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Neurally plausible mechanisms for learning selective and invariant representations. 学习选择性和不变表征的神经机制。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-08-18 DOI: 10.1186/s13408-020-00088-7
Fabio Anselmi, Ankit Patel, Lorenzo Rosasco

Coding for visual stimuli in the ventral stream is known to be invariant to object identity preserving nuisance transformations. Indeed, much recent theoretical and experimental work suggests that the main challenge for the visual cortex is to build up such nuisance invariant representations. Recently, artificial convolutional networks have succeeded in both learning such invariant properties and, surprisingly, predicting cortical responses in macaque and mouse visual cortex with unprecedented accuracy. However, some of the key ingredients that enable such success-supervised learning and the backpropagation algorithm-are neurally implausible. This makes it difficult to relate advances in understanding convolutional networks to the brain. In contrast, many of the existing neurally plausible theories of invariant representations in the brain involve unsupervised learning, and have been strongly tied to specific plasticity rules. To close this gap, we study an instantiation of simple-complex cell model and show, for a broad class of unsupervised learning rules (including Hebbian learning), that we can learn object representations that are invariant to nuisance transformations belonging to a finite orthogonal group. These findings may have implications for developing neurally plausible theories and models of how the visual cortex or artificial neural networks build selectivity for discriminating objects and invariance to real-world nuisance transformations.

已知腹侧流中视觉刺激的编码对保持对象身份的干扰变换是不变的。事实上,最近的许多理论和实验工作表明,视觉皮层面临的主要挑战是建立这种令人讨厌的不变表征。最近,人工卷积网络已经成功地学习了这些不变的特性,令人惊讶的是,它还以前所未有的准确性预测了猕猴和小鼠视觉皮层的皮层反应。然而,促成这种成功的一些关键因素——监督学习和反向传播算法——在神经学上是难以置信的。这使得很难将理解卷积网络的进展与大脑联系起来。相比之下,许多现有的关于大脑中不变表征的神经学理论都涉及无监督学习,并且与特定的可塑性规则密切相关。为了缩小这一差距,我们研究了一个简单-复杂单元模型的实例,并表明,对于一类广泛的无监督学习规则(包括Hebbian学习),我们可以学习对属于有限正交群的讨厌变换不变的对象表示。这些发现可能对发展神经上似是而非的理论和模型有启示意义,这些理论和模型是关于视觉皮层或人工神经网络如何建立选择性来区分物体和对现实世界中讨厌的转换的不变性。
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引用次数: 4
A sub-Riemannian model of the visual cortex with frequency and phase. 具有频率和相位的视觉皮层亚黎曼模型。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-07-29 DOI: 10.1186/s13408-020-00089-6
E Baspinar, A Sarti, G Citti

In this paper, we present a novel model of the primary visual cortex (V1) based on orientation, frequency, and phase selective behavior of V1 simple cells. We start from the first-level mechanisms of visual perception, receptive profiles. The model interprets V1 as a fiber bundle over the two-dimensional retinal plane by introducing orientation, frequency, and phase as intrinsic variables. Each receptive profile on the fiber is mathematically interpreted as rotated, frequency modulated, and phase shifted Gabor function. We start from the Gabor function and show that it induces in a natural way the model geometry and the associated horizontal connectivity modeling of the neural connectivity patterns in V1. We provide an image enhancement algorithm employing the model framework. The algorithm is capable of exploiting not only orientation but also frequency and phase information existing intrinsically in a two-dimensional input image. We provide the experimental results corresponding to the enhancement algorithm.

