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Wilson-Cowan Equations for Neocortical Dynamics. 新皮质动力学的Wilson-Cowan方程。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-01-04 DOI: 10.1186/s13408-015-0034-5
Jack D Cowan, Jeremy Neuman, Wim van Drongelen

In 1972-1973 Wilson and Cowan introduced a mathematical model of the population dynamics of synaptically coupled excitatory and inhibitory neurons in the neocortex. The model dealt only with the mean numbers of activated and quiescent excitatory and inhibitory neurons, and said nothing about fluctuations and correlations of such activity. However, in 1997 Ohira and Cowan, and then in 2007-2009 Buice and Cowan introduced Markov models of such activity that included fluctuation and correlation effects. Here we show how both models can be used to provide a quantitative account of the population dynamics of neocortical activity.We first describe how the Markov models account for many recent measurements of the resting or spontaneous activity of the neocortex. In particular we show that the power spectrum of large-scale neocortical activity has a Brownian motion baseline, and that the statistical structure of the random bursts of spiking activity found near the resting state indicates that such a state can be represented as a percolation process on a random graph, called directed percolation.Other data indicate that resting cortex exhibits pair correlations between neighboring populations of cells, the amplitudes of which decay slowly with distance, whereas stimulated cortex exhibits pair correlations which decay rapidly with distance. Here we show how the Markov model can account for the behavior of the pair correlations.Finally we show how the 1972-1973 Wilson-Cowan equations can account for recent data which indicates that there are at least two distinct modes of cortical responses to stimuli. In mode 1 a low intensity stimulus triggers a wave that propagates at a velocity of about 0.3 m/s, with an amplitude that decays exponentially. In mode 2 a high intensity stimulus triggers a larger response that remains local and does not propagate to neighboring regions.

1972-1973年,Wilson和Cowan引入了新皮层中突触耦合的兴奋性和抑制性神经元种群动态的数学模型。该模型只处理了激活和静止的兴奋性和抑制性神经元的平均数量,而没有说明这些活动的波动和相关性。然而,在1997年Ohira和Cowan,以及2007-2009年Buice和Cowan引入了包括波动和相关效应在内的此类活动的马尔可夫模型。在这里,我们展示了如何使用这两个模型来提供新皮层活动的种群动态的定量说明。我们首先描述了马尔可夫模型如何解释最近对新皮层的静息或自发活动的许多测量。特别是,我们表明大规模新皮层活动的功率谱具有布朗运动基线,并且在静息状态附近发现的峰值活动随机爆发的统计结构表明,这种状态可以表示为随机图上的渗透过程,称为定向渗透。其他数据表明,静息皮层表现出相邻细胞群之间的成对相关性,其振幅随距离缓慢衰减,而受刺激皮层表现出随距离迅速衰减的成对相关性。在这里,我们展示了马尔可夫模型如何解释这对相关性的行为。最后,我们展示了1972-1973年威尔逊-考恩方程如何解释最近的数据,这些数据表明至少有两种不同的皮层对刺激的反应模式。在模式1中,低强度刺激触发的波以约0.3 m/s的速度传播,其振幅呈指数衰减。在模式2中,一个高强度的刺激触发一个更大的反应,这个反应保持在局部,不传播到邻近的区域。
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引用次数: 81
Mathematical Frameworks for Oscillatory Network Dynamics in Neuroscience. 神经科学中振荡网络动力学的数学框架。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-01-06 DOI: 10.1186/s13408-015-0033-6
Peter Ashwin, Stephen Coombes, Rachel Nicks

The tools of weakly coupled phase oscillator theory have had a profound impact on the neuroscience community, providing insight into a variety of network behaviours ranging from central pattern generation to synchronisation, as well as predicting novel network states such as chimeras. However, there are many instances where this theory is expected to break down, say in the presence of strong coupling, or must be carefully interpreted, as in the presence of stochastic forcing. There are also surprises in the dynamical complexity of the attractors that can robustly appear-for example, heteroclinic network attractors. In this review we present a set of mathematical tools that are suitable for addressing the dynamics of oscillatory neural networks, broadening from a standard phase oscillator perspective to provide a practical framework for further successful applications of mathematics to understanding network dynamics in neuroscience.

