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[病例讨论] 纹状体

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发表于 2019-5-6 00:00:19 | 显示全部楼层 |阅读模式
纹状体[5](也称为新纹状体和纹状核)是前脑皮质下基底神经节中的细胞核(神经元簇)。纹状体是运动和奖励系统的关键组成部分;接受来自不同来源的谷氨酸能和多巴胺能输入;并作为其他基底神经节的主要输入。

在功能上,纹状体协调认知的多个方面,包括运动和行动计划,决策,动机,强化和奖励感知。[2] [3] [4]纹状体由尾状核和豆状核组成。[6] [7]豆状核由较大的壳核和较小的苍白球构成。[8]

在灵长类动物中,纹状体分为腹侧纹状体和背侧纹状体,基于功能和连接的细分。腹侧纹状体由伏隔核和嗅结节组成。背侧纹状体由尾状核和壳核组成。背侧纹状体中的白质,神经束(内囊)将尾状核和壳核分开。[4]在解剖学上,术语纹状体描述了灰色和白色物质的条纹(条纹)外观。[9]


视频:2分钟神经科学 - 纹状体


紫色=尾状和壳核,橙色=丘脑


显示皮质纹状体连接的纤维束成像

目录
1 结构
1.1 细胞类型
1.2 输入
1.3 目标
2 功能
3 临床意义
3.1 帕金森病
3.2 成瘾
3.3 双相情感障碍
3.4 自闭症谱系障碍
3.5 功能障碍
4 历史
5 其他动物
6 参考

结构
在MRI上看到的纹状体。 纹状体包括尾状核和豆状核,其中包括壳核和苍白球。


在MRI上看到的红色纹状体。纹状体包括尾状核(顶部)和豆状核(壳核(右)和左下苍白球)
纹状体是基底神经节的最大结构。根据功能和连接,纹状体分为腹侧和背侧分区。

腹侧纹状体由伏隔核和嗅结节组成。[4] [10]伏隔核由伏隔核和伏隔核组成,它们因神经群体而不同。嗅球结节接受来自嗅球的输入,但未被证明在处理气味方面起作用。[10]在非灵长类物种中,包括Calleja岛屿。[11]腹侧纹状体与边缘系统相关,并且被认为是决策和奖励相关行为的重要部分。[12] [13]

背侧纹状体由尾状核和壳核组成。

染色可以将纹状体分化为不同的区域,即纹状体或斑块和周围的基质;这在乙酰胆碱酯酶和钙结合蛋白的组分上尤其明显。对背侧纹状体进行了更多的研究,但在腹侧纹状体中也发现了隔室。在背侧纹状体中,纹状体占纹状体体积的10-15%。[14]

细胞类型


树突状棘刺在纹状体的中型多刺神经元上
纹状体中的细胞类型包括:

中型多刺神经元(MSNs),是纹状体的主要神经元。[2]它们是GABA能的,因此被归类为抑制性神经元。中型多刺投射神经元占人纹状体总神经元群体的95%[2]。中型多刺神经元有两种特征类型:D1型MSN和D2型MSN。[2] [4] [15] MSN亚群包含D1型和D2型受体,大约40%的纹状体MSN表达DRD1和DRD2 mRNA [2] [4] [15]
胆碱能中间神经元释放乙酰胆碱,其在纹状体中具有多种重要作用。在人类,其他灵长类动物和啮齿动物中,这些中间神经元对突出的环境刺激作出反应,其定型反应与黑质多巴胺能神经元的反应在时间上一致。[16] [17]大型阿司匹林胆碱能中间神经元本身受多巴胺通过D5多巴胺受体的影响。[18]
有许多类型的GABAergic中间神经元。[19]最着名的是表达小球蛋白的中间神经元,也称为快速中间神经元,它们参与主要神经元的强大前馈抑制。[20]此外,还有GABAergic中间神经元表达酪氨酸羟化酶,[21]生长抑素,一氧化氮合酶和神经肽-y。最近,已经详细描述了两种表达神经肽y的GABA能中间神经元,[22]其中一种将胆碱能中间神经元的同步活动转化为主要神经元的抑制[23]。纹状体的这些神经元不均匀分布。[19]
大脑中存在两个神经发生区域 - 侧脑室的脑室下区和齿状回。在与纹状体相邻的侧脑室中形成的成神经细胞整合在纹状体中。[24] [25]在缺血性中风后的人纹状体中已经注意到这一点。对纹状体造成的损伤刺激成神经细胞从脑室下区迁移到纹状体,在那里它们分化成成体神经元。[26] SVZ成神经细胞正常通过嗅球,但这种交通在缺血性中风后转移到纹状体。然而,很少有新发现的神经元存活。[27]

