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Exact mean-field models for spiking neural networks with adaptation. 带自适应脉冲神经网络的精确平均场模型。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-11-01 Epub Date: 2022-07-14 DOI: 10.1007/s10827-022-00825-9
Liang Chen, Sue Ann Campbell

Networks of spiking neurons with adaption have been shown to be able to reproduce a wide range of neural activities, including the emergent population bursting and spike synchrony that underpin brain disorders and normal function. Exact mean-field models derived from spiking neural networks are extremely valuable, as such models can be used to determine how individual neurons and the network they reside within interact to produce macroscopic network behaviours. In the paper, we derive and analyze a set of exact mean-field equations for the neural network with spike frequency adaptation. Specifically, our model is a network of Izhikevich neurons, where each neuron is modeled by a two dimensional system consisting of a quadratic integrate and fire equation plus an equation which implements spike frequency adaptation. Previous work deriving a mean-field model for this type of network, relied on the assumption of sufficiently slow dynamics of the adaptation variable. However, this approximation did not succeed in establishing an exact correspondence between the macroscopic description and the realistic neural network, especially when the adaptation time constant was not large. The challenge lies in how to achieve a closed set of mean-field equations with the inclusion of the mean-field dynamics of the adaptation variable. We address this problem by using a Lorentzian ansatz combined with the moment closure approach to arrive at a mean-field system in the thermodynamic limit. The resulting macroscopic description is capable of qualitatively and quantitatively describing the collective dynamics of the neural network, including transition between states where the individual neurons exhibit asynchronous tonic firing and synchronous bursting. We extend the approach to a network of two populations of neurons and discuss the accuracy and efficacy of our mean-field approximations by examining all assumptions that are imposed during the derivation. Numerical bifurcation analysis of our mean-field models reveals bifurcations not previously observed in the models, including a novel mechanism for emergence of bursting in the network. We anticipate our results will provide a tractable and reliable tool to investigate the underlying mechanism of brain function and dysfunction from the perspective of computational neuroscience.

具有适应性的尖峰神经元网络已被证明能够再现广泛的神经活动,包括支撑大脑紊乱和正常功能的突发性种群爆发和尖峰同步。源自脉冲神经网络的精确平均场模型是非常有价值的,因为这样的模型可以用来确定单个神经元及其所在网络如何相互作用以产生宏观网络行为。本文导出并分析了具有尖峰频率自适应的神经网络的一组精确平均场方程。具体来说,我们的模型是一个Izhikevich神经元网络,其中每个神经元由一个二维系统建模,该系统由二次积分和火焰方程以及实现峰值频率自适应的方程组成。先前的工作推导了这类网络的平均场模型,依赖于自适应变量的足够慢的动态假设。然而,这种近似并没有成功地建立宏观描述与现实神经网络之间的精确对应关系,特别是当自适应时间常数不大时。挑战在于如何获得包含自适应变量的平均场动力学的一组封闭的平均场方程。我们用洛伦兹解算结合矩闭的方法来解决这个问题,得到了热力学极限下的平均场系统。由此产生的宏观描述能够定性和定量地描述神经网络的集体动力学,包括单个神经元表现出异步强直放电和同步爆发的状态之间的转换。我们将该方法扩展到两个神经元群体的网络,并通过检查推导过程中施加的所有假设来讨论我们的平均场近似的准确性和有效性。我们的平均场模型的数值分岔分析揭示了以前未在模型中观察到的分岔,包括网络中出现破裂的新机制。我们期望我们的研究结果将为从计算神经科学的角度研究脑功能和功能障碍的潜在机制提供一个易于操作和可靠的工具。
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引用次数: 7
Probabilistic solvers enable a straight-forward exploration of numerical uncertainty in neuroscience models. 概率解算器可以直接探索神经科学模型中的数值不确定性。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-11-01 DOI: 10.1007/s10827-022-00827-7
Jonathan Oesterle, Nicholas Krämer, Philipp Hennig, Philipp Berens

