【摘要】： 研究背景和目的: 神經(jīng)干細(xì)胞是近年來神經(jīng)科學(xué)領(lǐng)域的研究熱點(diǎn)。神經(jīng)干細(xì)胞的自我更新和多向分化潛能為許多神經(jīng)系統(tǒng)疾病如帕金森氏病等的治療帶來了新的希望。但關(guān)鍵問題是誘導(dǎo)神經(jīng)干細(xì)胞定向分化為所需的神經(jīng)元的機(jī)制尚不明確。如何實(shí)現(xiàn)神經(jīng)干細(xì)胞定向誘導(dǎo)分化是神經(jīng)發(fā)育研究的核心問題。目前研究表明決定神經(jīng)干細(xì)胞定向分化的機(jī)制有兩種:一種是細(xì)胞發(fā)育、分化的表觀遺傳學(xué)機(jī)制,另一種是以各類細(xì)胞因子為主的外來信號(hào)的調(diào)控,即微環(huán)境。其中關(guān)鍵基因的調(diào)控對(duì)神經(jīng)干細(xì)胞的增殖分化至關(guān)重要,它決定著神經(jīng)干細(xì)胞向特定神經(jīng)元分化的方向。目前主要通過以下2個(gè)途徑展開研究: (1)在體研究,通過基因轉(zhuǎn)導(dǎo)與敲除,檢測這些基因在促進(jìn)神經(jīng)干細(xì)胞分化方向上的作用; (2)體外研究,通過各種細(xì)胞因子誘導(dǎo)特定的基因,檢測在誘導(dǎo)刺激下基因?qū)ι窠?jīng)干細(xì)胞的增殖與分化的影響。但是目前有關(guān)基因調(diào)控機(jī)制仍未完全清楚。因此對(duì)神經(jīng)干細(xì)胞基因調(diào)控機(jī)制的研究顯得十分重要。 室管膜前下區(qū)(Anterior subventricular zone,SVZa)是公認(rèn)的神經(jīng)干細(xì)胞較為集中的地方之一。從胚胎期到成年階段,此區(qū)的神經(jīng)干細(xì)胞在體內(nèi)背-腹信號(hào)的控制下,不斷沿著嘴側(cè)遷移流(Rostral migratory stream,RSM)向嗅球(Olfactory bulb,OB)遷移,最后在OB分化為具有OB表型特征的中間神經(jīng)元,其中就包括多巴胺能神經(jīng)元(dopamine neurons,DA neurons )。SVZa神經(jīng)干細(xì)胞除具有一般神經(jīng)干細(xì)胞的自我更新和多向分化潛能外,還具有以下特點(diǎn):(1)自產(chǎn)生起即具備發(fā)育為神經(jīng)元的潛能;(2)遷移路徑固定,高度局限化,長距離遷移;(3)在遷移過程中始終維持增殖狀態(tài)和神經(jīng)元分化潛能而不進(jìn)一步分化。故SVZa神經(jīng)干細(xì)胞成為研究神經(jīng)干細(xì)胞增殖、遷移和分化調(diào)控機(jī)制的較佳模型,并成為神經(jīng)干細(xì)胞研究的重點(diǎn)。 SVZa神經(jīng)干細(xì)胞在向OB遷移、分化的過程中,需要體內(nèi)外復(fù)雜的環(huán)境信號(hào)來調(diào)控,其中包括DLX5及骨形成蛋白-2(Bone morphogenetic protein-2,BMP-2)等。研究表明,DLX(distall-less homeobox)基因家族中的DLX5基因與OB中間神經(jīng)元的發(fā)育密切相關(guān)。既往通過基因敲除鼠發(fā)現(xiàn)DLX5基因在SVZa神經(jīng)干細(xì)胞向嗅球多巴胺能中間神經(jīng)元分化過程中起著重要的作用,是出生后神經(jīng)發(fā)生的必須基因。雖然在體實(shí)驗(yàn)的研究證實(shí)DLX5基因是生后OB多巴胺能中間神經(jīng)元的必須基因,但DLX5具體調(diào)節(jié)神經(jīng)干細(xì)胞分化機(jī)制目前還不清楚。而且由于DLX5基因敲除鼠在圍生期就死亡,以后的研究就集中在體外實(shí)驗(yàn)研究。 BMP-2作為骨形成蛋白家族(Bone morphogenetic proteins ,BMPs)中重要成員,是干細(xì)胞所處局部微環(huán)境中非常重要的一種細(xì)胞因子。發(fā)現(xiàn)其對(duì)不同發(fā)育階段和不同部位的神經(jīng)干細(xì)胞的作用并不相同,既可以促進(jìn)其向神經(jīng)元分化也可以促進(jìn)其向星形膠質(zhì)細(xì)胞分化。而在胚胎鼠SVZa ,BMP-2可以促進(jìn)神經(jīng)干細(xì)胞向神經(jīng)元分化,并且明顯促進(jìn)其向多巴胺能神經(jīng)元分化。 研究已經(jīng)證實(shí)DLX5基因是OB多巴胺能中間神經(jīng)元分化的必須基因,那么什么基因可以調(diào)節(jié)DLX5對(duì)SVZa神經(jīng)干細(xì)胞向多巴胺能神經(jīng)元分化呢?結(jié)合既往研究:BMP-2及DLX5在SVZa神經(jīng)干細(xì)胞向OB多巴胺能神經(jīng)元分化中的重要作用; BMP-2明顯促進(jìn)骨組織DLX5的表達(dá)。那么在SVZa神經(jīng)干細(xì)胞向多巴胺能神經(jīng)元分化中,BMP-2是否作為DLX5的上游調(diào)節(jié)因子,通過調(diào)節(jié)DLX5的表達(dá)從而促進(jìn)SVZa神經(jīng)干細(xì)胞向多巴胺能神經(jīng)元分化呢?因此,本研究首先分離培養(yǎng)出SVZa神經(jīng)干細(xì)胞作為研究對(duì)象,觀察BMP-2及其拮抗劑Noggin對(duì)SVZa神經(jīng)干細(xì)胞向多巴胺能神經(jīng)元分化的影響,并分別用免疫熒光染色和RT-PCR檢測這個(gè)過程中DLX5表達(dá)水平的變化,證明BMP-2通過DLX5途徑誘導(dǎo)SVZa神經(jīng)干細(xì)胞向多巴胺能神經(jīng)元分化,希望為神經(jīng)干細(xì)胞的定向分化研究提供一些實(shí)驗(yàn)資料。 方法: 1、運(yùn)用細(xì)胞培養(yǎng)技術(shù)對(duì)SVZa神經(jīng)干細(xì)胞進(jìn)行分離培養(yǎng),并將收獲的細(xì)胞克隆球行免疫細(xì)胞學(xué)的Nestin鑒定,然后將細(xì)胞克隆球分化后行NSE、GFAP、Oligo-2的免疫細(xì)胞學(xué)鑒定,以明確其具有干細(xì)胞特性和多向分化潛能。 2、將同批次體外培養(yǎng)所得的SVZa神經(jīng)干細(xì)胞,分別以對(duì)照組、BMP-2組、BMP-2+Noggin組和Noggin組誘導(dǎo),分別用免疫熒光染色和RT PCR檢測不同處理組DLX5蛋白和DLX5 mRNA表達(dá)情況。 3、將同批次體外培養(yǎng)所得的SVZa神經(jīng)干細(xì)胞,分別以對(duì)照組、BMP-2組、BMP-2+Noggin組和Noggin組誘導(dǎo),通過免疫熒光染色和流式細(xì)胞技術(shù)檢測不同處理組SVZa神經(jīng)干細(xì)胞分化為多巴胺能神經(jīng)元的比例。 