Marmor-Kollet Neta, Schuldiner Oren
Department of Molecular Cell Biology, Weizmann Institute of Sciences, Rehovot, 76100, Israel.
Dev Neurobiol. 2016 Mar;76(3):262-76. doi: 10.1002/dneu.22312. Epub 2015 Jun 10.
The molecular mechanisms regulating intrinsic axon growth potential during development or following injury remain largely unknown despite their vast importance. Here, we have established a neurite sprouting assay of primary cultured mushroom body (MB) neurons. We used the MARCM technique to both mark and manipulate MB neurons, enabling us to quantify the sprouting abilities of single WT and mutant neurons originating from flies at different developmental stages. Sprouting of dissociated MB neurons was dependent on wnd, the DLK ortholog, a conserved gene that is required for axon regeneration. Next, and as expected, we found that the sprouting ability of adult MB neurons was significantly decreased. In contrast, and to our surprise, we found that pupal-derived neurons exhibit increased sprouting compared with neurons derived from larvae, suggesting the existence of an elevated growth potential state. We then contrasted the molecular requirements of neurite sprouting to developmental axon regrowth of MB ɣ neurons, a process that we have previously shown requires the nuclear receptor UNF acting via the target of rapamycin (TOR) pathway. Strikingly, we found that while TOR was required for neurite sprouting, UNF was not. In contrast, we found that PTEN inhibits sprouting in adult neurons, suggesting that TOR is regulated by the PI3K/PTEN pathway during sprouting and by UNF during developmental regrowth. Interestingly, the PI3K pathway as well as Wnd were not required for developmental regrowth nor for initial axon outgrowth suggesting that axon growth during circuit formation, remodeling, and regeneration share some molecular components but differ in others.
尽管其极为重要,但在发育过程中或损伤后调节轴突内在生长潜力的分子机制仍 largely 未知。在这里,我们建立了原代培养的蘑菇体(MB)神经元的神经突萌发试验。我们使用 MARCM 技术来标记和操纵 MB 神经元,使我们能够量化源自不同发育阶段果蝇的单个野生型和突变型神经元的萌发能力。解离的 MB 神经元的萌发依赖于 wnd,即 DLK 的直系同源物,这是一种轴突再生所需的保守基因。接下来,正如预期的那样,我们发现成年 MB 神经元的萌发能力显著降低。相比之下,令我们惊讶的是,我们发现蛹源神经元与幼虫源神经元相比表现出增加的萌发,这表明存在一种升高的生长潜力状态。然后,我们将神经突萌发的分子需求与 MBɣ 神经元的发育性轴突再生进行了对比,我们之前已经表明这个过程需要核受体 UNF 通过雷帕霉素靶标(TOR)途径发挥作用。令人惊讶的是,我们发现虽然 TOR 是神经突萌发所必需的,但 UNF 不是。相反,我们发现 PTEN 抑制成年神经元的萌发,这表明 TOR 在萌发过程中受 PI3K/PTEN 途径调节,而在发育性再生过程中受 UNF 调节。有趣的是,PI3K 途径以及 Wnd 在发育性再生和初始轴突生长过程中都不是必需的,这表明在神经回路形成、重塑和再生过程中的轴突生长共享一些分子成分,但在其他方面有所不同。