Understanding neural computation on the mechanistic level requires models of neurons and neuronal networks. To analyze such models one typically has to solve coupled ordinary differential equations (ODEs), which describe the dynamics of the underlying neural system. These ODEs are solved numerically with deterministic ODE solvers that yield single solutions with either no, or only a global scalar error indicator on precision. It can therefore be challenging to estimate the effect of numerical uncertainty on quantities of interest, such as spike-times and the number of spikes. To overcome this problem, we propose to use recently developed sampling-based probabilistic solvers, which are able to quantify such numerical uncertainties. They neither require detailed insights into the kinetics of the models, nor are they difficult to implement. We show that numerical uncertainty can affect the outcome of typical neuroscience simulations, e.g. jittering spikes by milliseconds or even adding or removing individual spikes from simulations altogether, and demonstrate that probabilistic solvers reveal these numerical uncertainties with only moderate computational overhead.

在机制层面上理解神经计算需要神经元和神经网络的模型。要分析这种模型,通常必须求解耦合常微分方程(ode),它描述了底层神经系统的动力学。这些ODE是用确定性ODE求解器进行数值求解的,该求解器产生单个解,在精度上没有或只有全局标量误差指示器。因此,估计数值不确定性对感兴趣的数量(如峰值时间和峰值数量)的影响可能具有挑战性。为了克服这个问题,我们建议使用最近开发的基于抽样的概率求解器,它能够量化这种数值不确定性。它们既不需要详细了解模型的动力学,也不难以实现。我们表明数值不确定性可以影响典型神经科学模拟的结果,例如毫秒级的抖动尖峰,甚至从模拟中添加或删除单个尖峰,并证明概率解算器只需要适度的计算开销就可以揭示这些数值不确定性。
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引用次数: 0
Homogeneous inhibition is optimal for the phase precession of place cells in the CA1 field. 均匀抑制对于CA1区中定位细胞的相位进动是最佳的。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-07-05 DOI: 10.1007/s10827-023-00855-x
Georgy Vandyshev, Ivan Mysin

Place cells are hippocampal neurons encoding the position of an animal in space. Studies of place cells are essential to understanding the processing of information by neural networks of the brain. An important characteristic of place cell spike trains is phase precession. When an animal is running through the place field, the discharges of the place cells shift from the ascending phase of the theta rhythm through the minimum to the descending phase. The role of excitatory inputs to pyramidal neurons along the Schaffer collaterals and the perforant pathway in phase precession is described, but the role of local interneurons is poorly understood. Our goal is estimating of the contribution of field CA1 interneurons to the phase precession of place cells using mathematical methods. The CA1 field is chosen because it provides the largest set of experimental data required to build and verify the model. Our simulations discover optimal parameters of the excitatory and inhibitory inputs to the pyramidal neuron so that it generates a spike train with the effect of phase precession. The uniform inhibition of pyramidal neurons best explains the effect of phase precession. Among interneurons, axo-axonal neurons make the greatest contribution to the inhibition of pyramidal cells.

位置细胞是海马神经元,编码动物在太空中的位置。对位置细胞的研究对于理解大脑神经网络对信息的处理至关重要。位置细胞尖峰序列的一个重要特征是相位进动。当动物在场地中奔跑时,场地细胞的放电从θ节律的上升期通过最小值转移到下降期。对沿Schaffer络和穿孔通路的锥体神经元的兴奋性输入在相位进动中的作用进行了描述,但对局部中间神经元的作用知之甚少。我们的目标是使用数学方法估计场CA1中间神经元对位置细胞相位进动的贡献。之所以选择CA1字段,是因为它提供了构建和验证模型所需的最大实验数据集。我们的模拟发现了锥体神经元兴奋性和抑制性输入的最佳参数,从而使其产生具有相位进动影响的尖峰序列。锥体神经元的均匀抑制最好地解释了相位进动的影响。在中间神经元中,轴突神经元对锥体细胞的抑制作用最大。
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引用次数: 0
Comparing performance between a deep neural network and monkeys with bilateral removals of visual area TE in categorizing feature-ambiguous stimuli. 比较深度神经网络和双侧去除视觉区域TE的猴子在对特征模糊刺激进行分类方面的表现。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-05-17 DOI: 10.1007/s10827-023-00854-y
Narihisa Matsumoto, Mark A G Eldridge, J Megan Fredericks, Kaleb A Lowe, Barry J Richmond

