本文選題：NLRC5　切入點(diǎn)：白痢沙門氏菌　出處：《揚(yáng)州大學(xué)》2015年博士論文　論文類型：學(xué)位論文

【摘要】：NOD樣受體家族CARD結(jié)構(gòu)域包含分子5 (NLR family, CARD domain containing 5, NLRC5)在調(diào)控天然免疫和細(xì)胞免疫中發(fā)揮重要作用,到目前為止,NLRC5已經(jīng)被證實(shí)在人和小鼠中可以調(diào)控MHCI類分子的表達(dá),并對NF-κB以及I型干擾素分子通路具有抑制性調(diào)控作用。白痢沙門氏菌病是我國家禽養(yǎng)殖重要的細(xì)菌性疾病之一,該病的爆發(fā)給家禽生產(chǎn)造成嚴(yán)重的經(jīng)濟(jì)損失。白痢沙門氏菌的隱性感染和攜帶狀態(tài)的形成會導(dǎo)致重復(fù)發(fā)病進(jìn)而防治困難。白痢沙門氏菌侵入雞體內(nèi)后,會抑制MHC Class I以及誘導(dǎo)IL10表達(dá),這一機(jī)制是阻斷清除的過程從而導(dǎo)致攜帶狀態(tài)和持續(xù)感染狀態(tài)的原因之一。雞NLRC5分子可能與沙門氏菌清除的分子機(jī)理有關(guān)。截止目前雛雞感染白痢沙門氏菌后組織水平的高通量研究尚無報(bào)道,其他菌株由于致病性和病理機(jī)制的差異僅能提供部分參考,因此有必要專門對雞白痢沙門氏菌感染過程中的表達(dá)譜變化進(jìn)行系統(tǒng)研究,并在此基礎(chǔ)上進(jìn)一步研究家禽NLRC5及相關(guān)基因的表達(dá)規(guī)律以及啟動子缺失對NLRC5表達(dá)的影響,從而分析其受調(diào)控的可能機(jī)制,以幫助建立家禽的NLRC5調(diào)控的機(jī)制模型,更好的理解家禽白痢沙門氏菌隱性感染形成的機(jī)制以及細(xì)菌性病原清除的機(jī)理。研究內(nèi)容和結(jié)果如下。1.第二章第一節(jié),比較不同沙門氏菌菌種感染導(dǎo)致雛雞的脾臟和盲腸發(fā)生的表達(dá)譜變化。通過安捷倫雞全基因組表達(dá)譜芯片,分別對7dpi感染了白痢、腸炎還有poly(I:C)的雛雞脾臟和盲腸進(jìn)行了表達(dá)譜分析,同時檢測IL-la, IFNy, IgG和IgM濃度,腸道載菌量和免疫熒光鑒定脾臟載菌量。結(jié)果表明,病原刺激激發(fā)了系統(tǒng)性沙門氏菌感染的主要特征性免疫反應(yīng)。在白痢沙門氏菌感染組中,脾臟表達(dá)譜富集分析結(jié)果顯示,差異表達(dá)基因主要富集于淋巴細(xì)胞的分化和增殖過程,因而呈現(xiàn)出獲得性免疫的有效激活狀態(tài)；其中與Thl淋巴以及NK細(xì)胞激活有關(guān)的重要分子諸如IL15, CD28等均呈現(xiàn)下調(diào),而與B淋巴細(xì)胞活化有關(guān)的基因例如IL7呈現(xiàn)上調(diào)。這表明細(xì)胞免疫的清除作用一定程度上已經(jīng)受到了抑制,而B細(xì)胞所參與的抗體反應(yīng)被激活。而感染白痢沙門氏菌組的盲腸組織中,下調(diào)基因主要與天然性免疫系統(tǒng)有關(guān),表明相關(guān)天然性免疫反應(yīng)或細(xì)胞免疫反應(yīng)活性的減弱,推測可能與抗體的保護(hù)作用增強(qiáng)有關(guān)。第二章第二節(jié),在證實(shí)白痢沙門氏菌感染過程在7dpi造成細(xì)胞免疫功能衰弱的基礎(chǔ)上,然后利用時間序列表達(dá)譜分析白痢沙門氏菌感染雛雞后2小時、4小時、8小時、24小時、3天、5天、7天、12天、21天脾臟組織中重要的分子事件。同時測定體重、脾臟指數(shù)、法氏囊指數(shù)、胸腺指數(shù)、以及IL-1α、IFNγ、IgG和IgM濃度。通過WGCNA的表型關(guān)聯(lián)分析,以及探針間的共表達(dá)特征,韋恩分析等方法挖掘?qū)е码[性感染和攜帶的差異表達(dá)基因。時序表達(dá)譜的分析初步探明了在白痢沙門氏菌感染的關(guān)鍵時期脾臟組織中表達(dá)譜發(fā)生的變化,觀察到了天然免疫到體液免疫發(fā)展變化過程中脾臟組織中重要的分子事件,包括淋巴細(xì)胞的激活,增殖和遷移事件；并鑒定出大量關(guān)鍵調(diào)控基因,例如MHC class I B-F, ANXA13, IL18, BF2, TNFRSF13B, BLB2以及CD28等；富集通路中的基因呈現(xiàn)兩類高度負(fù)相關(guān)的功能團(tuán)；差異表達(dá)基因中的節(jié)點(diǎn)基因的表達(dá)模式與脾臟指數(shù)高度正相關(guān)；在不同階段中免疫反應(yīng)有關(guān)的通路表現(xiàn)出與其他信號通路和代謝通路不同的關(guān)聯(lián)特征；差異表達(dá)基因大多數(shù)均可以定位在有報(bào)道的與家禽抗體滴度有關(guān)的QTL區(qū)域內(nèi)；NLRC5是差異表達(dá)基因；3dpi-5dpi是天然性免疫抑制和細(xì)胞免疫激活的關(guān)鍵階段,也是病原清除的主要時期,該階段的差異表達(dá)基因表達(dá)趨勢反應(yīng)了雛雞對病原做出免疫反應(yīng)的特征；NLRC5的表達(dá)模式表明NLRC5的翻譯和激發(fā)的下游通路促進(jìn)了細(xì)胞免疫反應(yīng)和病原的清除作用,但同時可能也受到了負(fù)調(diào)控因素的調(diào)控。第二章第三節(jié),進(jìn)而基于上述試驗(yàn)提供的表達(dá)譜芯片數(shù)據(jù),以及NLRC5對MHCI已知的調(diào)控作用的假設(shè),開展針對NLRC5在表達(dá)譜數(shù)據(jù)中的共表達(dá)分析,嘗試找到重要的共表達(dá)分子。共表達(dá)分析結(jié)果顯示,NLRC5與TAP1和IL21R高度共表達(dá),其中作為MHCI區(qū)域中的分子TAP1可能在受到NF-κB調(diào)控的同時也受到NLRC5的影響。2.第三章第一節(jié),鑒于NLRC5基因功能的重要性及其在雞白痢沙門氏菌感染過程中的表達(dá)與調(diào)控功能,本研究進(jìn)一步對雞NLRC5基因功能進(jìn)行研究。首先,克隆了雞NLRC5的全長CDS以及UTR序列,并對其進(jìn)行細(xì)致的生物信息學(xué)分析。在組織水平對有報(bào)道可能與NLRC5功能相關(guān)的基因進(jìn)行熒光定量的檢測和分析。第三章第二節(jié),隨后基于克降的片段構(gòu)建過表達(dá)和干擾載體,轉(zhuǎn)染DF1細(xì)胞系,并構(gòu)建穩(wěn)定表達(dá)株。