本文選題：水牛 + 泌乳期和非泌乳期　； 參考：《廣西大學》2016年博士論文

【摘要】：奶水牛是熱帶亞熱帶地區(qū)重要的產(chǎn)奶動物,水牛奶占世界牛奶供給量的5%以上。水牛奶的乳脂率和乳蛋白含量是荷斯坦奶牛的2.22和1.72倍。因此,研究奶水牛泌乳生理的基因調(diào)控網(wǎng)絡可以揭示奶水牛高營養(yǎng)形成的分子機制。MicroRNAs(miRs)是一類非編碼的19-25nt的小RNA,通過與靶基因mRNA結合阻遏靶mRNA翻譯或降解靶mRNA來調(diào)控蛋白的表達。據(jù)估算,30%的蛋白編碼基因受到miR的調(diào)控。研究發(fā)現(xiàn)miRs在動物細胞分化、增殖和凋亡過程中發(fā)揮重要作用。成年動物乳腺組織可以經(jīng)歷細胞增殖、分化、去分化和凋亡的循環(huán),是研究乳腺組織泌乳和生理的理想分子機制模型。對人和小鼠的研究結果表明人的乳腺特異性miRs有23個,而小鼠的乳腺特異性miRs只有9個,表明miRs在乳腺中發(fā)揮重要作用。目前研究主要是關于miRs在其他產(chǎn)奶家畜泌乳生理中的作用,水牛的泌乳期和非泌乳乳腺組織miRs表達譜尚無報導。本研究首次對水牛泌乳期和非泌乳期乳腺組織中miRs的表達譜和作用進行了研究,構建了奶水牛泌乳期和非泌乳期乳腺組織的miR表達譜,分析和驗證了 18個miR的差異表達模式,對篩選出的差異顯著表達bbu-miR-103和novel-miR-57及其靶向基因和功能進行了研究,以便為闡明奶水牛泌乳物質(zhì)及能量代謝通路的分子作用機制奠定工作基礎。1.水牛泌乳期和非泌乳期乳腺組織miRNA差異表達譜的構建及分析采集水牛泌乳高峰期和非泌乳期(干乳期)的乳腺樣本,利用Solexa高通量測序技術構建了水牛泌乳期以及非泌乳期乳腺組織2個miRNA表達譜,獲得非泌乳期和泌乳期18nt-31nt的12,569,467和12,768,110條高質(zhì)量序列。統(tǒng)計顯示,中國沼澤型水牛泌乳和非泌乳期的sRNA分布的寬度模式介于18-31nt之間,在22nt達到一個高峰值,其中泌乳期乳腺組織中22nt序列占總sRNA數(shù)量的33.4%。在miRbase17.0庫中檢測到成熟miRs和pre-miR分別是676個和662個,歸屬500個microRNA基因家族。本研究在水牛乳腺組織測序確認的成熟miRs和全部pre-miR分別為359個和363個,歸屬為259個miRs家族;從新發(fā)現(xiàn)的262個候選miR中鑒定出230個水牛新miRs,其中5個是水牛特有的miRs。對所有鑒定的miRNA的第一個核苷酸偏好性分析顯示,U是19nt和25nt miRs的5'端最普遍的核苷酸(94.15%和97.90%)。統(tǒng)計其在染色體上的分布情況,發(fā)現(xiàn)68.77%的已知miR和84.69%的新miR位于常染色體上,成功定位于乳腺組織染色體上的基因間隔區(qū)。已知miR主要分布在21號和X染色體,分別為74和37個占總數(shù)量的635個的11.65%和5.83%。新發(fā)現(xiàn)的miR,在21號和X染色體上分別為38個和32個�？俶iR分布在常染色體和X染色體上,密度從0.09到1.05 miRs/Mbp不等。21號染色體是miRs主要表達的染色體,分布最多,總miRs,泌乳期表達miRs和非泌乳期miRs分別為1.05個/Mbp,1.34個/Mbp和1.35個/Mbp。泌乳期和非泌乳期miRs在相同染色體上的分布密度基本相同。2.水牛泌乳期和非泌乳期乳腺組織miRs差異表達譜分析在非泌乳期,bbu-miR-148a,bbu-let-7b,bbu-let-7a,bbu-miR-21,bbu-miR-143,bbu-miR-200c,bbu-miR-26a,bbu-miR-200a 和 bbu-let-7f 是主要表達 miRs,組成了 總已知miRNA序列的53.8%,每個miRs讀數(shù)大于20,000個序列,表明它們是非泌乳期組織中高豐度表達miRs。在泌乳期發(fā)現(xiàn)有7個高表達的miRs(bbu-let-7b,bbu-let-7a,bbu-miR-26a,bbu-miR-125b,bbu-miR-21,bbu-miR-29a 和 bbu-let-7c),每個有 20,000 多條序列表達。比較兩個樣品組,bbu-miR-148a,bbu-miR-143,bbu-miR-200a,bbu-miR-141 和bbu-miR-30a-5p等這些miRs在泌乳期的表達量下降到非泌乳期的一半以下。而另外一些 miR,如 bbu-miR-26a,bbu-miR-29a,bbu-miR-125b,bbu-let-7c and bbu-miR-99a 在泌乳期比非泌乳期表現(xiàn)出多余或等于2倍序列豐度。根據(jù)KEGG對20個差異高表達miRNA的預測靶基因進行了功能分類。結果109個預測靶基因都被標記在MAPK信號通路,其他重要途徑有Jak-STAT、催乳素(PRL)信號轉(zhuǎn)導和胰島素信號通路等。3.泌乳期和非泌乳期差異表達miRNAs篩選及QRT-PCR驗證差異分析兩個時期miRs的表達譜,篩選出18個miRs并運用QRT-PCR對其表達量進行驗證。將其分為高豐度、低豐度和未知miRs:(a)9個高表達的已知的miR:bbu-miR-30a-5p,bbu-miR-141,bbu-miR-101,bbu-miR-103,bbu-miR-148a,bbu-miR-29a,bbu-miR-125b,bbu-miR-497 和 bbu-miR-125a;(b)4 個低豐度表達的 miR:bbu-miR-490,bbu-miR-217,bbu-miR-592 和 bbu-miR-2370*;(c)5 個高表達的未知 miR:novel-miR-57,novel-39,novel-148,novel-76 和 novel-123b。QRT-PCR 驗證結果顯示,bbu-miR-103,bbu-miR-125a、bbu-miR-30a-5p 和 bbu-miR-148a 在泌乳期表達量高于非泌乳期,其表達量分別為非泌乳期的2.43、3.43、4.06和5.29倍(P0.05);bbu-miR-29a在非泌乳期表達量為泌乳期的0.3倍,顯著低于泌乳期(P0.05);bbu-miR-141,bbu-miR-125b,bbu-miR-497和bbu-miR-101的表達量在泌乳期高于非泌乳期,但差異不顯著(P0.05)。