弱耦合相位振荡器理论的工具对神经科学界产生了深远的影响,让人们深入了解了从中心模式生成到同步等各种网络行为,并预测了嵌合体等新型网络状态。然而,在很多情况下,这一理论会被打破,例如在强耦合的情况下,或者必须仔细解释,例如在随机强迫的情况下。在吸引子的动态复杂性方面也会出现令人惊讶的情况--例如,异链网络吸引子。在这篇综述中,我们介绍了一套适用于解决振荡神经网络动力学问题的数学工具,从标准相位振荡器的角度出发,为进一步成功应用数学理解神经科学中的网络动力学提供了一个实用框架。
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引用次数: 0
Entrainment Ranges for Chains of Forced Neural and Phase Oscillators. 强迫神经和相位振荡器链的夹带范围。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-04-18 DOI: 10.1186/s13408-016-0038-9
Nicole Massarelli, Geoffrey Clapp, Kathleen Hoffman, Tim Kiemel

Sensory input to the lamprey central pattern generator (CPG) for locomotion is known to have a significant role in modulating lamprey swimming. Lamprey CPGs are known to have the ability to entrain to a bending stimulus, that is, in the presence of a rhythmic signal, the CPG will change its frequency to match the stimulus frequency. Bending experiments in which the lamprey spinal cord has been removed and mechanically bent back and forth at a single point have been used to determine the range of frequencies that can entrain the CPG rhythm. First, we model the lamprey locomotor CPG as a chain of neural oscillators with three classes of neurons and sinusoidal forcing representing edge cell input. We derive a phase model using the connections described in the neural model. This results in a simpler model yet maintains some properties of the neural model. For both the neural model and the derived phase model, entrainment ranges are computed for forcing at different points along the chain while varying both intersegmental coupling strength and the coupling strength between the forcer and chain. Entrainment ranges for chains with nonuniform intersegmental coupling asymmetry are larger when forcing is applied to the middle of the chain than when it is applied to either end, a result that is qualitatively similar to the experimental results. In the limit of weak coupling in the chain, the entrainment results of the neural model approach the entrainment results for the derived phase model. Both biological experiments and the robustness of non-monotonic entrainment ranges as a function of the forcing position across different classes of CPG models with nonuniform asymmetric coupling suggest that a specific property of the intersegmental coupling of the CPG is key to entrainment.

七鳃鳗中央模式发生器(CPG)运动的感觉输入在调节七鳃鳗游泳中起着重要作用。众所周知,七鳃鳗CPG具有弯曲刺激的能力,也就是说,在有节奏的信号存在时,CPG会改变其频率以匹配刺激频率。弯曲实验中,七鳃鳗的脊髓被移除,在一个点上机械地前后弯曲,以确定可以携带CPG节律的频率范围。首先,我们将七鳃鳗运动CPG建模为一个神经振荡链,其中有三类神经元和代表边缘细胞输入的正弦强迫。我们利用神经模型中描述的连接推导出相位模型。这导致了一个更简单的模型,但保持了神经模型的一些属性。对于神经模型和推导的相位模型,分别计算了在链上不同位置的力的夹带范围,同时改变了段间耦合强度和力与链之间的耦合强度。对于具有非均匀节段间耦合不对称的链,在链的中间施加力时,夹带范围比在任何一端施加力时都要大,这一结果在质量上与实验结果相似。在链的弱耦合极限下,神经模型的夹带结果接近于推导相模型的夹带结果。生物实验和非单调夹带范围的鲁棒性作为不同类型非均匀非对称耦合CPG模式强迫位置的函数表明,CPG的节段间耦合的特定性质是夹带的关键。
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引用次数: 9
Wave Generation in Unidirectional Chains of Idealized Neural Oscillators. 理想神经振荡器单向链中的波产生。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-04-08 DOI: 10.1186/s13408-016-0037-x
Bastien Fernandez, Stanislav M Mintchev