本文基于初级视觉皮层简单细胞的定向、频率和相位选择行为,提出了一种新的初级视觉皮层(V1)模型。我们从视觉感知的第一级机制,接受性轮廓开始。该模型通过引入方向、频率和相位作为固有变量,将V1解释为二维视网膜平面上的纤维束。光纤上的每个接收剖面在数学上被解释为旋转、调频和相移的Gabor函数。我们从Gabor函数开始,并证明它以自然的方式诱导了V1中神经连接模式的模型几何和相关的水平连接建模。我们提出了一种基于模型框架的图像增强算法。该算法不仅可以利用二维输入图像中固有的方向信息,还可以利用其固有的频率和相位信息。给出了与增强算法相对应的实验结果。
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引用次数: 10
Methods to assess binocular rivalry with periodic stimuli. 方法评价周期性刺激下的双眼竞争。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-06-15 DOI: 10.1186/s13408-020-00087-8
Farzaneh Darki, James Rankin

Binocular rivalry occurs when the two eyes are presented with incompatible stimuli and perception alternates between these two stimuli. This phenomenon has been investigated in two types of experiments: (1) Traditional experiments where the stimulus is fixed, (2) eye-swap experiments in which the stimulus periodically swaps between eyes many times per second (Logothetis et al. in Nature 380(6575):621-624, 1996). In spite of the rapid swapping between eyes, perception can be stable for many seconds with specific stimulus parameter configurations. Wilson introduced a two-stage, hierarchical model to explain both types of experiments (Wilson in Proc. Natl. Acad. Sci. 100(24):14499-14503, 2003). Wilson's model and other rivalry models have been only studied with bifurcation analysis for fixed inputs and different types of dynamical behavior that can occur with periodically forcing inputs have not been investigated. Here we report (1) a more complete description of the complex dynamics in the unforced Wilson model, (2) a bifurcation analysis with periodic forcing. Previously, bifurcation analysis of the Wilson model with fixed inputs has revealed three main types of dynamical behaviors: Winner-takes-all (WTA), Rivalry oscillations (RIV), Simultaneous activity (SIM). Our results have revealed richer dynamics including mixed-mode oscillations (MMOs) and a period-doubling cascade, which corresponds to low-amplitude WTA (LAWTA) oscillations. On the other hand, studying rivalry models with numerical continuation shows that periodic forcing with high frequency (e.g. 18 Hz, known as flicker) modulates the three main types of behaviors that occur with fixed inputs with forcing frequency (WTA-Mod, RIV-Mod, SIM-Mod). However, dynamical behavior will be different with low frequency periodic forcing (around 1.5 Hz, so-called swap). In addition to WTA-Mod and SIM-Mod, cycle skipping, multi-cycle skipping and chaotic dynamics are found. This research provides a framework for either assessing binocular rivalry models to check consistency with empirical results, or for better understanding neural dynamics and mechanisms necessary to implement a minimal binocular rivalry model.

双眼竞争发生时,两只眼睛呈现不相容的刺激和感知交替这两个刺激。这种现象已经在两种类型的实验中进行了研究:(1)刺激固定的传统实验,(2)眼睛交换实验,刺激每秒在眼睛之间周期性地交换多次(Logothetis et al. in Nature 380(6575):621-624, 1996)。尽管眼睛之间的快速交换,知觉可以在特定的刺激参数配置下稳定数秒。威尔逊引入了一个两阶段的分层模型来解释这两种类型的实验(威尔逊在Proc. Natl。科学通报,2009(4):379 - 379。威尔逊的模型和其他竞争模型只研究了固定输入的分岔分析,而周期性强迫输入可能发生的不同类型的动态行为尚未研究。在这里,我们报告(1)在非强制Wilson模型中更完整地描述了复杂动力学,(2)具有周期强迫的分岔分析。在此之前,固定输入的Wilson模型的分岔分析揭示了三种主要的动态行为:赢家通吃(WTA)、竞争振荡(RIV)和同步活动(SIM)。我们的研究结果揭示了更丰富的动力学,包括混合模式振荡(MMOs)和周期倍级联,对应于低振幅WTA (LAWTA)振荡。另一方面,研究具有数值延续的竞争模式表明,高频周期性强迫(例如18 Hz,称为闪烁)调节了固定频率强迫输入(WTA-Mod, RIV-Mod, SIM-Mod)时发生的三种主要行为。然而,在低频周期性强迫(约1.5 Hz,即所谓的交换)下,动力学行为将有所不同。除了WTA-Mod和SIM-Mod之外,还发现了周期跳跃、多周期跳跃和混沌动力学。本研究为评估双眼竞争模型以检查与实证结果的一致性,或更好地理解实现最小双眼竞争模型所需的神经动力学和机制提供了一个框架。
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引用次数: 4
Understanding the dynamics of biological and neural oscillator networks through exact mean-field reductions: a review. 通过精确平均场还原理解生物和神经振荡器网络的动力学:综述。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-05-27 DOI: 10.1186/s13408-020-00086-9
Christian Bick, Marc Goodfellow, Carlo R Laing, Erik A Martens