低豐度的miR-490和miR-592在泌乳期表達量分別為非泌乳期的4.96和3.82倍,差異顯著(P0.05)。在高豐度表達的5個的未知miRNA中,novel-39,novel-148,novel-76和novel-123b在泌乳期和非泌乳期差異不顯著(P0.05),Novel-miR-57在泌乳期比非泌乳期高29.79倍(P0.05)。因此,選擇miR-103和novel-miR-57作為下一步主要研究對象。4.Bbu-miR-103靶向基因PANK3調(diào)控乳脂代謝的研究基于人免疫缺陷慢病毒系統(tǒng)的載體骨架,構建了 LPEZX-PRE-MIR-103-1前體表達克隆載體(8219 bp),攜帶 Bbu-miR-103 前體序列。將 LPEZX-PRE-MIR-103-1 與對空載體LPEZX-MIR-NC分別與NRF和VSVG在293T細胞中進行包裝和擴繁,獲得了感染滴度分別為3.42×106 PFU/mL和3.47×106 PFU/mL的復制缺陷型病毒。用組織塊法培養(yǎng)了乳腺上皮細胞,用等量對照和miR-103病毒原液感染乳腺上皮細胞48 h后,在顯微鏡下觀察有綠色熒光蛋白發(fā)光,表明miR-103及空載對照病毒顆粒已經(jīng)成功地進入了水牛乳腺上皮細胞中,代謝產(chǎn)生了過量miR-103。同時化學法合成miR抑制劑并進行轉(zhuǎn)染試驗。結果發(fā)現(xiàn)過表達和抑制表達miR-103與乳腺細胞靶基因PANK3表達呈現(xiàn)負相關。因此,推測PANK3是水牛miR-103作用的靶基因。過表達miR-103,下調(diào)了 PANK3的表達,顯著提高了 SREBP1c和ACACA的表達。測定了過表達miR-103載體對水牛脂肪代謝通路8個關鍵的步驟相關基因的影響。結果發(fā)現(xiàn),bbu-miR-103主要作用基因范圍從脂肪酸從頭合成開始、延伸至甘油三酯合成、乳脂合成和分泌以及脂肪酸吸收為止,對乳脂后續(xù)的轉(zhuǎn)運代謝無較大影響。5.Novel-miR-57在Bcap-37和水牛乳腺上皮細胞上的靶基因—DOK4的狩獵及驗證Novel-miR-57是在泌乳期和非泌乳期乳腺組織中篩選出的相對表達量差異最大的新發(fā)現(xiàn)miR,但在miRBase數(shù)據(jù)庫檢索不到類似或同族的miR。為此,我們運用MiRscan對Novel-miR-57的二級結構進行了預測。結果顯示,Novel-miR-57可能有8個莖環(huán)結構,結合自由能為-28.0kcal/mol,Novel-miR-57的成熟序列位于miR的第一個莖環(huán)上。使用自編軟件ensembl(v80)注釋mRNA截取3'UTR,篩選結合自由能小于-20kcal/mol的mRNA 3'UTR與miR成熟序列的5'端的2-8種子序列進行配對,找到34個水牛轉(zhuǎn)錄組mRNA可能是Novel-miR-57的作用靶基因。這些基因包括CYP7B1、CACNG3、DOK4、COL17A1、和 ESN1 等基因,對這些基因進行了 GO assignment和KEGG信號通路預測分析。用水牛泌乳和非泌乳期乳腺組織cDNA添加線蟲miR-39作為外參進行QRT-PCR定量分析,尋找差異表達靶基因。結果發(fā)現(xiàn)有6個基因檢測不到,RCL1、UBE3C和NFRKB等10個基因在非泌乳期和泌乳期相對表達量相似或高于泌乳期;BRMSIL、ACTL8和ADORO等11個基因在泌乳期和非泌乳期相對表達量相似或高于非泌乳期;DLX3、CANCNG3、NFKBID、C17orf53、RTN1和FBXO10等7個基因在非泌乳期表達量遠高于泌乳期,分別為128.03、144.23、146.24、160.96、160.38、274.40 和 326.24 倍,差異極顯著(P0.01),可能為主要靶基因�；瘜W合成Novel-miR-57的類似物Mimics和抑制劑Inhibitor,探討Novel-miR-57的作用。結果發(fā)現(xiàn),Bcap-37細胞系添加Novel-miR-57后能顯著抑制DOK4基因的表達(P0.01),而Inhibitor能顯著提高DOK4基因的表達(P0.01);而在水牛乳腺細胞添加100nM的Novel-miR-57后能顯著促進DOK4基因的表達(P0.01),添加200nM的Inhibitor能顯著抑制DOK4基因的表達(P0.01),與對Bcap-37細胞的作用相反。揭示了 DOK4是Novel-miR-57的作用靶基因,且Novel-miR-57能夠根據(jù)不同細胞系不同生理條件來上調(diào)和下調(diào)DOK4基因的表達,最終對泌乳新陳代謝通路產(chǎn)生作用。結論:高差異表達Bbu-miR-103在水牛乳腺組織中的靶基因是PANK3。Bbu-miR-103通過從頭合成途徑促進脂肪酸合成,對水牛乳腺上皮細胞的甘油三脂合成、乳脂滴合成和分泌、脂肪酸激活等3個乳脂代謝步驟有促進作用。Novel-miR-57在Bcap-37細胞和水牛乳腺中的靶基因為DOK4,與水牛乳腺上皮細胞的分化有關。
[Abstract]:Milk buffalo is an important milking animal in tropical subtropical region. Milk and milk account for more than 5% of the world's milk supply. The milk fat rate and milk protein content are 2.22 and 1.72 times of Holstein cows. Therefore, the study of the molecular mechanism.MicroRNAs (miRs) of the high nutrition formation of milk and buffalo can be revealed by the study of the gene regulation network of milk buffalo lactating physiology. It is a small RNA of a class of non coded 19-25nt, which regulates protein expression by binding to target gene mRNA to inhibit target mRNA translation or degradation of target mRNA. It is estimated