結(jié)果: 1、在體外無血清培養(yǎng)的條件下,可以培養(yǎng)出大量懸浮生長的細(xì)胞克隆球,Nestin免疫熒光染色陽性,分化后的NSE、GFAP、Oligo-2的免疫細(xì)胞學(xué)鑒定反應(yīng)也呈陽性反應(yīng)。 2、經(jīng)對(duì)照組、BMP-2組、BMP-2+Noggin組和Noggin組誘導(dǎo)72小時(shí)后,分別行免疫熒光染色和RT PCR檢測,結(jié)果顯示BMP-2組中鏡下可見DLX5表達(dá)明顯增強(qiáng),而對(duì)照組、BMP-2+Noggin組和Noggin組的DLX5表達(dá)無明顯變化;BMP-2組中DLX5 mRNA表達(dá)量也明顯高于對(duì)照組,p0.05,差異有統(tǒng)計(jì)學(xué)意義,而對(duì)照組、BMP-2+Noggin組和Noggin組DLX5mRNA表達(dá)量組間兩兩比較,差異無顯著性,p0.05。 3、經(jīng)對(duì)照組、BMP-2組、BMP-2+Noggin組和Noggin組誘導(dǎo)分化72小時(shí)后,分別行免疫熒光染色及流式細(xì)胞儀檢測,結(jié)果顯示BMP-2組可見較多TH+細(xì)胞,而對(duì)照組、BMP-2 +Noggin組和Noggin組的TH +細(xì)胞明顯較少;流式細(xì)胞結(jié)果也提示BMP-2組TH+神經(jīng)元分化比例較對(duì)照組明顯增高,p0.05,差異有統(tǒng)計(jì)學(xué)意義;而對(duì)照組、BMP-2+Noggin組和Noggin組TH +神經(jīng)元分化比例組間兩兩比較,差異無顯著性,p0.05。 結(jié)論: 1、從E15天Wistar大鼠SVZa腦組織中成功分離得到具有不斷增殖和自我更新能力,并可以分化為神經(jīng)元、星形膠質(zhì)細(xì)胞和少突膠質(zhì)細(xì)胞的神經(jīng)干細(xì)胞。 2、BMP-2明顯促進(jìn)SVZa神經(jīng)干細(xì)胞DLX5蛋白和DLX5mRNA表達(dá),而且這種作用能被其拮抗劑Noggin所拮抗。 3、BMP-2明顯促進(jìn)SVZa神經(jīng)干細(xì)胞分化為多巴胺能神經(jīng)元,并且這種作用同樣能被其拮抗劑Noggin所拮抗。在SVZa,BMP-2可能通過上調(diào)DLX5表達(dá)從而促進(jìn)神經(jīng)干細(xì)胞向TH+神經(jīng)元分化。
[Abstract]:Background and purpose of the study: Neural stem cells are the field of neuroscience in recent years The self-renewal and multiple-to-differentiation potential of neural stem cells bring new treatment to many nervous system diseases, such as Parkinson's disease Hopefully, the key issue is that the mechanism for inducing the directional differentiation of neural stem cells into the desired neurons is not yet It is clear how to realize the directional induction and differentiation of neural stem cells is the core of the neurodevelopmental study. The present study indicates that there are two mechanisms to determine the directional differentiation of neural stem cells: one is the epigenetic mechanism of cell development and differentiation, and the other is the regulation of external signals dominated by various cytokines. The regulation of the key genes is crucial to the proliferation and differentiation of neural stem cells, which determines the differentiation of neural stem cells to specific neurons. direction. The study was conducted mainly by (1) in vivo studies, by gene transduction and knock-out, to detect the role of these genes in promoting the differentiation of neural stem cells, (2) in vitro studies, by the induction of specific cytokines by various cytokines, Gene, to detect the proliferation and differentiation of the gene to neural stem cells under the induction of stimulation However, the current regulation mechanism of the gene is still incomplete. It is clear that the research on the regulation mechanism of neural stem cell gene is very important It is important that the anterior inferior zone (SVZa) of the ependymal membrane is a well-recognized neural stem cell. One of the places from the embryonic stage to the adult stage, the neural stem cells of this region, under the control of the in vivo back-abdominal signal, migrate continuously along the mouth-side migration flow (RSM) to the olfactory bulb (OB), and finally, in the OB differentiation to have the OB phenotype characteristic the intermediate