Many biological and neural systems can be seen as networks of interacting periodic processes. Importantly, their functionality, i.e., whether these networks can perform their function or not, depends on the emerging collective dynamics of the network. Synchrony of oscillations is one of the most prominent examples of such collective behavior and has been associated both with function and dysfunction. Understanding how network structure and interactions, as well as the microscopic properties of individual units, shape the emerging collective dynamics is critical to find factors that lead to malfunction. However, many biological systems such as the brain consist of a large number of dynamical units. Hence, their analysis has either relied on simplified heuristic models on a coarse scale, or the analysis comes at a huge computational cost. Here we review recently introduced approaches, known as the Ott-Antonsen and Watanabe-Strogatz reductions, allowing one to simplify the analysis by bridging small and large scales. Thus, reduced model equations are obtained that exactly describe the collective dynamics for each subpopulation in the oscillator network via few collective variables only. The resulting equations are next-generation models: Rather than being heuristic, they exactly link microscopic and macroscopic descriptions and therefore accurately capture microscopic properties of the underlying system. At the same time, they are sufficiently simple to analyze without great computational effort. In the last decade, these reduction methods have become instrumental in understanding how network structure and interactions shape the collective dynamics and the emergence of synchrony. We review this progress based on concrete examples and outline possible limitations. Finally, we discuss how linking the reduced models with experimental data can guide the way towards the development of new treatment approaches, for example, for neurological disease.

许多生物和神经系统都可以看作是由相互作用的周期性过程组成的网络。重要的是,它们的功能性,即这些网络能否发挥其功能,取决于网络新出现的集体动力学。振荡的同步性是这种集体行为最突出的例子之一,既与功能有关,也与功能障碍有关。了解网络结构和相互作用以及单个单元的微观特性如何塑造新出现的集体动力学,对于找到导致功能失调的因素至关重要。然而,许多生物系统(如大脑)由大量动态单元组成。因此,对它们的分析要么依赖于粗略尺度上的简化启发式模型,要么需要付出巨大的计算成本。在此,我们回顾了最近引入的方法,即所谓的奥特-安通森和瓦塔纳贝-斯特罗加茨还原法,它允许人们通过连接小尺度和大尺度来简化分析。这样,就得到了简化的模型方程,这些方程仅通过少数几个集体变量就能精确描述振荡器网络中每个子群的集体动力学。由此得到的方程是新一代模型:它们不是启发式的,而是精确地将微观和宏观描述联系起来,因此能准确捕捉底层系统的微观特性。同时,这些模型非常简单,无需大量计算即可进行分析。近十年来,这些还原方法在理解网络结构和相互作用如何塑造集体动力学和同步性的出现方面发挥了重要作用。我们根据具体实例回顾了这一进展,并概述了可能存在的局限性。最后,我们将讨论如何将简化模型与实验数据联系起来,从而为开发新的治疗方法(如神经疾病)提供指导。
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引用次数: 0
Spatially extended balanced networks without translationally invariant connectivity. 没有平移不变连通性的空间扩展平衡网络。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-05-13 DOI: 10.1186/s13408-020-00085-w
Christopher Ebsch, Robert Rosenbaum