输入


额叶皮层到纹状体到丘脑通路的简化图 - 前纹状体回路


基底神经节的主要电路概述。纹状体以蓝色显示。图片显示已叠加的2个冠状切片,包括所涉及的基底神经节结构。 +和 - 箭头处的符号分别表示该途径是兴奋的还是抑制的。绿色箭头指的是兴奋性谷氨酸能通路,红色箭头指的是抑制性GABA能通路,绿松石箭头指的是直接通路上兴奋的多巴胺能通路和间接通路的抑制作用。
[28]就细胞轴突而言,最大的连接来自皮层。新皮质的许多部分支配背侧纹状体。投射到纹状体的皮层锥体神经元位于II-VI层,最密集的投影来自V层。[29]它们主要终止于多刺神经元的树突棘。它们是谷氨酸能的,令人兴奋的纹状体神经元。

纹状体被视为具有自己的内部微电路。[30]腹侧纹状体接受来自大脑皮层和边缘结构的多个区域的直接输入,例如杏仁核,丘脑和海马,以及内嗅皮层和颞下回。[31]它的主要输入是基底神经节系统。此外,中脑边缘通路从腹侧被盖区突出到腹侧纹状体的伏隔核[32]。

另一个众所周知的传入是由黑质致密部神经元引起的黑质纹状体连接。虽然皮质轴突主要在多刺神经元的脊柱头上突触,但黑质轴突突触主要在脊柱轴上。在灵长类动物中,丘脑纹状体传入来自丘脑的中央 - 束旁复合体(参见灵长类动物基底神经节系统)。这种传入是谷氨酸能的。真正的层内神经元的参与更加有限。纹状体还接受来自基底神经节的其他元件的传入,例如丘脑底核(谷氨酸能)或外部苍白球(GABA能)。

目标
更多信息:中型多刺神经元
腹侧纹状体的主要输出突出到腹侧苍白球,然后是丘脑的内侧背核,这是前纹状体回路的一部分。此外,腹侧纹状体突出到苍白球和黑质网状物。其他一些产出包括对延长的杏仁核,下丘脑外侧和脑桥脑核的投射。[33]

来自背侧和腹侧组成部分的纹状体输出主要由中型多刺神经元(MSN)组成,这是一种投射神经元,具有两种主要表型:表达D2样受体的“间接”MSN和表达D1的“直接”MSN。类受体[2] [4]

基底神经节的主要核是纹状体,通过纹状体纤维通路直接投射到苍白球。[34]由于有髓纤维,striato-pallidal路径具有发白的外观。该投射依次包括外部苍白球(GPe),内部苍白球(GPi),黑质致密部(SNc)和黑质网状结构(SNr)。这种投射的神经元受到来自背侧纹状体的GABA能神经突触的抑制。在这些目标中,GPe不会在系统外发送轴突。其他人将轴突送到上丘。另外两个包括丘脑的输出,形成两个独立的通道:一个通过苍白球的内部区域到丘脑的腹侧口腔核,从那里到皮质辅助运动区域,另一个通过黑质到腹侧前部。丘脑的核,从那里到额叶皮质和动眼神经皮质。

功能
腹侧纹状体,特别是伏核,主要介导奖赏,认知,强化和动机显着性,而背侧纹状体主要介导涉及运动功能,某些执行功能(如抑制控制和冲动)和刺激反应的认知。学习; [2] [3] [4] [35] [36]存在一定程度的重叠,因为背侧纹状体也是奖励系统的一个组成部分,与伏隔核一起,调节编码与未来奖励获取相关的新运动程序(例如,条件运动对奖励线索的反应)。[3] [35]