在此基礎(chǔ)上首先進(jìn)行細(xì)胞免疫熒光鑒定,然后對細(xì)胞進(jìn)行了LPS刺激,以觀察相關(guān)基因表達(dá)變化的趨勢。最后在穩(wěn)定表達(dá)細(xì)胞株上對NLRC5以及信號通路中的關(guān)鍵基因的表達(dá)模式進(jìn)行了比較和分析,并提出了NLRC5及重要相關(guān)基因的假設(shè)調(diào)控機(jī)制。NLRC5克隆和分析結(jié)果顯示,雞NLRC5與哺乳動物NLRC5基因具有一定的同源性,推測可能和哺乳動物該基因功能一致,可以進(jìn)入核內(nèi)發(fā)揮作用,并受到表觀遺傳修飾的影響。NLRC5及其信號通路中的關(guān)鍵基因在組織和細(xì)胞中的表達(dá)研究發(fā)現(xiàn),NF-κB2與NLRC5有顯著的線性共表達(dá)關(guān)系,雞的STAT1與NLRC5沒有明顯的共表達(dá)特征,推測可能對NLRC5的調(diào)控作用較復(fù)雜或可能沒有調(diào)控作用,而STAT3可能作為NF-κB2的負(fù)調(diào)控因子負(fù)反饋調(diào)控了NLRC5的表達(dá)。綜上,除MHCI外,NLRC5可能調(diào)控了STAT3和IL18的轉(zhuǎn)錄表達(dá)。家禽NLRC5可能具有與哺乳動物不同的表達(dá)調(diào)控機(jī)制。3.第四章,為進(jìn)一步弄清NLRC5受到調(diào)控的具體過程,以及進(jìn)一步證明雞NLRC5對MHCI的調(diào)控作用,進(jìn)而幫助更好的開展SPI-2 Ⅲ型分泌系統(tǒng)對隱性感染和攜菌狀態(tài)形成作用的研究奠定基礎(chǔ),本研究初步探索了NLRC5起始密碼子前4372bp區(qū)域不同片段的活性,并分析其潛在的可能調(diào)控原理。NLRC5的系列缺失結(jié)果證明,NLRC5可能具有2個有活性的啟動子區(qū)域,并且通過617到1448之間的區(qū)域隔離兩個啟動子的活性；第二個啟動子活性要強(qiáng)于第一個啟動子活性,并且在該區(qū)域內(nèi)發(fā)現(xiàn)了NF-κB等有報(bào)道與NLRC5表達(dá)調(diào)控有密切相關(guān)的轉(zhuǎn)錄因子的結(jié)合基序,1448到4372的片段可能是脾臟組織中的發(fā)揮作用的主要啟動子,轉(zhuǎn)錄表達(dá)主要從2108開始的NLRC5分子片段；下游調(diào)控可能較為復(fù)雜并且下游元件對NLRC5的表達(dá)有顯著的影響；1-2108的活性僅為1448-4372的50%左右,這表明1448-2108的活性要比1828到2108的活性低,因而推測,1828到2213的活性會進(jìn)一步增強(qiáng),甚至超過1448到4372的活性,因此認(rèn)為NLRC5的核心啟動子主要為1828到2213的片段,該區(qū)間包含的元件可能有STAT1、NF-κB、P300、MRF2、FAC1、 PAX5、CRFB、CPBP、NF-AT1、MRF-2、AML1、FAC1、CRX、RelA-p65。其中STAT1和NF-κB均于轉(zhuǎn)錄起始位點(diǎn)前182bp起始的位置被鑒定出來。未來,研究需確認(rèn)家禽NLRC5對MHCI調(diào)控的機(jī)制,通過更進(jìn)一步的ChIP,乃至ChIP-Seq對NLRC5結(jié)合啟動子區(qū)域進(jìn)行全基因組的分析,結(jié)合RNA-Seq分析可以更準(zhǔn)確的弄清該基因是否對STAT3, MHCI以及IL18等基因存在調(diào)控作用,再通過EMSA,點(diǎn)突變以及敲除和過表達(dá)實(shí)驗(yàn)來進(jìn)一步的確認(rèn)NLRC5可能存在的調(diào)控機(jī)制。最終轉(zhuǎn)入白痢沙門氏菌SPI-2 Ⅲ型分泌系統(tǒng)對巨噬細(xì)胞中MHCI有關(guān)通路的調(diào)控作用的研究,以期揭示白痢沙門氏菌隱性感染和攜帶狀態(tài)形成的具體機(jī)制。
[Abstract]:NOD like receptor family, CARD domain containing 5 (NLR family, CARD domain containing 5, NLRC5) play an important role in the innate immune cells and immune regulation, so far, NLRC5 has been demonstrated in human and mouse can regulate the expression of MHCI molecules, and has inhibitory effects on the expression of NF- B and type I interferon pathways. Salmonella pullorum disease is one of the most important bacterial diseases in China poultry breeding, the disease outbreak caused serious economic losses to poultry production. Recessive Salmonella infection and carrier status form will lead to repeated and difficult to control. The incidence of Salmonella invasion of chicken in vivo, Class could inhibit MHC induced expression of IL10 and I, this mechanism is blocking the process of elimination resulting in one of the reasons for carrying state and persistent infection. Chicken NLRC5 molecules and Salmonella The molecular mechanism of removal. As of high-throughput studies currently infected with Salmonella after tissue level has not been reported, due to differences in other strains of pathogenic and pathological mechanism can only provide some reference, so it is necessary to systematically study the spectrum changes