Pay attention to the selection of mammals commonly used in animal experiments for the study on visual mechanism

doi:10.3877/cma.j.issn.2095-2007.2019.04.001

Abstract

Abstract:

The understanding of retinal ganglion cells (RGC) is an important premise for us to explore the visual mechanism. In light of various kinds of RGC, the classification and naming methods of them are more complicated and non-uniform in different mammalian species. On the one hand, there are intrinsic connections and commonalities between different classification methods, suggesting the homology of visual signal transmission in the evolution of species; on the other hand, the distribution of ganglion cells on the retina in different species has a similar pattern. Therefore, it is necessary to pick up the difference and simlarity among various mammals in the classification of RGC. This review describes the classification and characteristics of ganglion cells in several types of species that are important in retinal research, and summarizes the intrinsic homology between different classification methods, which helps us to deepen the understanding of the visual formation process.

Key words: Mammal, Retina, Ganglion cell, Classification

Jiajia Wang, Yining Guo, Xuemin Li. Pay attention to the selection of mammals commonly used in animal experiments for the study on visual mechanism[J]. Chinese Journal of Ophthalmologic Medicine(Electronic Edition), 2019, 09(04): 193-198.

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References 46

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细胞类型	别称	形态	亚类	同源性	分布	通路	功能
P型神经节细胞	侏儒节细胞(midget cell)	中小胞体，小树突野	—	灵长类动物特化的结果	主要集中在中央凹，随离心率增加，占比减少	投射到膝状体小细胞层的P通路	传导有色视觉，传导L和M-视锥细胞的红色、绿色信号(中长波敏感)；感受野较小，高空间敏感性，可对静止的物体进行精细感知；低时间敏感性，反应速度较慢；低对比增益
M型神经节细胞	伞状节细胞(parasol cell)	大胞体，大树突野	光滑单分层神经节细胞	与猫α-Y神经节细胞同源，与兔G11型神经节细胞同源	主要分布在视网膜的周边部，随离心率增加，占比增大	投射到膝状体大细胞层的M通路	传导无色视觉，感受野较大；低空间敏感性，对细节信息感知能力较弱，可对物体的运动信息进行有效感知；高时间敏感性，反应速度较快，高对比增益
K型神经节细胞	双分层神经节细胞	树突在视网膜内丛状层中分层	小型双分层神经节细胞；大型双分层神经节细胞；弥漫性神经节细胞	可能与兔G3型神经节细胞同源	—	小型双分层神经节细胞对应K细胞通路	小型双分层神经节细胞与颜色感知有关，传递S-视锥细胞的蓝色信号(短波敏感)

细胞类型	别称	形态	亚类	同源性	分布	通路	功能
P型神经节细胞	侏儒节细胞(midget cell)	中小胞体，小树突野	—	灵长类动物特化的结果	主要集中在中央凹，随离心率增加，占比减少	投射到膝状体小细胞层的P通路	传导有色视觉，传导L和M-视锥细胞的红色、绿色信号(中长波敏感)；感受野较小，高空间敏感性，可对静止的物体进行精细感知；低时间敏感性，反应速度较慢；低对比增益
M型神经节细胞	伞状节细胞(parasol cell)	大胞体，大树突野	光滑单分层神经节细胞	与猫α-Y神经节细胞同源，与兔G11型神经节细胞同源	主要分布在视网膜的周边部，随离心率增加，占比增大	投射到膝状体大细胞层的M通路	传导无色视觉，感受野较大；低空间敏感性，对细节信息感知能力较弱，可对物体的运动信息进行有效感知；高时间敏感性，反应速度较快，高对比增益
K型神经节细胞	双分层神经节细胞	树突在视网膜内丛状层中分层	小型双分层神经节细胞；大型双分层神经节细胞；弥漫性神经节细胞	可能与兔G3型神经节细胞同源	—	小型双分层神经节细胞对应K细胞通路	小型双分层神经节细胞与颜色感知有关，传递S-视锥细胞的蓝色信号(短波敏感)