Networks of neurons in the cerebral cortex exhibit a balance between excitation (positive input current) and inhibition (negative input current). Balanced network theory provides a parsimonious mathematical model of this excitatory-inhibitory balance using randomly connected networks of model neurons in which balance is realized as a stable fixed point of network dynamics in the limit of large network size. Balanced network theory reproduces many salient features of cortical network dynamics such as asynchronous-irregular spiking activity. Early studies of balanced networks did not account for the spatial topology of cortical networks. Later works introduced spatial connectivity structure, but were restricted to networks with translationally invariant connectivity structure in which connection probability depends on distance alone and boundaries are assumed to be periodic. Spatial connectivity structure in cortical network does not always satisfy these assumptions. We use the mathematical theory of integral equations to extend the mean-field theory of balanced networks to account for more general dependence of connection probability on the spatial location of pre- and postsynaptic neurons. We compare our mathematical derivations to simulations of large networks of recurrently connected spiking neuron models.

大脑皮层的神经元网络表现出兴奋(正输入电流)和抑制(负输入电流)之间的平衡。平衡网络理论为这种兴奋-抑制平衡提供了一个简洁的数学模型,该模型使用随机连接的模型神经元网络,在大网络规模的限制下,平衡作为网络动力学的稳定不动点实现。平衡网络理论再现了皮层网络动力学的许多显著特征,如异步-不规则尖峰活动。早期对平衡网络的研究没有考虑到皮质网络的空间拓扑结构。后来的研究引入了空间连接结构,但仅限于具有平动不变连接结构的网络,其中连接概率仅取决于距离,并且假定边界是周期性的。皮层网络的空间连通性结构并不总是满足这些假设。我们使用积分方程的数学理论来扩展平衡网络的平均场理论,以说明连接概率对突触前和突触后神经元的空间位置的更一般的依赖。我们将我们的数学推导与循环连接的脉冲神经元模型的大型网络的模拟进行比较。
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引用次数: 0
Geometry of color perception. Part 1: structures and metrics of a homogeneous color space. 色彩感知的几何学。第1部分:均匀色彩空间的结构和度量。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-05-12 DOI: 10.1186/s13408-020-00084-x
Edoardo Provenzi

This is the first half of a two-part paper dealing with the geometry of color perception. Here we analyze in detail the seminal 1974 work by H.L. Resnikoff, who showed that there are only two possible geometric structures and Riemannian metrics on the perceived color space [Formula: see text] compatible with the set of Schrödinger's axioms completed with the hypothesis of homogeneity. We recast Resnikoff's model into a more modern colorimetric setting, provide a much simpler proof of the main result of the original paper, and motivate the need of psychophysical experiments to confute or confirm the linearity of background transformations, which act transitively on [Formula: see text]. Finally, we show that the Riemannian metrics singled out by Resnikoff through an axiom on invariance under background transformations are not compatible with the crispening effect, thus motivating the need of further research about perceptual color metrics.

这是一篇由两部分组成的论文的第一部分,该论文涉及颜色感知的几何。在这里,我们详细分析1974年由H.L. Resnikoff的开创性工作,他表明在感知颜色空间上只有两种可能的几何结构和黎曼度量(公式:见文本)与Schrödinger的公理集兼容,该公理集由同质性假设完成。我们将Resnikoff的模型重新塑造成一个更现代的色度设置,为原始论文的主要结果提供了一个更简单的证明,并激发了心理物理实验的需要,以反驳或证实背景变换的线性,背景变换对[公式:见文本]起传递作用。最后,我们证明了Resnikoff通过背景变换下的不变性公理挑选出的黎曼度量与脆化效果不兼容,从而激发了对感知颜色度量的进一步研究的必要性。
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引用次数: 25
Correction to: Linking demyelination to compound action potential dispersion with a spike-diffuse-spike approach. 更正:用尖峰-扩散-尖峰方法将脱髓鞘与复合动作电位分散联系起来。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-04-20 DOI: 10.1186/s13408-020-00083-y
Richard Naud, André Longtin

Following publication of the original article (Naud and Longtin in J Math Neurosci 9:3, 2019), the authors noticed a mistake in the first paragraph within "Altered propagation".