in the expression of Salmonella pullorum infection, affecting the expression of law as well as promoter deletion and on the basis of further study of NLRC5 in poultry and related gene expression of NLRC5, so as to analyze the possible mechanism by regulation, mechanism model of NLRC5 regulation of poultry to help establish a better understanding of the poultry Salmonella infection mechanism and the mechanism of the formation of recessive bacterial pathogen clearance. Research content.1. and the results are as follows: the first section of the second chapter, comparison of different strains of Salmonella infection leads to changes in the expression of chicken spleen and cecum by spectrum. Agilent chicken whole genome microarray, respectively for 7dpi and poly infection diarrhea, enteritis (I:C) in spleen and cecum expression spectrum analysis, simultaneous detection of IL-la, IFNy, IgG and IgM concentration, intestinal bacteria loading and immunofluorescence identification of spleen microbial load. The results showed that the pathogen triggers system Salmonella infection. The main characteristics of the immune response in Salmonella pullorum infection group, spleen expression enrichment analysis showed that the expression of differentiation and proliferation of genes mainly enriched in lymphocytes, thus showing a valid activation state of acquired immunity; one with the Thl and NK lymph cell activation related important molecules such as IL15, CD28 and B were down regulated, lymphocyte activation related genes such as IL7 increased. This indicates that the removal of cell immune function to a certain extent has been suppressed System, antibody response and B cells in the cecal tissue is activated. Infection of Salmonella group, down regulated genes mainly related to innate immunity, that natural immunity or weaken the cellular immune response activity, presumably related with protective antibodies enhanced. The second chapter second section. In that process in 7dpi cause the underlying cellular immune function weakened on Salmonella infection, and then use the time series expression analysis of 2 hours of chickens infected with Salmonella after 4 hours, 8 hours, 24 hours, 3 days, 5 days, 7 days, 12 days, 21 days were important molecular events tissue. Simultaneous determination of body weight, spleen index, bursal index, thymus index, and IL-1 alpha, IFN gamma, IgG and IgM concentration. Through the analysis of WGCNA and phenotype correlation, co expression characteristics between the probes, Wayne analysis method of mining cause latent infection The difference of gene expression and carrying. Temporal expression spectrum analysis preliminarily expression changes in spleen tissue critical period of Salmonella infection, observed natural immunity to humoral immunity is an important molecular event in the process of development and change in spleen tissue, including lymphocyte activation, proliferation and migration events; and to identify a large