that 30% of the protein encoding gene is regulated by miR. Studies have found that miRs plays an important role in the process of differentiation, proliferation and apoptosis in animal cells. Adult mammary tissues can undergo fine experience. The cycle of cell proliferation, differentiation, dedifferentiation and apoptosis is an ideal molecular mechanism for the study of mammary tissue lactation and physiology. The results of human and mouse studies show that there are 23 human breast specific miRs and only 9 mammary specific miRs in mice, indicating that miRs plays an important role in the mammary gland. The present study is mainly about miRs in it The miRs expression profiles of the lactating and non lactation mammary tissues of buffalo were not reported. The expression and function of miRs in the lactation and non lactation tissues of buffalo were first studied for the first time. The miR expression profiles of milk buffalo lactation and non lactation mammary tissues were constructed and analyzed and verified. The differential expression patterns of 18 miR were used to express bbu-miR-103 and novel-miR-57 and their target genes and functions, so as to lay a work foundation for clarifying the molecular mechanism of lactating substance and energy metabolism pathway of milk Buffalo,.1. buffalo lactation and non lactating breast tissue miRNA differential expression profiles By constructing and analyzing mammary samples of buffalo lactation peak and non lactation period (dry milk period), 2 miRNA expression profiles of buffalo lactation and non lactation mammary tissues were constructed by Solexa high throughput sequencing technology, and 12569467 and 12768110 high quality sequences of 18nt-31nt in non lactation and lactation period were obtained. The width pattern of sRNA distribution in the lactation and non lactation period of the lustrate buffalo is between 18-31nt and a high peak in 22nt, of which the 22nt sequence of the mammary gland in the lactation period is 676 and 662 respectively, which belong to 500 microRNA gene families in the miRbase17.0 library, and belong to 500 microRNA genes. Bovine mammary gland tissue sequencing confirmed that mature miRs and total pre-miR were 359 and 363, belonging to 259 miRs families, and 230 New Buffalo miRs were identified from 262 newly found candidate miR, 5 of which were the first nucleotide bias analysis of all identified miRNA by Buffalo specific miRs., and U was the largest 5'end of 19nt and 25nt miRs. The common nucleotides (94.15% and 97.90%). The distribution of the chromosomes on the chromosomes shows that 68.77% of the known miR and 84.69% of the new miR are located on the autosomes, which are located successfully on the gene spacers on the chromosomes of the mammary tissues. The known miR is mainly distributed on the 21 and X