion of the acetycholine sensitive, M-current. We study bifurcations in the parameter space consisting of the applied current [Formula: see text], the maximal conductance of the M-current [Formula: see text] and the conductance of the leak current [Formula: see text]. We give precise conditions for the model that ensure the existence of a Bogdanov-Takens (BT) point and show that such a point can occur by varying [Formula: see text] and [Formula: see text]. We discuss the case when the BT point becomes a Bogdanov-Takens-cusp (BTC) point and show that such a point can occur in the three-dimensional parameter space. The results of the bifurcation analysis are applied to different neuronal models and are verified and supplemented by numerical bifurcation diagrams generated using the package MATCONT. We conclude that there is a transition in the neuronal excitability type organised by the BT point and the neuron switches from Class-I to Class-II as conductance of the M-current increases.

在这项工作中,我们考虑了一个基于一般电导的神经元模型,其中包括乙酰胆碱敏感的m电流。我们研究了由施加电流[公式:见文]、m电流的最大电导[公式:见文]和漏电流的电导[公式:见文]组成的参数空间中的分岔。我们给出了保证波格丹诺夫- takens (BT)点存在的模型的精确条件,并表明这样的点可以通过改变[公式:见文]和[公式:见文]而出现。讨论了BT点成为波格丹诺夫-塔肯斯尖点(BTC)的情况,并证明了这种点在三维参数空间中可以出现。分岔分析的结果应用于不同的神经元模型,并通过使用MATCONT包生成的数值分岔图进行验证和补充。我们的结论是,随着m电流电导的增加,由BT点组织的神经元兴奋性类型发生了转变,神经元从i类转换到ii类。
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引用次数: 3
Retroactive interference model of forgetting. 遗忘的追溯干扰模式
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-01-23 DOI: 10.1186/s13408-021-00102-6
Antonios Georgiou, Mikhail Katkov, Misha Tsodyks

Memory and forgetting constitute two sides of the same coin, and although the first has been extensively investigated, the latter is often overlooked. A possible approach to better understand forgetting is to develop phenomenological models that implement its putative mechanisms in the most elementary way possible, and then experimentally test the theoretical predictions of these models. One such mechanism proposed in previous studies is retrograde interference, stating that a memory can be erased due to subsequently acquired memories. In the current contribution, we hypothesize that retrograde erasure is controlled by the relevant "importance" measures such that more important memories eliminate less important ones acquired earlier. We show that some versions of the resulting mathematical model are broadly compatible with the previously reported power-law forgetting time course and match well the results of our recognition experiments with long, randomly assembled streams of words.

记忆和遗忘是一枚硬币的两面,虽然前者已被广泛研究,但后者却常常被忽视。为了更好地理解遗忘,一种可行的方法是建立现象学模型,以最基本的方式实现遗忘的假定机制,然后通过实验检验这些模型的理论预测。以往研究中提出的逆行干扰就是这样一种机制,即记忆会因随后获得的记忆而被抹去。在当前的研究中,我们假设逆行消除是由相关的 "重要性 "指标控制的,因此,较重要的记忆会消除较不重要的早期获得的记忆。我们的研究表明,由此产生的数学模型的某些版本与之前报道的幂律遗忘时间过程基本一致,并且与我们用随机组合的长单词流进行的识别实验的结果非常吻合。
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引用次数: 0
On the potential role of lateral connectivity in retinal anticipation. 关于侧连通性在视网膜预期中的潜在作用。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-01-09 DOI: 10.1186/s13408-020-00101-z
Selma Souihel, Bruno Cessac

We analyse the potential effects of lateral connectivity (amacrine cells and gap junctions) on motion anticipation in the retina. Our main result is that lateral connectivity can-under conditions analysed in the paper-trigger a wave of activity enhancing the anticipation mechanism provided by local gain control (Berry et al. in Nature 398(6725):334-338, 1999; Chen et al. in J. Neurosci. 33(1):120-132, 2013). We illustrate these predictions by two examples studied in the experimental literature: differential motion sensitive cells (Baccus and Meister in Neuron 36(5):909-919, 2002) and direction sensitive cells where direction sensitivity is inherited from asymmetry in gap junctions connectivity (Trenholm et al. in Nat. Neurosci. 16:154-156, 2013). We finally present reconstructions of retinal responses to 2D visual inputs to assess the ability of our model to anticipate motion in the case of three different 2D stimuli.