In the canonical view of visual processing the neural representation of complex objects emerges as visual information is integrated through a set of convergent, hierarchically organized processing stages, ending in the primate inferior temporal lobe. It seems reasonable to infer that visual perceptual categorization requires the integrity of anterior inferior temporal cortex (area TE). Many deep neural networks (DNNs) are structured to simulate the canonical view of hierarchical processing within the visual system. However, there are some discrepancies between DNNs and the primate brain. Here we evaluated the performance of a simulated hierarchical model of vision in discriminating the same categorization problems presented to monkeys with TE removals. The model was able to simulate the performance of monkeys with TE removals in the categorization task but performed poorly when challenged with visually degraded stimuli. We conclude that further development of the model is required to match the level of visual flexibility present in the monkey visual system.

在视觉处理的经典观点中,复杂物体的神经表示是随着视觉信息通过一组收敛的、分层组织的处理阶段整合而出现的,最终在灵长类动物的下颞叶结束。似乎可以合理地推断,视觉感知分类需要前颞下皮层(TE区)的完整性。许多深度神经网络(DNN)的结构是为了模拟视觉系统中层次处理的规范视图。然而,DNN和灵长类动物的大脑之间存在一些差异。在这里,我们评估了一个模拟的视觉层次模型在区分去除TE的猴子遇到的相同分类问题方面的性能。该模型能够模拟去除TE的猴子在分类任务中的表现,但在受到视觉退化刺激的挑战时表现不佳。我们得出的结论是,需要进一步开发该模型,以匹配猴子视觉系统中存在的视觉灵活性水平。
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引用次数: 0
Hierarchical processing underpins competition in tactile perceptual bistability. 分级处理是触觉-知觉双稳态竞争的基础。
IF 2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-05-19 DOI: 10.1007/s10827-023-00852-0
Farzaneh Darki, Andrea Ferrario, James Rankin

Ambiguous sensory information can lead to spontaneous alternations between perceptual states, recently shown to extend to tactile perception. The authors recently proposed a simplified form of tactile rivalry which evokes two competing percepts for a fixed difference in input amplitudes across antiphase, pulsatile stimulation of the left and right fingers. This study addresses the need for a tactile rivalry model that captures the dynamics of perceptual alternations and that incorporates the structure of the somatosensory system. The model features hierarchical processing with two stages. The first and the second stages of model could be located at the secondary somatosensory cortex (area S2), or in higher areas driven by S2. The model captures dynamical features specific to the tactile rivalry percepts and produces general characteristics of perceptual rivalry: input strength dependence of dominance times (Levelt's proposition II), short-tailed skewness of dominance time distributions and the ratio of distribution moments. The presented modelling work leads to experimentally testable predictions. The same hierarchical model could generalise to account for percept formation, competition and alternations for bistable stimuli that involve pulsatile inputs from the visual and auditory domains.