neurons, including the dopaminergic neurons (dopamine netrons, da neu, The SVZa neural stem cells have the following characteristics in addition to the self-renewal and multi-directional differentiation potential of general neural stem cells: (1) the potential for development as a neuron is self-generated; (2) the migration path is fixed and highly localized, Long-distance migration; (3) always maintain the proliferative state and the potential of the neuron differentiation during the migration, and not The SVZa neural stem cell is a better model for the study of the mechanism of the proliferation, migration and differentiation of neural stem cells and becomes a neural stem cell. The focus of the study was that the SVZa neural stem cells, in the process of migration and differentiation to OB, needed to be regulated by a complex in-vivo environment signal, including DLX5 and bone morphogenetic protein-2 (Bone morphogenetic protein-2). and BMP-2, etc. The study shows that the DLX5 gene in the DLX (distall-less homoerox) gene family and the OB middle nerve It is closely related to the development of meta. In the past, the gene knockout mice have found that the DLX5 gene plays an important role in the differentiation of the dopaminergic neurons of the olfactory bulb in the SVZa neural stem cell, and is the postnatal god. Although the study of in vivo experiments confirmed that the DLX5 gene is a necessary gene for OB dopaminergic intermediate neurons, the DLX5 specifically regulates the differentiation of neural stem cells. The mechanism is not yet clear. And since the DLX5 knockout mice died during the perinatal period, the later study set In vitro, BMP-2 is an important member of bone morphogenetic protein (BMPs) and is the local microenvironment of stem cells. It is a very important cytokine. It is found that its effects on the neural stem cells at different developmental stages and in different parts are not the same, which can promote the differentiation of the neurons to the neurons. In embryonic rat SVZa, BMP-2 can promote the differentiation of neural stem cells into neurons and promote the differentiation of neural stem cells. The study has confirmed that the DLX5 gene is an essential gene for OB dopaminergic neuron differentiation, so what gene can regulate the DLX5 to the SVZa nerve The differentiation of stem cells to dopaminergic neurons? Combined with previous studies: BMP-2 and DLX5 play an important role in the differentiation of the neural stem cells of SVZa to OB dopaminergic neurons; BMP-2 The expression of the DLX5 in the bone tissue is obviously promoted. In the differentiation of the SVZa neural stem cell into the dopaminergic neuron, whether the BMP-2 is the upstream regulatory factor of the DLX5, and the expression of the DLX5 is adjusted to promote the SVZa nerve. The effect of BMP-2 and its antagonist Noggin on the differentiation of the dopaminergic neurons of the SVZa neural stem cells was observed and the effect of the antagonist Noggin on the differentiation of the dopaminergic neurons was observed, and this was detected by immunofluorescent staining and