chromosomes, divided into 74 and 37 of 635 of the total number of 11.65% and 5.83%.. The newly discovered miR was 38 and 32 on the 21 and X chromosomes. The total miR was distributed on the autosomes and the X chromosomes, and the density from 0.09 to 1.05 miRs/Mbp was the main chromosomes of miRs, the most distributed, the total miRs, the miRs and the non lactation miRs were 1.05 /Mbp, 1.34 /Mbp and 1.35 secreting periods. The distribution density of miRs on the same chromosome in milk and non lactation period was basically the same as that of.2. buffalo lactating and non lactating mammary gland tissue miRs differential expression profiles in non lactation period, bbu-miR-148a, bbu-let-7b, bbu-let-7a, bbu-miR-21, bbu-miR-143, bbu-miR-200c, bbu-miR-26a, bbu-miR-200a and bbu-let-7f were mainly expressed. 53.8% of the total known miRNA sequences, each miRs reading is greater than 20000 sequences, indicating that they are high abundance expression of miRs. in the non lactation period and found 7 high expression miRs (bbu-let-7b, bbu-let-7a, bbu-miR-26a, bbu-miR-125b, bbu-miR-21, bbu-miR-29a and bbu-let-7c) in lactation period, each of which has more than 20000 sequences. The two sample groups, such as bbu-miR-148a, bbu-miR-143, bbu-miR-200a, bbu-miR-141 and bbu-miR-30a-5p, decreased to less than half of the lactation period, and some other miR, such as bbu-miR-26a, bbu-miR-29a, bbu-miR-125b, bbu-let-7c and, showed excess or equal to 2 in lactation period than non lactation period. Sequence abundance. According to KEGG, the functional classification of the predicted target genes of 20 differentially expressed miRNA was carried out. Results 109 target genes were labeled in MAPK signaling pathway, other important pathways were Jak-STAT, PRL signal transduction and insulin signaling pathway and other.3. lactation and non lactation period differential expression miRNAs screening and QRT-PCR Verify the expression profile of miRs in the two period of difference analysis, select 18 miRs and verify its expression using QRT-PCR, and divide it into 9 known miR:bbu-miR-30a-5p, bbu-miR-141, bbu-miR-101, bbu-miR-103, bbu-miR-148a, bbu-miR-29a, bbu-miR-125b, bbu-miR-125b, and miRs:. 125A; (b) 4 low abundances of miR:bbu-miR-490, bbu-miR-217, bbu-miR-592 and bbu-miR-2370*; (c) 5 highly expressed unknown miR:novel-miR-57, novel-39, novel-148, novel-76 and novel-123b.QRT-PCR verification results showed that the expression level in lactation is higher than that of non lactation. The expression of bbu-miR-29a in non lactation period was 2.43,3.43,4.06 and 5.29 times (P0.05), and the expression of bbu-miR-29a in non lactation period was 0.3 times of lactation period, significantly lower than lactation period (P0.05); bbu-miR-141, bbu-miR-125b, bbu-miR-497 and bbu-miR-101 were higher than non lactation period in