模糊的感官信息会导致感知状态之间的自发变化,最近显示这种变化延伸到触觉。作者最近提出了一种简化形式的触觉对抗,它唤起了两种相互竞争的感知,即在左手和右手手指的反相脉动刺激中输入振幅的固定差异。这项研究解决了对触觉竞争模型的需求,该模型捕捉感知变化的动态,并结合了体感系统的结构。该模型的特点是分两个阶段进行分层处理。模型的第一和第二阶段可以位于次级体感皮层(S2区域),或者位于S2驱动的更高区域。该模型捕捉了触觉竞争感知特有的动态特征,并产生了感知竞争的一般特征:支配时间的输入强度依赖性(Levelt命题II)、支配时间分布的短尾偏斜度和分布矩的比率。所提出的建模工作导致了可通过实验测试的预测。同样的层次模型可以广义地解释双稳态刺激的感知形成、竞争和交替,双稳态刺激涉及来自视觉和听觉领域的脉动输入。
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引用次数: 0
Transmission of delta band (0.5-4 Hz) oscillations from the globus pallidus to the substantia nigra pars reticulata in dopamine depletion. 多巴胺耗竭时从苍白球到黑质网状部的δ带(0.5-4 Hz)振荡的传输。
IF 1.5 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-06-02 DOI: 10.1007/s10827-023-00853-z
Timothy C Whalen, John E Parker, Aryn H Gittis, Jonathan E Rubin

Parkinson's disease (PD) and animal models of PD feature enhanced oscillations in several frequency bands in the basal ganglia (BG). Past research has emphasized the enhancement of 13-30 Hz beta oscillations. Recently, however, oscillations in the delta band (0.5-4 Hz) have been identified as a robust predictor of dopamine loss and motor dysfunction in several BG regions in mouse models of PD. In particular, delta oscillations in the substantia nigra pars reticulata (SNr) were shown to lead oscillations in motor cortex (M1) and persist under M1 lesion, but it is not clear where these oscillations are initially generated. In this paper, we use a computational model to study how delta oscillations may arise in the SNr due to projections from the globus pallidus externa (GPe). We propose a network architecture that incorporates inhibition in SNr from oscillating GPe neurons and other SNr neurons. In our simulations, this configuration yields firing patterns in model SNr neurons that match those measured in vivo. In particular, we see the spontaneous emergence of near-antiphase active-predicting and inactive-predicting neural populations in the SNr, which persist under the inclusion of STN inputs based on experimental recordings. These results demonstrate how delta oscillations can propagate through BG nuclei despite imperfect oscillatory synchrony in the source site, narrowing down potential targets for the source of delta oscillations in PD models and giving new insight into the dynamics of SNr oscillations.

帕金森病(PD)和帕金森病动物模型的特征是基底神经节(BG)中几个频带的振荡增强。过去的研究强调了13-30Hzβ振荡的增强。然而,最近,在PD小鼠模型中,δ带(0.5-4 Hz)的振荡已被确定为多巴胺损失和几个BG区域运动功能障碍的有力预测因子。特别是,黑质网状部(SNr)的δ振荡被证明会导致运动皮层(M1)的振荡,并在M1损伤下持续存在,但尚不清楚这些振荡最初是在哪里产生的。在本文中,我们使用一个计算模型来研究由于苍白球(GPe)的投影,SNr中可能会出现德尔塔振荡。我们提出了一种网络结构,该结构结合了振荡GPe神经元和其他SNr神经元对SNr的抑制。在我们的模拟中,这种配置在模型SNr神经元中产生的放电模式与体内测量的相匹配。特别是,我们在SNr中看到了近反相主动预测和非主动预测神经群体的自发出现,这些神经群体在基于实验记录的STN输入的情况下持续存在。这些结果表明,尽管震源位置的振荡同步性不完美,但德尔塔振荡如何通过BG核传播,缩小了PD模型中德尔塔振荡源的潜在目标,并对SNr振荡的动力学提供了新的见解。
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引用次数: 0
Correction to: Probabilistic solvers enable a straight-forward exploration of numerical uncertainty in neuroscience models. 更正:概率求解器能够直接探索神经科学模型中的数值不确定性。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 DOI: 10.1007/s10827-023-00856-w
Jonathan Oesterle, Nicholas Krämer, Philipp Hennig, Philipp Berens
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引用次数: 1
Selective neural stimulation by leveraging electrophysiological differentiation and using pre-pulsing and non-rectangular waveforms. 利用电生理分化以及预脉冲和非矩形波形,进行选择性神经刺激。
IF 2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2022-04-13 DOI: 10.1007/s10827-022-00818-8
Bemin Ghobreal, Farzan Nadim, Mesut Sahin