number of key regulatory genes, such as MHC class, I B-F, ANXA13, IL18, BF2, TNFRSF13B, BLB2 and CD28; enrichment pathway gene shows two kinds of highly negative related functional groups; differential expression and expression pattern of genes in the spleen index node highly positive correlation; in different stages of the immune response the pathway showed different characteristics with other signaling pathways and metabolic pathways; differentially expressed genes were reported in most can locate the anti body and poultry The titer of the QTL region; NLRC5 is differentially expressed genes; 3dpi-5dpi is the key stage of natural immune suppression and cell immune activation, and also is the important time to remove pathogens, gene expression in response to a trend feature chicken's immune response to pathogenic differential expression of the stage; the expression pattern of NLRC5 showed that NLRC5 translation and excitation the downstream pathway promoted scavenging reactions and pathogenic immune cells, but may also be the regulation of negative regulatory factors. The second chapter third section, and then the expression test based on the microarray data, and assuming the role of NLRC5 in regulation of the known MHCI, to carry out targeted expression of NLRC5 in the co expression data in spectrum analysis and try to find the important co expression of molecular. Analysis results showed that the co expression of NLRC5 and TAP1 and IL21R highly co expressed, as TAP1 molecules in MHCI region can be Can be NF- K B regulation but also by the effect of.2. third NLRC5 in the first chapter, in view of the importance of NLRC5 gene and its function in the process of expression and regulation function of Salmonella pullorum infection, this study further study of chicken NLRC5 gene. Firstly, cloning the full-length CDS and chicken NLRC5 the sequence of UTR, and carries on the detailed analysis of the biological information. At the organizational level of coverage may be correlated with NLRC5 fluorescence quantitative gene detection and analysis. The third chapter second section, then the fragment was constructed based on clone overexpression and RNAi vector, transfected DF1 cell line, and to build a stable expression strain. Firstly, on the basis of immunofluorescence staining, then the cells were stimulated with LPS, to observe the expression of genes related to changes in trend. Finally, expression cell line of NLRC5 and the signal pathway in the stable The expression patterns of key genes were compared and analyzed, and put forward the hypothesis of the regulatory mechanism of.NLRC5 cloning and analysis of the results of NLRC5 and the key genes showed that chicken NLRC5 and mammalian NLRC5 genes have homology, and the mammalian gene function may be inferred by entering the nucleus can play a role, and by the expression of the table key epigenetic modifications of.NLRC5 gene and its signal pathway in cell and tissue found that NF- kappa B2 and NLRC5. The