We investigate the dynamics of unidirectional semi-infinite chains of type-I oscillators that are periodically forced at their root node, as an archetype of wave generation in neural networks. In previous studies, numerical simulations based on uniform forcing have revealed that trajectories approach a traveling wave in the far-downstream, large time limit. While this phenomenon seems typical, it is hardly anticipated because the system does not exhibit any of the crucial properties employed in available proofs of existence of traveling waves in lattice dynamical systems. Here, we give a full mathematical proof of generation under uniform forcing in a simple piecewise affine setting for which the dynamics can be solved explicitly. In particular, our analysis proves existence, global stability, and robustness with respect to perturbations of the forcing, of families of waves with arbitrary period/wave number in some range, for every value of the parameters in the system.

我们研究了在其根节点周期性强制的i型振荡器的单向半无限链的动力学,作为神经网络中波产生的原型。在之前的研究中,基于均匀强迫的数值模拟表明,轨迹在远下游的大时间限制内接近行波。虽然这种现象看起来很典型,但很难预料到,因为该系统没有表现出晶格动力系统中行波存在的现有证明中所使用的任何关键性质。在此,我们给出了在一个简单的分段仿射设置下均匀强迫下的生成的完整数学证明,该设置下的动力学可以显式求解。特别是,我们的分析证明了在一定范围内具有任意周期/波数的波族的存在性,全局稳定性和关于强迫扰动的鲁棒性,对于系统中的每个参数值。
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引用次数: 2
Responses of Leaky Integrate-and-Fire Neurons to a Plurality of Stimuli in Their Receptive Fields. 漏性整合-放电神经元对其感受野中多种刺激的反应。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-05-23 DOI: 10.1186/s13408-016-0040-2
Kang Li, Claus Bundesen, Susanne Ditlevsen

A fundamental question concerning the way the visual world is represented in our brain is how a cortical cell responds when its classical receptive field contains a plurality of stimuli. Two opposing models have been proposed. In the response-averaging model, the neuron responds with a weighted average of all individual stimuli. By contrast, in the probability-mixing model, the cell responds to a plurality of stimuli as if only one of the stimuli were present. Here we apply the probability-mixing and the response-averaging model to leaky integrate-and-fire neurons, to describe neuronal behavior based on observed spike trains. We first estimate the parameters of either model using numerical methods, and then test which model is most likely to have generated the observed data. Results show that the parameters can be successfully estimated and the two models are distinguishable using model selection.

关于视觉世界如何在我们的大脑中呈现的一个基本问题是,当皮质细胞的经典接受野包含多种刺激时,它是如何反应的。人们提出了两种相反的模型。在反应平均模型中,神经元对所有个体刺激的加权平均作出反应。相比之下,在概率混合模型中,细胞对多个刺激作出反应,就好像只有一个刺激存在一样。在这里,我们将概率混合和响应平均模型应用于泄漏的整合和激活神经元,以描述基于观察到的尖峰序列的神经元行为。我们首先用数值方法估计两个模型的参数,然后测试哪个模型最有可能产生观测到的数据。结果表明,采用模型选择方法可以很好地估计出两种模型的参数,并且可以区分两种模型。
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引用次数: 3
Neural Field Models with Threshold Noise. 带有阈值噪声的神经场模型
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-12-01 Epub Date: 2016-03-02 DOI: 10.1186/s13408-016-0035-z
Rüdiger Thul, Stephen Coombes, Carlo R Laing

The original neural field model of Wilson and Cowan is often interpreted as the averaged behaviour of a network of switch like neural elements with a distribution of switch thresholds, giving rise to the classic sigmoidal population firing-rate function so prevalent in large scale neuronal modelling. In this paper we explore the effects of such threshold noise without recourse to averaging and show that spatial correlations can have a strong effect on the behaviour of waves and patterns in continuum models. Moreover, for a prescribed spatial covariance function we explore the differences in behaviour that can emerge when the underlying stationary distribution is changed from Gaussian to non-Gaussian. For travelling front solutions, in a system with exponentially decaying spatial interactions, we make use of an interface approach to calculate the instantaneous wave speed analytically as a series expansion in the noise strength. From this we find that, for weak noise, the spatially averaged speed depends only on the choice of covariance function and not on the shape of the stationary distribution. For a system with a Mexican-hat spatial connectivity we further find that noise can induce localised bump solutions, and using an interface stability argument show that there can be multiple stable solution branches.