在原始文章发表后(Naud和Longtin在J Math Neurosci, 2019年9月3日),作者注意到“Altered propagation”的第一段中有一个错误。
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引用次数: 0
Mesoscopic population equations for spiking neural networks with synaptic short-term plasticity. 具有突触短期可塑性的脉冲神经网络的介观种群方程。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-04-06 DOI: 10.1186/s13408-020-00082-z
Valentin Schmutz, Wulfram Gerstner, Tilo Schwalger

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). The extended theory offers an approximate mean-field dynamics for the synaptic input currents arising from populations of spiking neurons and synapses undergoing Tsodyks-Markram STP. The approximate mean-field dynamics accounts for both finite number of synapses and correlation between the two synaptic variables of the model (utilization and available resources) and its numerical implementation is simple. Comparisons with Monte Carlo simulations of the microscopic model show that in both feedforward and recurrent networks, the mesoscopic mean-field model accurately reproduces the first- and second-order statistics of the total synaptic input into a postsynaptic neuron and accounts for stochastic switches between Up and Down states and for population spikes. The extended mesoscopic population theory of spiking neural networks with STP may be useful for a systematic reduction of detailed biophysical models of cortical microcircuits to numerically efficient and mathematically tractable mean-field models.

从生物神经网络的粗粒度微观模型中获得神经活动的介观模型是迈向大脑多尺度模型的重要一步。在这里,我们将静态突触的介观种群动态理论扩展到具有短期可塑性(STP)的动态突触的情况。扩展理论提供了一个近似的平均场动力学的突触输入电流产生的群体尖峰神经元和突触经历Tsodyks-Markram STP。近似平均场动力学既考虑了有限突触数,又考虑了模型的两个突触变量(利用率和可用资源)之间的相关性,其数值实现简单。与蒙特卡罗模拟微观模型的比较表明,在前馈和循环网络中,介观平均场模型准确地再现了突触后神经元总突触输入的一阶和二阶统计量,并解释了上下状态之间的随机切换和种群峰值。带STP的脉冲神经网络的扩展介观种群理论可能有助于系统地将皮层微电路的详细生物物理模型简化为数值上有效且数学上易于处理的平均场模型。
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引用次数: 20
Phase-dependence of response curves to deep brain stimulation and their relationship: from essential tremor patient data to a Wilson-Cowan model. 深部脑刺激反应曲线的相位依赖性及其关系:从本质性震颤患者数据到威尔逊-考恩模型。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-03-30 DOI: 10.1186/s13408-020-00081-0
Benoit Duchet, Gihan Weerasinghe, Hayriye Cagnan, Peter Brown, Christian Bick, Rafal Bogacz

Essential tremor manifests predominantly as a tremor of the upper limbs. One therapy option is high-frequency deep brain stimulation, which continuously delivers electrical stimulation to the ventral intermediate nucleus of the thalamus at about 130 Hz. Constant stimulation can lead to side effects, it is therefore desirable to find ways to stimulate less while maintaining clinical efficacy. One strategy, phase-locked deep brain stimulation, consists of stimulating according to the phase of the tremor. To advance methods to optimise deep brain stimulation while providing insights into tremor circuits, we ask the question: can the effects of phase-locked stimulation be accounted for by a canonical Wilson-Cowan model? We first analyse patient data, and identify in half of the datasets significant dependence of the effects of stimulation on the phase at which stimulation is provided. The full nonlinear Wilson-Cowan model is fitted to datasets identified as statistically significant, and we show that in each case the model can fit to the dynamics of patient tremor as well as to the phase response curve. The vast majority of top fits are stable foci. The model provides satisfactory prediction of how patient tremor will react to phase-locked stimulation by predicting patient amplitude response curves although they were not explicitly fitted. We also approximate response curves of the significant datasets by providing analytical results for the linearisation of a stable focus model, a simplification of the Wilson-Cowan model in the stable focus regime. We report that the nonlinear Wilson-Cowan model is able to describe response to stimulation more precisely than the linearisation.