代谢型多巴胺受体存在于多刺神经元和皮质轴突末端上。由激活这些多巴胺受体引发的第二信使级联可以在短期和长期内调节突触前和突触后功能。[37] [38]在人类中,纹状体由与奖赏相关的刺激激活,但也受到厌恶,新颖,[39]意外或强烈刺激以及与此类事件相关的线索的激活。[40] fMRI证据表明,纹状体反应的这些刺激物的共同特性在呈现条件下是显著的。[41] [42]许多其他大脑区域和电路也与奖励相关,例如额叶区域。纹状体的功能图揭示了与广泛分布的大脑皮层区域的相互作用,这些区域对于各种功能是重要的。[43]

纹状体和前额皮质之间的相互作用与行为有关,特别是双系统模型提出的青少年发育。[44]

临床意义
帕金森病
帕金森病导致背侧纹状体(和其他基底神经节)的多巴胺能神经支配丧失和一系列后果。纹状体萎缩还涉及亨廷顿氏病和运动障碍,如舞蹈病,舞蹈病和运动障碍。[45]这些也被描述为基底神经节的电路紊乱。[46]

成瘾
成瘾是一种大脑奖赏系统的紊乱,是由于腹侧纹状体D1型中型多刺神经元中转录因子DeltaFosB(ΔFosB)的过度表达而产生的。 ΔFosB是一种诱导型基因,由于反复过量服用成瘾药物或过度暴露于其他成瘾性刺激物而逐渐在伏隔核中表达[47] [48]。

双相情感障碍
已经观察到PDE10A基因的变体的纹状体表达与一些I型双相障碍患者之间的关联。其他基因的变异,DISC1和GNAS,与双相II障碍有关。[49]

自闭症谱系障碍
自闭症谱系障碍(ASD)的特点是认知缺乏能力和对社会系统的理解不足。这种不灵活的行为起源于前额叶皮质以及纹状体回路中的缺陷。[50]纹状体缺陷似乎特别有助于ASD患者的运动,社交和沟通障碍。在通过翻译因子4E的真核起始过表达而诱导具有ASD样表型的小鼠中,已经表明这些缺陷似乎源于在纹状体中储存和处理信息的能力降低,这导致了困难。从形成新的电机模式,以及脱离现有电机模式。[51]

功能障碍
腹侧纹状体功能障碍可导致多种疾病,最显着的是抑郁症和强迫症。由于其参与奖励途径,腹侧纹状体也涉及在成瘾中起关键作用。已经确定腹侧纹状体通过多巴胺能刺激强烈参与介导药物,特别是兴奋剂的增强作用。[52]

历史
在十七和十八世纪,术语“纹状体”被用来指定半球的许多不同的,深的,皮下的元素。[53] 1941年,Cécile和Oskar Vogt通过为纹状体元素构建的所有元素(参见灵长类动物基底神经节系统)提出术语纹状体来简化命名:尾状核,壳核和眼底纹[54]腹侧部分连接两者一起腹侧向内囊的下部。

比较解剖学家比较脊椎动物之间的皮质下结构这一术语neostriatum是伪造的,因为它被认为是纹状体的系统发育上较新的部分。 该术语仍被某些来源使用,包括医学主题标题。[55]

其他动物
在鸟类中,使用的术语是古纹状体,并且在新的鸟类术语列表(截至2002年)中,新纹状体已经改为nidopallium。[56]

在非灵长类物种中,Calleja岛包括在腹侧纹状体中。[11]

另见
Cortico-basal ganglia-thalamo-cortical loop
List of regions in the human brain
Striatopallidal fibres

参考
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Functional neuroimaging in humans demonstrates activation of the prefrontal cortex and caudate nucleus (part of the striatum) in tasks that demand inhibitory control of behavior. ...
The brain reward circuitry that is targeted by addictive drugs normally mediates the pleasure and strengthening of behaviors associated with natural reinforcers, such as food, water, and sexual contact. Dopamine neurons in the VTA are activated by food and water, and dopamine release in the NAc is stimulated by the presence of natural reinforcers, such as food, water, or a sexual partner. ...
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Table 1
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