Efforts on selective neural stimulation have concentrated on segregating axons based on their size and geometry. Nonetheless, axons of the white matter or peripheral nerves may also differ in their electrophysiological properties. The primary objective of this study was to investigate the possibility of selective activation of axons by leveraging an assumed level of diversity in passive (Cm & Gleak) and active membrane properties (Ktemp & Gnamax). First, the stimulus waveforms with hyperpolarizing (HPP) and depolarizing pre-pulsing (DPP) were tested on selectivity in a local membrane model. The default value of membrane capacitance (Cm) was found to play a critical role in sensitivity of the chronaxie time (Chr) and rheobase (Rhe) to variations of all the four membrane parameters. Decreasing the default value of Cm, and thus the passive time constant of the membrane, amplified the sensitivity to the active parameters, Ktemp and GNamax, on Chr. The HPP waveform could selectively activate neurons even if they were diversified by membrane leakage (Gleak) only, and produced higher selectivity than DPP when parameters are varied in pairs. Selectivity measures were larger when the passive parameters (Cm & Gleak) were varied together, compared to the active parameters. Second, this novel mechanism of selectivity was investigated with non-rectangular waveforms for the stimulating phase (and HPP) in the same local membrane model. Simulation results suggest that Kt2 is the most selective waveform followed by Linear and Gaussian waveforms. Traditional rectangular pulse was among the least selective of all. Finally, a compartmental axon model confirmed the main findings of the local model that Kt2 is the most selective, but rank ordered the other waveforms differently. These results suggest a potentially novel mechanism of stimulation selectivity, leveraging electrophysiological variations in membrane properties, that can lead to various neural prosthetic applications.

选择性神经刺激的工作主要集中在根据轴突的大小和几何形状对其进行分离。然而,白质或周围神经的轴突在电生理特性上也可能存在差异。本研究的主要目的是利用被动膜特性(Cm 和 Gleak)和主动膜特性(Ktemp 和 Gnamax)的假定多样性水平,研究选择性激活轴突的可能性。首先,在局部膜模型中测试了超极化(HPP)和去极化预脉动(DPP)刺激波形的选择性。结果发现,膜电容(Cm)的默认值对计时时间(Chr)和流变基(Rhe)对所有四个膜参数变化的敏感性起着关键作用。降低 Cm 的默认值,从而降低膜的被动时间常数,会放大主动参数 Ktemp 和 GNamax 对 Chr 的敏感性。 HPP 波形可以选择性地激活神经元,即使它们只因膜泄漏(Gleak)而多样化,当参数成对变化时,其选择性高于 DPP。与主动参数相比,当被动参数(Cm 和 Gleak)一起变化时,选择性更强。其次,在相同的局部膜模型中,对刺激阶段(和 HPP)的非矩形波形研究了这种新的选择性机制。模拟结果表明,Kt2 是选择性最强的波形,其次是线性波形和高斯波形。传统矩形脉冲的选择性最小。最后,分区轴突模型证实了局部模型的主要发现,即 Kt2 的选择性最强,但对其他波形的排序有所不同。这些结果表明,利用膜特性的电生理变化,一种潜在的新型刺激选择性机制可能会带来各种神经假体应用。
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引用次数: 0
Improving a cortical pyramidal neuron model's classification performance on a real-world ecg dataset by extending inputs. 通过扩展输入改进皮层锥体神经元模型在真实世界心电图数据集上的分类性能。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-05-06 DOI: 10.1007/s10827-023-00851-1
Ilknur Kayikcioglu Bozkir, Zubeyir Ozcan, Cemal Kose, Temel Kayikcioglu, Ahmet Enis Cetin