威尔逊和考恩(Wilson and Cowan)的原始神经场模型通常被解释为具有开关阈值分布的开关样神经元网络的平均行为,从而产生了在大规模神经元建模中非常普遍的经典的西格码群体发射率函数。在本文中,我们在不求助于平均的情况下探索了这种阈值噪声的影响,结果表明空间相关性会对连续模型中的波和模式的行为产生强烈影响。此外,对于规定的空间协方差函数,我们还探讨了当基本静态分布从高斯分布变为非高斯分布时可能出现的行为差异。对于具有指数衰减空间相互作用的系统中的行进前沿解,我们利用界面方法将瞬时波速作为噪声强度的序列展开进行分析计算。由此我们发现,对于弱噪声,空间平均速度只取决于协方差函数的选择,而不取决于静态分布的形状。对于具有墨西哥帽空间连通性的系统,我们进一步发现,噪声会诱发局部凹凸解,并利用界面稳定性论证表明,可能存在多个稳定解分支。
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引用次数: 0
Stochastic Network Models in Neuroscience: A Festschrift for Jack Cowan. Introduction to the Special Issue 神经科学中的随机网络模型:杰克·考恩的经典著作。特刊简介
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2016-04-04 DOI: 10.1186/s13408-016-0036-y
P. Bressloff, B. Ermentrout, O. Faugeras, P. Thomas
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引用次数: 3
Shifting Spike Times or Adding and Deleting Spikes-How Different Types of Noise Shape Signal Transmission in Neural Populations. 移动尖峰时间或增加和删除尖峰——不同类型的噪声如何影响神经群体中的信号传输。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2015-12-01 Epub Date: 2015-01-12 DOI: 10.1186/2190-8567-5-1
Sergej O Voronenko, Wilhelm Stannat, Benjamin Lindner

We study a population of spiking neurons which are subject to independent noise processes and a strong common time-dependent input. We show that the response of output spikes to independent noise shapes information transmission of such populations even when information transmission properties of single neurons are left unchanged. In particular, we consider two Poisson models in which independent noise either (i) adds and deletes spikes (AD model) or (ii) shifts spike times (STS model). We show that in both models suprathreshold stochastic resonance (SSR) can be observed, where the information transmitted by a neural population is increased with addition of independent noise. In the AD model, the presence of the SSR effect is robust and independent of the population size or the noise spectral statistics. In the STS model, the information transmission properties of the population are determined by the spectral statistics of the noise, leading to a strongly increased effect of SSR in some regimes, or an absence of SSR in others. Furthermore, we observe a high-pass filtering of information in the STS model that is absent in the AD model. We quantify information transmission by means of the lower bound on the mutual information rate and the spectral coherence function. To this end, we derive the signal-output cross-spectrum, the output power spectrum, and the cross-spectrum of two spike trains for both models analytically.

我们研究了一群受独立噪声过程和强公共时变输入影响的尖峰神经元。我们表明,即使单个神经元的信息传输特性保持不变,输出尖峰对独立噪声的响应也会影响这些群体的信息传输。特别地,我们考虑了两个泊松模型,其中独立噪声要么(i)增加和删除尖峰(AD模型),要么(ii)移动尖峰时间(STS模型)。我们表明,在这两种模型中都可以观察到超阈值随机共振(SSR),其中神经种群传递的信息随着独立噪声的增加而增加。在AD模型中,SSR效应的存在具有鲁棒性,且与种群大小或噪声谱统计量无关。在STS模型中,种群的信息传递特性由噪声的谱统计量决定,导致SSR效应在某些制度下显著增加,或在其他制度下不存在。此外,我们在STS模型中观察到AD模型中不存在的信息高通滤波。我们通过互信息率的下界和谱相干函数来量化信息传输。为此,我们导出了两种模型的信号输出交叉频谱、输出功率谱和两个尖峰串的交叉频谱。
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引用次数: 13
Uncertainty Propagation in Nerve Impulses Through the Action Potential Mechanism. 通过动作电位机制的神经冲动不确定性传播。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2015-12-01 Epub Date: 2015-01-12 DOI: 10.1186/2190-8567-5-3
Aldemar Torres Valderrama, Jeroen Witteveen, Maria Navarro, Joke Blom

