Yadav Shubhash C, Tetenborg Stephan, Dedek Karin
Animal Navigation/Neurosensorics, Institute for Biology and Environmental Sciences, University of Oldenburg, Oldenburg, Germany.
Research Center Neurosensory Science, University of Oldenburg, Oldenburg, Germany.
Front Mol Neurosci. 2019 Apr 23;12:99. doi: 10.3389/fnmol.2019.00099. eCollection 2019.
In the mammalian retina, amacrine cells represent the most diverse cell class and are involved in the spatio-temporal processing of visual signals in the inner plexiform layer. They are connected to bipolar, other amacrine and ganglion cells, forming complex networks via electrical and chemical synapses. The small-field A8 amacrine cell was shown to receive non-selective glutamatergic input from OFF and ON cone bipolar cells at its bistratified dendrites in sublamina 1 and 4 of the inner plexiform layer. Interestingly, it was also shown to form electrical synapses with ON cone bipolar cells, thus resembling the rod pathway-specific AII amacrine cell. In contrast to the AII cell, however, the electrical synapses of A8 cells are poorly understood. Therefore, we made use of the Ier5-GFP mouse line, in which A8 cells are labeled by GFP, to study the gap junction composition and frequency in A8 cells. We found that A8 cells form <20 gap junctions per cell and these gap junctions consist of connexin36. Connexin36 is present at both OFF and ON dendrites of A8 cells, preferentially connecting A8 cells to type 1 OFF and type 6 and 7 ON bipolar cells and presumably other amacrine cells. Additionally, we show that the OFF dendrites of A8 cells co-stratify with the processes of dopaminergic amacrine cells from which they may receive GABAergic input via GABA receptor subunit α3. As we found A8 cells to express dopamine receptor D (but not D), we also tested whether A8 cell coupling is modulated by D receptor agonists and antagonists as was shown for the coupling of AII cells. However, this was not the case. In summary, our data suggests that A8 coupling is differently regulated than AII cells and may even be independent of ambient light levels and serve signal facilitation rather than providing a separate neuronal pathway.
在哺乳动物视网膜中,无长突细胞是种类最多样化的细胞类型,参与内网状层视觉信号的时空处理。它们与双极细胞、其他无长突细胞和神经节细胞相连,通过电突触和化学突触形成复杂的网络。研究表明,小视野A8无长突细胞在其位于内网状层1层和4层的双分层树突处接收来自OFF和ON视锥双极细胞的非选择性谷氨酸能输入。有趣的是,还发现它与ON视锥双极细胞形成电突触,因此类似于视杆通路特异性的AII无长突细胞。然而,与AII细胞不同的是,A8细胞的电突触目前了解甚少。因此,我们利用Ier5-GFP小鼠品系(其中A8细胞由绿色荧光蛋白标记)来研究A8细胞中的缝隙连接组成和频率。我们发现,每个A8细胞形成的缝隙连接少于20个,这些缝隙连接由连接蛋白36组成。连接蛋白36存在于A8细胞的OFF和ON树突上,优先将A8细胞与1型OFF双极细胞以及6型和7型ON双极细胞以及可能的其他无长突细胞相连。此外,我们发现A8细胞的OFF树突与多巴胺能无长突细胞的突起共同分层,A8细胞可能通过GABA受体亚基α3从多巴胺能无长突细胞接收GABA能输入。由于我们发现A8细胞表达多巴胺受体D(但不表达D),我们还测试了A8细胞的耦合是否像AII细胞的耦合那样受到D受体激动剂和拮抗剂的调节。然而,情况并非如此。总之,我们的数据表明,A8细胞的耦合调节方式与AII细胞不同,甚至可能与环境光照水平无关,其作用是促进信号传递而非提供一条独立的神经元通路。