Pyramidal neurons display a variety of active conductivities and complex morphologies that support nonlinear dendritic computation. Given growing interest in understanding the ability of pyramidal neurons to classify real-world data, in our study we applied both a detailed pyramidal neuron model and the perceptron learning algorithm to classify real-world ECG data. We used Gray coding to generate spike patterns from ECG signals as well as investigated the classification performance of the pyramidal neuron's subcellular regions. Compared with the equivalent single-layer perceptron, the pyramidal neuron performed poorly due to a weight constraint. A proposed mirroring approach for inputs, however, significantly boosted the classification performance of the neuron. We thus conclude that pyramidal neurons can classify real-world data and that the mirroring approach affects performance in a way similar to non-constrained learning.

金字塔神经元表现出各种活跃的导电性和复杂的形态,支持非线性树突计算。鉴于人们对理解锥体神经元对真实世界数据进行分类的能力越来越感兴趣,在我们的研究中,我们应用了详细的锥体神经元模型和感知器学习算法来对真实世界的ECG数据进行分类。我们使用格雷编码从ECG信号中生成尖峰模式,并研究了锥体神经元亚细胞区域的分类性能。与等效的单层感知器相比,由于权重限制,金字塔神经元表现不佳。然而,所提出的输入镜像方法显著提高了神经元的分类性能。因此,我们得出结论,锥体神经元可以对真实世界的数据进行分类,镜像方法以类似于非约束学习的方式影响性能。
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引用次数: 0
Functional architecture of M1 cells encoding movement direction. M1细胞编码运动方向的功能架构。
IF 1.2 4区 医学 Q3 MATHEMATICAL & COMPUTATIONAL BIOLOGY Pub Date : 2022-08-01 Epub Date: 2023-06-07 DOI: 10.1007/s10827-023-00850-2
Caterina Mazzetti, Alessandro Sarti, Giovanna Citti

In this paper we propose a neurogeometrical model of the behaviour of cells of the arm area of the primary motor cortex (M1). We will mathematically express as a fiber bundle the hypercolumnar organization of this cortical area, first modelled by Georgopoulos (Georgopoulos et al., 1982; Georgopoulos, 2015). On this structure, we will consider the selective tuning of M1 neurons of kinematic variables of positions and directions of movement. We will then extend this model to encode the notion of fragments introduced by Hatsopoulos et al. (2007) which describes the selectivity of neurons to movement direction varying in time. This leads to consider a higher dimensional geometrical structure where fragments are represented as integral curves. A comparison with the curves obtained through numerical simulations and experimental data will be presented. Moreover, neural activity shows coherent behaviours represented in terms of movement trajectories pointing to a specific pattern of movement decomposition Kadmon Harpaz et al. (2019). Here, we will recover this pattern through a spectral clustering algorithm in the subriemannian structure we introduced, and compare our results with the neurophysiological one of Kadmon Harpaz et al. (2019).

在本文中,我们提出了一个初级运动皮层(M1)臂区细胞行为的神经几何模型。我们将把该皮层区域的超体积组织以纤维束的形式进行数学表达,该组织首先由Georgopoulos建模(Georgepoulos等人,1982;Georgeopoulos,2015)。在这个结构上,我们将考虑运动位置和方向的运动学变量的M1神经元的选择性调谐。然后,我们将扩展这个模型,对Hatsopoulos等人引入的片段概念进行编码。(2007)描述了神经元对随时间变化的运动方向的选择性。这导致考虑更高维度的几何结构,其中碎片表示为积分曲线。将与通过数值模拟和实验数据获得的曲线进行比较。此外,神经活动显示出以运动轨迹表示的连贯行为,指向运动分解的特定模式Kadmon-Harpaz等人。(2019)。在这里,我们将通过我们引入的亚黎曼结构中的光谱聚类算法来恢复这种模式,并将我们的结果与Kadmon Harpaz等人的神经生理学结果进行比较。(2019)。
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引用次数: 3
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