RT-PCR, respectively. The changes of the expression level of the DLX5 in the process demonstrated that the BMP-2 induces the differentiation of the SVZa neural stem cells to the dopaminergic neurons through the DLX5 pathway and is expected to be a neural stem cell. directional point Methods:1. The SVZa neural stem cells were isolated and cultured by cell culture, and the harvested cells were identified by Nestin, and then the cells were cloned into NSE. The Immune Cytological Evaluation of GFAP and Oligo-2 (2) The SVZa neural stem cells obtained from in-vitro culture of the same batch were cultured in the control group, the BMP-2 group, the BMP-2 + Noggin group and the Noggin group, respectively. The expression of DLX5 and DLX5 mRNA in the same batch was compared with that of the control group, the BMP-2 group, the BMP-2 + Noggin group and the Noggin group. co-processing The ratio of the group of SVZa neural stem cells to the dopaminergic neurons was determined. The results were as follows:1. Under the condition of no serum culture in vitro, a large number of cell clone balls of suspended growth could be cultured, and the Nestin immunofluorescence staining was positive and differentiated. The results showed that the expression of DLX5 in BMP-2 group was significantly enhanced by RT-PCR, and the results showed that the expression of DLX5 was significantly enhanced in BMP-2 group. There was no significant change in the expression of DLX5 in the group, BMP-2 + Noggin group and Noggin group, and the amount of DLX5 mRNA in the BMP-2 group was significantly higher than that of the control group, p0.05, the difference was of statistical significance, and the control group, BMP-2 + Noggin group and No. The expression of DLX5mRNA in ggin group was not significant between the two groups. The difference was not significant, p0.05.3. After 72 hours of induction and differentiation of the control group, the BMP-2 group, the BMP-2 + Noggin group and the Noggin group, the results showed that the BMP-2 group showed more TH + fine. In the control group, the TH + cells of the control group, the BMP-2 + Noggin group and the Noggin group were significantly less. The flow cytometry also suggested that the proportion of TH + neurons in the BMP-2 group was significantly higher than that in the control group. Group and N There was no significant difference between the ratio of TH and neuron in the goggin group, and the difference was not significant, p0.05. Conclusion:1. The successful separation of the brain of the SVZa from the Wistar rats from E15 days. neural stem cells with constant proliferation and self-renewal and can be differentiated into neurons, astrocytes, and oligodendrocytes. BMP-2 significantly promotes S VZa neural stem cell DLX5 protein and DLX5mRNA expression, and this effect can be antagonized by its antagonist Noggin.3, BM P-2 significantly promotes the differentiation of SVZa neural stem cells into dopaminergic neurons, and this effect can also be used as an antagonist, N,
【學(xué)位授予單位】：第三軍醫(yī)大學(xué)
【學(xué)位級(jí)別】：碩士
【學(xué)位授予年份】：2010
【分類號(hào)】：R329

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