lactation period, but the difference was not significant (P0.05). MiR-490 and low abundance of miR-490 were not significant. The expression of miR-592 in lactation period was 4.96 and 3.82 times of non lactation period, respectively (P0.05). In 5 unknown miRNA with high abundance, novel-39, novel-148, novel-76 and novel-123b were not significantly different in lactation and non lactation period (P0.05), Novel-miR-57 was 29.79 times higher than non lactation period (P0.05). Therefore, select miR-103 And novel-miR-57 as the next major research object,.4.Bbu-miR-103 targeting gene PANK3 regulation of lipid metabolism, based on the carrier skeleton of the human immunodeficiency lentivirus system, constructed the LPEZX-PRE-MIR-103-1 precursor expression clone carrier (8219 BP), carrying the Bbu-miR-103 precursor sequence. LPEZX-PRE-MIR-103-1 and LPEZX-M to the empty body LPEZX-M. IR-NC was packed and expanded in 293T cells respectively with NRF and VSVG, and the infection titers were 3.42 * 106 PFU/mL and 3.47 * 106 PFU/mL respectively. The mammary epithelial cells were cultured by tissue block method. After infection of 48 h of mammary epithelial cells with equal control and miR-103 virus, the green fluores were observed under the microscope. Photoluminescence shows that miR-103 and unloaded control virus particles have been successfully entered into the buffalo mammary epithelial cells. Metabolic production of excessive miR-103. and chemical synthesis of miR inhibitors and transfection tests have been carried out. The results showed that overexpression and inhibition of expression of miR-103 were negatively correlated with the expression of PANK3 in mammary gland cells. Therefore, it was presumed that the expression of miR-103 was negatively correlated with the expression of PANK3 in the mammary gland cells. PANK3 is the target gene for the action of buffalo miR-103. Over expression of miR-103, down down the expression of PANK3 and significantly improving the expression of SREBP1c and ACACA. The effects of over expressed miR-103 vectors on the 8 key steps related genes of the buffalo fat metabolism pathway were determined. The results showed that the bbu-miR-103 main function gene range was synthesized from ab initio from fatty acids. Beginning, extending to triglyceride synthesis, milk fat synthesis and secretion, and fatty acid absorption, the hunt for the target gene of.5.Novel-miR-57 on Bcap-37 and buffalo mammary epithelial cells - the hunting and verifying that Novel-miR-57 is the relative expression of Novel-miR-57 in lactation and non lactation mammary tissues The new discovery miR has the largest difference in quantity, but we do not find similar or family miR. in the miRBase database. We use MiRscan to predict the two structure of Novel-miR-57. The result shows that Novel-miR-57 