We investigate the propagation of probabilistic uncertainty through the action potential mechanism in nerve cells. Using the Hodgkin-Huxley (H-H) model and Stochastic Collocation on Sparse Grids, we obtain an accurate probabilistic interpretation of the deterministic dynamics of the transmembrane potential and gating variables. Using Sobol indices, out of the 11 uncertain parameters in the H-H model, we unravel two main uncertainty sources, which account for more than 90 % of the fluctuations in neuronal responses, and have a direct biophysical interpretation. We discuss how this interesting feature of the H-H model allows one to reduce greatly the probabilistic degrees of freedom in uncertainty quantification analyses, saving CPU time in numerical simulations and opening possibilities for probabilistic generalisation of other deterministic models of great importance in physiology and mathematical neuroscience.

我们通过神经细胞的动作电位机制来研究概率不确定性的传播。利用霍奇金-赫胥黎(H-H)模型和稀疏网格上的随机配置,我们获得了跨膜电位和门控变量的确定性动力学的精确概率解释。利用Sobol指数,在H-H模型的11个不确定参数中,我们揭示了两个主要的不确定源,它们占神经元响应波动的90%以上,并且具有直接的生物物理解释。我们讨论了H-H模型的这个有趣的特征如何允许人们在不确定性量化分析中大大降低概率自由度,节省数值模拟中的CPU时间,并为生理学和数学神经科学中非常重要的其他确定性模型的概率推广提供可能性。
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引用次数: 10
Stochastic Synchronization in Purkinje Cells with Feedforward Inhibition Could Be Studied with Equivalent Phase-Response Curves. 用等效相位响应曲线研究具有前馈抑制的浦肯野细胞的随机同步。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2015-12-01 Epub Date: 2015-06-19 DOI: 10.1186/s13408-015-0025-6
Sergio Verduzco-Flores

Simple-spike synchrony between Purkinje cells projecting to a common neuron in the deep cerebellar nucleus is emerging as an important factor in the encoding of output information from cerebellar cortex. A phenomenon known as stochastic synchronization happens when uncoupled oscillators synchronize due to correlated inputs. Stochastic synchronization is a viable mechanism through which simple-spike synchrony could be generated, but it has received scarce attention, perhaps because the presence of feedforward inhibition in the input to Purkinje cells makes insights difficult. This paper presents a method to account for feedforward inhibition so the usual mathematical approaches to stochastic synchronization can be applied. The method consists in finding a single Phase Response Curve, called the equivalent PRC, that accounts for the effects of both excitatory inputs and delayed feedforward inhibition from molecular layer interneurons. The results suggest that a theory of stochastic synchronization for the case of feedforward inhibition may not be necessary, since this case can be approximately reduced to the case of inputs characterized by a single PRC. Moreover, feedforward inhibition could in many situations increase the level of synchrony experienced by Purkinje cells.

浦肯野细胞与小脑深部核共同神经元之间的单峰同步是编码小脑皮层输出信息的重要因素。当非耦合振荡器由于相关输入而同步时,就会发生随机同步现象。随机同步是一种可行的机制,通过它可以产生简单的脉冲同步,但它很少受到关注,也许是因为在浦肯野细胞的输入中存在前馈抑制,这使得人们很难深入了解。本文提出了一种考虑前馈抑制的方法,从而可以应用通常的随机同步数学方法。该方法包括找到一个称为等效PRC的单相响应曲线,该曲线可以解释来自分子层中间神经元的兴奋性输入和延迟前馈抑制的影响。结果表明,前馈抑制情况下的随机同步理论可能不是必要的,因为这种情况可以近似地简化为以单个PRC为特征的输入情况。此外,前馈抑制可以在许多情况下增加浦肯野细胞的同步性水平。
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引用次数: 2
期刊
Journal of Mathematical Neuroscience
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