我们分析了侧连通性(无突细胞和间隙连接)对视网膜运动预期的潜在影响。我们的主要结果是,在论文中分析的条件下,横向连通性可以触发一波活动,增强局部增益控制提供的预期机制(Berry等人,Nature 398(6725):334-338, 1999;神经科学学报,33(1):120-132,2013。我们通过实验文献中研究的两个例子来说明这些预测:微分运动敏感细胞(Baccus和Meister在Neuron 36(5):909-919, 2002)和方向敏感细胞,其中方向敏感性遗传自间隙连接的不对称性(Trenholm等人在Nat. Neurosci. 16:154-156, 2013)。我们最后展示了视网膜对2D视觉输入的反应重建,以评估我们的模型在三种不同的2D刺激情况下预测运动的能力。
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引用次数: 5
Noisy network attractor models for transitions between EEG microstates. 脑电微态转换的噪声网络吸引子模型。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2021-01-04 DOI: 10.1186/s13408-020-00100-0
Jennifer Creaser, Peter Ashwin, Claire Postlethwaite, Juliane Britz

The brain is intrinsically organized into large-scale networks that constantly re-organize on multiple timescales, even when the brain is at rest. The timing of these dynamics is crucial for sensation, perception, cognition, and ultimately consciousness, but the underlying dynamics governing the constant reorganization and switching between networks are not yet well understood. Electroencephalogram (EEG) microstates are brief periods of stable scalp topography that have been identified as the electrophysiological correlate of functional magnetic resonance imaging defined resting-state networks. Spatiotemporal microstate sequences maintain high temporal resolution and have been shown to be scale-free with long-range temporal correlations. Previous attempts to model EEG microstate sequences have failed to capture this crucial property and so cannot fully capture the dynamics; this paper answers the call for more sophisticated modeling approaches. We present a dynamical model that exhibits a noisy network attractor between nodes that represent the microstates. Using an excitable network between four nodes, we can reproduce the transition probabilities between microstates but not the heavy tailed residence time distributions. We present two extensions to this model: first, an additional hidden node at each state; second, an additional layer that controls the switching frequency in the original network. Introducing either extension to the network gives the flexibility to capture these heavy tails. We compare the model generated sequences to microstate sequences from EEG data collected from healthy subjects at rest. For the first extension, we show that the hidden nodes 'trap' the trajectories allowing the control of residence times at each node. For the second extension, we show that two nodes in the controlling layer are sufficient to model the long residence times. Finally, we show that in addition to capturing the residence time distributions and transition probabilities of the sequences, these two models capture additional properties of the sequences including having interspersed long and short residence times and long range temporal correlations in line with the data as measured by the Hurst exponent.

大脑在本质上被组织成大规模的网络,在多个时间尺度上不断重组,即使大脑处于休息状态。这些动态的时间对感觉、知觉、认知和最终的意识至关重要,但控制网络之间不断重组和切换的潜在动态尚未得到很好的理解。脑电图(EEG)微状态是稳定的头皮地形的短暂时期,已被确定为功能磁共振成像定义的静息状态网络的电生理相关。时空微态序列保持高时间分辨率,并具有无标度的长程时间相关性。以前对EEG微状态序列建模的尝试未能捕捉到这一关键特性,因此无法完全捕捉到动态;本文响应了对更复杂的建模方法的需求。我们提出了一个动态模型,该模型显示了代表微观状态的节点之间的噪声网络吸引子。利用四个节点之间的可激网络,我们可以再现微观状态之间的跃迁概率,但不能再现重尾停留时间分布。我们对该模型进行了两个扩展:首先,在每个状态处增加一个隐藏节点;第二层是控制原始网络中交换频率的附加层。在网络中引入任意一种扩展都可以灵活地捕获这些重尾。我们将模型生成的序列与从健康受试者在休息时收集的脑电图数据的微状态序列进行比较。对于第一个扩展,我们证明了隐藏节点“捕获”了允许控制每个节点停留时间的轨迹。对于第二次扩展,我们证明了控制层中的两个节点足以模拟长停留时间。最后,我们表明,除了捕获序列的停留时间分布和转移概率外,这两个模型还捕获了序列的其他属性,包括与Hurst指数测量的数据相一致的长、短停留时间和长范围时间相关性。
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引用次数: 7
期刊
Journal of Mathematical Neuroscience
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