may have 8 stem ring structures, and the free energy is -28.0kcal/mol, and the mature sequence of Novel-miR-57 is located on the first stem ring of miR. Using the self-made software Ensembl (V80) annotation mRNA to intercept the 3'UTR and screen the 2-8 seed sequences of mRNA 3'UTR with the free energy less than -20kcal/mol and the 5'end of the mature miR sequence, and find that the mRNA of the 34 buffalo transcriptome may be the Novel-miR-57 target gene. These genes were predicted by GO assignment and KEGG signaling pathway. QRT-PCR quantitative analysis was carried out by adding nematode miR-39 in the breast tissue of buffalo and non lactation period cDNA to find the differential expression target gene. The results showed that 6 genes were not detected, and 10 genes such as RCL1, UBE3C and NFRKB were found in non lactation and lactation phases. The expressions of 11 genes, such as BRMSIL, ACTL8 and ADORO, were similar or higher than non lactation periods in lactation and non lactation periods, and 7 genes, such as DLX3, CANCNG3, NFKBID, C17orf53, RTN1 and FBXO10, were much higher in non lactation periods than in lactation period, which were 128.03144.23146.24160.96160.38274.40 and 326.2, respectively. 4 times, the difference is very significant (P0.01), may be the main target gene. Chemical synthesis of Novel-miR-57 analogues Mimics and inhibitor Inhibitor, to explore the role of Novel-miR-57. The results showed that the Bcap-37 cell line adding Novel-miR-57 can significantly inhibit the expression of DOK4 gene (P0.01), and Inhibitor can significantly increase the DOK4 gene expression (P0.01); and The addition of 100nM Novel-miR-57 to the buffalo mammary cells can significantly promote the expression of DOK4 gene (P0.01). The addition of 200nM Inhibitor can significantly inhibit the expression of the DOK4 gene (P0.01), which is contrary to the effect on Bcap-37 cells. It is revealed that DOK4 is the target gene of Novel-miR-57, and Novel-miR-57 can be based on different physiological conditions in different cell lines. To up-regulation and downregulate the expression of DOK4 gene and ultimately to the metabolic pathway of lactation. Conclusion: the target gene of high differential expression of Bbu-miR-103 in Buffalo mammary gland is that PANK3.Bbu-miR-103 promotes fatty acid synthesis through the ab initio pathway, glycerol three fat synthesis, lipid droplet synthesis and secretion, and fat drop in the mammary gland epithelial cells of Buffalo 3 lipid metabolism steps, such as acid activation, can promote the role of.Novel-miR-57 in the Bcap-37 cells and the buffalo mammary gland because of DOK4, which is related to the differentiation of the mammary epithelial cells of buffalo.
【學位授予單位】：廣西大學
【學位級別】：博士
【學位授予年份】：2016
【分類號】：S823.83

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