电生理学分类	形态学分类	亚类	同源性	功能特征
X型神经节细胞	β细胞：中型细胞体，小树突野	β-ON和β-OFF两类	与兔G4型神经节细胞同源	小感受野，对亮度和空间线性刺激产生持续反应，传递速度慢，可传递颜色、精细图案信号；成像视觉
Y型神经节细胞	α细胞：大型细胞体，大树突野	α-ON和α-OFF两类	与灵长类M神经节细胞同源；与兔G11型神经节细胞同源	大感受野，对空间非线性刺激产生瞬态反应(倍频响应)，传递速度快，可传递运动信息；非成像视觉
W型神经节细胞	γ细胞：中小型细胞体，大树突野	η细胞：小型细胞体，中等树突野；θ细胞：小型细胞体，中等树突野；θ-ON和θ-OFF两类	γ细胞与兔G8型神经节细胞同源；η细胞与兔G5型神经节细胞同源	较大的感受野，传递速度最慢；负责成像视觉或非成像视觉是不确定的

电生理学分类	形态学分类	亚类	同源性	功能特征
X型神经节细胞	β细胞：中型细胞体，小树突野	β-ON和β-OFF两类	与兔G4型神经节细胞同源	小感受野，对亮度和空间线性刺激产生持续反应，传递速度慢，可传递颜色、精细图案信号；成像视觉
Y型神经节细胞	α细胞：大型细胞体，大树突野	α-ON和α-OFF两类	与灵长类M神经节细胞同源；与兔G11型神经节细胞同源	大感受野，对空间非线性刺激产生瞬态反应(倍频响应)，传递速度快，可传递运动信息；非成像视觉
W型神经节细胞	γ细胞：中小型细胞体，大树突野	η细胞：小型细胞体，中等树突野；θ细胞：小型细胞体，中等树突野；θ-ON和θ-OFF两类	γ细胞与兔G8型神经节细胞同源；η细胞与兔G5型神经节细胞同源	较大的感受野，传递速度最慢；负责成像视觉或非成像视觉是不确定的

分型^*	形态特点	电生理特点与功能特性	同源性
G1型	兔视网膜中最小的神经节细胞，扁平胞体，致密、狭小的树突野，树突野直径大约为150~200 μm	局部边缘检测器
G2型	与G1树突分支类似，但更大而稀疏，树突野直径大约200 μm，且多位于单一平面内，分布不对称	—	与猫ζ细胞形态相似
G3型	小型双分层神经节细胞，其3~4个初级树突在视网膜内丛状层S4和S5分层，树突野直径大约为200~300 μm	负责有色视觉，对短波敏感，传递S-视锥细胞的蓝色信号	与灵长类动物中发现的小型双分层神经节细胞相同
G4型	树突野略大于G2和G3，其2~4个初级树突上布满细而短的分支，次级树突尖端肿胀为膨体	有ON和OFF两类	形态学上与猫β型细胞同源
G5型	树突野更大一些，直径大约为300 μm，外观辐射状，树突野中树突分布均匀	几乎均为OFF细胞	形态学与猫η型细胞同源
G6型	中型双分层树突野神经元，树突分布不均匀，相互交叉，树突野直径为300~500 μm，在内丛状层S1(OFF亚层)、S4和S5层(ON亚层)中多次分层	瞬态均匀检测器，其维持响应被所有类型的视觉刺激瞬时抑制，其ON树突的面积通常是OFF的两倍^[23]	分支模式类似猫ε型细胞
G7型	双分层的中型树突野神经元，在内丛状层的ON和OFF亚层中分层，树突环状分支，呈蜂窝样外观；与ON-OFF型DSGC类似的一种新型的双稳态瞬态ON-OFF型RGC近年来也被描述，其与ON-OFF型DSGC共分层，对闪烁的亮暗瞬时刺激作出响应，但与ON-OFF型DSGC不同的是，双稳态瞬态ON-OFF细胞对移动的刺激没有任何方向偏好^[24]	为兔视网膜中被广泛描述的ON-OFF型DSGC\^{[25,26,27,28]}	—
G8型	树突分支更大而稀疏，树突野直径为400 μm，多在S4层分层	—	形态学与猫γ型细胞同源
G9型	在内丛状层S1和S2的OFF亚层分层	—	形态与猫视网膜δ-OFF神经节细胞相似
G10型	与G11型类似，但初级树突比G11型更细，次级树突更长，在内丛状层不同水平分支；可进一步分为两种亚型，一种与无长突细胞耦联，一种不与无长突细胞耦联，表明其GABA依赖性定向信号可以独立于星爆型无长突细胞的机制产生^[29]	为先前研究的ON型DSGC^[25,30,31]	形态与猫δ-ON同源
G11型	兔视网膜中最大的神经节细胞	有ON和OFF两类，为瞬态反应细胞	与猫RGC中的α-Y细胞同源

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