本质性震颤主要表现为上肢震颤。高频深部脑刺激是一种治疗方法,它以大约 130 赫兹的频率持续向丘脑腹侧中间核提供电刺激。持续的刺激可能会导致副作用,因此希望找到既能减少刺激又能保持临床疗效的方法。其中一种策略是锁定相位的脑深部刺激,包括根据震颤的相位进行刺激。为了推进优化深部脑刺激的方法,同时深入了解震颤回路,我们提出了这样一个问题:锁定相位刺激的效果可以用典型的威尔逊-考恩模型来解释吗?我们首先分析了患者数据,发现在一半的数据集中,刺激效果与提供刺激的相位有显著的相关性。我们将完整的非线性威尔逊-科文(Wilson-Cowan)模型拟合到被确定为具有统计学意义的数据集中,结果表明,在每种情况下,该模型都能拟合患者震颤的动态以及相位响应曲线。绝大多数顶部拟合都是稳定的病灶。虽然没有明确拟合患者的振幅反应曲线,但该模型通过预测患者的振幅反应曲线,令人满意地预测了患者震颤对锁相刺激的反应。我们还通过提供稳定病灶模型线性化的分析结果,对重要数据集的反应曲线进行了近似分析。我们报告说,非线性威尔逊-科文模型比线性化模型能够更精确地描述对刺激的反应。
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引用次数: 0
Sparse identification of contrast gain control in the fruit fly photoreceptor and amacrine cell layer. 果蝇光感受器和无毛细胞层对比度增益控制的稀疏识别。
IF 2.3 4区 医学 Q1 Neuroscience Pub Date : 2020-02-12 DOI: 10.1186/s13408-020-0080-5
Aurel A Lazar, Nikul H Ukani, Yiyin Zhou

The fruit fly's natural visual environment is often characterized by light intensities ranging across several orders of magnitude and by rapidly varying contrast across space and time. Fruit fly photoreceptors robustly transduce and, in conjunction with amacrine cells, process visual scenes and provide the resulting signal to downstream targets. Here, we model the first step of visual processing in the photoreceptor-amacrine cell layer. We propose a novel divisive normalization processor (DNP) for modeling the computation taking place in the photoreceptor-amacrine cell layer. The DNP explicitly models the photoreceptor feedforward and temporal feedback processing paths and the spatio-temporal feedback path of the amacrine cells. We then formally characterize the contrast gain control of the DNP and provide sparse identification algorithms that can efficiently identify each the feedforward and feedback DNP components. The algorithms presented here are the first demonstration of tractable and robust identification of the components of a divisive normalization processor. The sparse identification algorithms can be readily employed in experimental settings, and their effectiveness is demonstrated with several examples.

果蝇的自然视觉环境通常以光强度为特征,光强度范围跨越几个数量级,并且在空间和时间上快速变化对比度。果蝇的光感受器强有力地传递并与无毛细胞一起处理视觉场景,并将产生的信号提供给下游目标。在这里,我们模拟了视觉处理的第一步在光感受器-腺分泌细胞层。我们提出了一种新的分裂归一化处理器(DNP)来模拟发生在光感受器-腺细胞层的计算。DNP明确地模拟了无毛细胞的光感受器前馈和时间反馈加工路径以及时空反馈路径。然后,我们正式表征了DNP的对比度增益控制,并提供了稀疏识别算法,可以有效地识别每个前馈和反馈DNP组件。这里提出的算法是分裂归一化处理器组件的可处理和鲁棒识别的第一个演示。稀疏识别算法可以很容易地应用于实验环境,并通过几个例子证明了它们的有效性。
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引用次数: 8
期刊
Journal of Mathematical Neuroscience
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