Prithviraj Ranjini, Trunova Svetlana, Giniger Edward
National Institute of Neurological Disorders and Stroke, National Institutes of Health, Bethesda, MD 20892, USA.
J Vis Exp. 2012 Jul 27(65):4270. doi: 10.3791/4270.
We describe a method for ex vivo culturing of whole Drosophila brains. This can be used as a counterpoint to chronic genetic manipulations for investigating the cell biology and development of central brain structures by allowing acute pharmacological interventions and live imaging of cellular processes. As an example of the technique, prior work from our lab(1) has shown that a previously unrecognized subcellular compartment lies between the axonal and somatodendritic compartments of axons of the Drosophila central brain. The development of this compartment, referred to as the axon initial segment (AIS)(2), was shown genetically to depend on the neuron-specific cyclin-dependent kinase, Cdk5. We show here that ex vivo treatment of wild-type Drosophila larval brains with the Cdk5-specific pharmacological inhibitors roscovitine and olomoucine(3) causes acute changes in actin organization, and in localization of the cell-surface protein Fasciclin 2, that mimic the changes seen in mutants that lack Cdk5 activity genetically. A second example of the ex vivo culture technique is provided for remodeling of the connections of embryonic mushroom body (MB) gamma neurons during metamorphosis from larva to adult. The mushroom body is the center of olfactory learning and memory in the fly(4), and these gamma neurons prune their axonal and dendritic branches during pupal development and then re-extend branches at a later timepoint to establish the adult innervation pattern(5). Pruning of these neurons of the MB has been shown to occur via local degeneration of neurite branches(6), by a mechanism that is triggered by ecdysone, a steroid hormone, acting at the ecdysone receptor B1(7), and that is dependent on the activity of the ubiquitin-proteasome system(6). Our method of ex vivo culturing can be used to interrogate further the mechanism of developmental remodeling. We found that in the ex vivo culture setting, gamma neurons of the MB recapitulated the process of developmental pruning with a time course similar to that in vivo. It was essential, however, to wait until 1.5 hours after puparium formation before explanting the tissue in order for the cells to commit irreversibly to metamorphosis; dissection of animals at the onset of pupariation led to little or no metamorphosis in culture. Thus, with appropriate modification, the ex vivo culture approach can be applied to study dynamic as well as steady state aspects of central brain biology.
我们描述了一种对完整果蝇大脑进行离体培养的方法。通过允许进行急性药理学干预和对细胞过程进行实时成像,该方法可作为慢性基因操作的补充手段,用于研究中枢脑结构的细胞生物学和发育。作为该技术的一个例子,我们实验室之前的工作(1)表明,果蝇中枢脑轴突的轴突和体树突区室之间存在一个先前未被识别的亚细胞区室。这个被称为轴突起始段(AIS)(2)的区室的发育在遗传学上被证明依赖于神经元特异性细胞周期蛋白依赖性激酶Cdk5。我们在此表明,用Cdk5特异性药理学抑制剂roscovitine和olomoucine(3)对野生型果蝇幼虫大脑进行离体处理,会导致肌动蛋白组织以及细胞表面蛋白Fasciclin 2的定位发生急性变化,这些变化与在基因上缺乏Cdk5活性的突变体中观察到的变化相似。离体培养技术的另一个例子是关于从幼虫到成虫变态过程中胚胎蘑菇体(MB)γ神经元连接的重塑。蘑菇体是果蝇嗅觉学习和记忆的中心(4),这些γ神经元在蛹发育过程中修剪其轴突和树突分支,然后在稍后的时间点重新延伸分支以建立成虫的神经支配模式(5)。MB中这些神经元的修剪已被证明是通过神经突分支的局部退化发生的(6),其机制由蜕皮激素触发,蜕皮激素是一种类固醇激素,作用于蜕皮激素受体B1(7),并且依赖于泛素 - 蛋白酶体系统的活性(6)。我们的离体培养方法可用于进一步探究发育重塑的机制。我们发现,在离体培养环境中,MB的γ神经元以与体内相似的时间进程重现了发育修剪过程。然而,至关重要的是要等到化蛹后1.5小时再取出组织,以便细胞不可逆转地进入变态;在化蛹开始时解剖动物,在培养中几乎不会发生或根本不会发生变态。因此,经过适当修改,离体培养方法可应用于研究中枢脑生物学的动态以及稳态方面。