co expression of STAT1 and NLRC5 was linear, the chicken has no obvious co expression characteristics, presumably the regulation of NLRC5 is complex or there may be no regulation, and STAT3 may act as a negative regulator NF- kappa B2 negative feedback regulation of the expression of NLRC5. In conclusion, in addition to MHCI, NLRC5 may regulate the expression of STAT3 and IL18. NLRC5 may have to feed poultry The mechanisms that regulate the expression of.3. in the fourth chapter, different animal milk, the specific process for the further understanding of NLRC5 regulated, and further prove that the control effect of chicken NLRC5 on MHCI, SPI-2 in type III secretion system to lay the foundation for research on latent infection and formation of bacteria carrying state and help better, this study investigated the NLRC5 start codon before the activity of different fragments of 4372bp region, and to analyze the potential regulation principle of.NLRC5 series deletion results show that NLRC5 may have 2 active promoter region, and by 617 to 1448 of the area between the isolation of two promoter activity; second promoter activity than the first promoter the activity in this area, and found that the NF- kappa B binding motif is reported and the expression and regulation of NLRC5 transcription factors closely related to the 1448 to 4372 of the fragments may be the spleen The main start organization play a role in the transcriptional expression of NLRC5 fragments, mainly from the beginning of 2108; the downstream regulation may be more complex and have a significant impact on the expression of downstream components of NLRC5; 1-2108 of the activity was only 1448-4372 of about 50%, which indicates that the activity of the 1448-2108 than the 1828 to 2108 and low activity. That 1828 to 2213 activity will be further enhanced, even more than 1448 to 4372 of the activity, so that the NLRC5 core promoter is 1828 to 2213 pieces, the interval contains the element may have STAT1, P300, NF- kappa B, MRF2, FAC1, PAX5, CRFB, CPBP, NF-AT1, MRF-2, AML1 FAC1, CRX, RelA-p65., STAT1 and NF- were in the kappa B transcription start site of 182bp before the starting position was identified. The future research should confirm the mechanism of poultry NLRC5 on regulation of MHCI, by further ChIP and ChIP-Seq binding to the promoter of NLRC5 Regional analysis of the whole genome, RNA-Seq analysis can more accurately find out whether the gene of STAT3, MHCI and IL18 are regulated by EMSA gene, point mutation and knockout and overexpression experiments to further confirm the NLRC5 regulation may exist. Regulation of the final turn to Salmonella pullorum SPI-2 type III secretion system of macrophages in the MHCI pathway, in order to reveal the latent infection of Salmonella and carrier status of the formation of the specific mechanism.

【學(xué)位授予單位】：揚(yáng)州大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2015
【分類號】：S852.4

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