本文選題：棗 + 糖��； 參考：《西北農林科技大學》2016年博士論文

【摘要】：棗(Ziziphus jujuba Mill.)是鼠李科最重要的經濟樹種,其果實營養(yǎng)豐富,含糖量高,素有“木本糧食”之稱。果實糖酸含量影響品質及商品價值。本研究通過測定糖、酸和抗壞血酸等生理指標,結合基因組數據,利用轉錄組測序和熒光實時定量PCR分析,從分子水平上闡明了棗和酸棗糖、酸、抗壞血酸形成與積累的遺傳與生理本質,深化了對棗和酸棗糖酸積累差異的理解。同時,利用全基因組重測序技術研究了酸棗到棗的糖酸馴化,明確糖酸馴化關鍵基因。本研究為棗果實品質形成和新品種選育提供分子理論基礎。主要研究結論如下:1.蔗糖是棗果實糖積累的最主要形式,積累時期始于白熟期。研究鑒定了5個棗蔗糖合成關鍵基因(ZjSPS1、ZjSPS2、ZjSS1、ZjSS2和ZjSS3)。糖代謝關鍵基因中,轉化酶類基因(vINV、nINV和cINV)在棗果實的低表達是其蔗糖含量高于果糖和葡萄糖含量的重要原因;而轉化酶基因及己糖基因(HK3和HK6)在酸棗的高表達降低了其糖分積累。本研究鑒定了棗果實發(fā)育過程中糖分積累的關鍵基因,揭示了果實糖分以蔗糖為主的重要原因,為果實糖分積累期的管理及提高含糖量提供生產指導。2.本研究共鑒定得到83個糖轉運蛋白基因,其中兩個參與山梨醇的運輸。ZjSWEET2和ZjSWEET15在花和果實中特異表達�；虻谋磉_模式揭示ZjSUC2和ZjSWEE2促進了果實的蔗糖積累;單糖轉運蛋白ZjSTP12,ZjSTP16,ZjpGlcT3,SWEET15,ZjSWEET20和ZjTMT2基因也對果實糖分積累起到重要作用。與其直系同源基因(葡萄和番茄)相比,ZjSTP12,ZjSTP16,ZjSUC2和ZjSWEET2特異在棗果實中高表達,這可能是棗果實含糖量高于其他果實的重要原因。另外,糖轉運蛋白基因在果實發(fā)育不同階段的葉片表達水平也具有差異性。本研究深化了糖轉運蛋白生物學功能的理解,明確了棗果實糖轉運機理,為紅棗優(yōu)質栽培和新品種選育提供了理論基礎。3.棗果實的有機酸類型主要為蘋果酸,酸棗的為蘋果酸和檸檬酸;本研究鑒定的9個棗和酸棗表達顯著差異的酸代謝關鍵基因是酸棗果實含酸量高于棗的重要原因。此外,ZjCS3促進了棗和酸棗檸檬酸的積累,果實中特異表達的蘋果酸合酶ZjMS及蘋果酸脫氫酶ZjMDH12促進棗和酸棗的蘋果酸合成,ZjMT3和ZjMT11促進了蘋果酸的運輸。本研究揭示了野生型酸棗味道極酸的重要原因,探究了導致有機酸積累的關鍵基因,為定向選育過程中淘汰酸性口味的種質提供分子基礎。4.基于32個樣品的全基因組重測序,找到了4個酸代謝關鍵基因、13個糖轉運蛋白及3個糖代謝相關基因。馴化關鍵基因的選擇,揭示了棗在栽培化過程中,糖代謝基因尤其是糖轉運蛋白的受選擇,對棗果實高含糖量的積累具有重要作用;酸代謝基因的選擇效應對馴化過程中酸積累的抑制具有重要貢獻。酸棗到棗糖酸馴化機制的解析,對于優(yōu)化種質、提升含糖量及利用酸棗多樣性選育新品種提供了分子依據,為其他果樹的風味馴化研究提供理論模型。5.棗抗壞血酸(AsA)主要在果實發(fā)育早期合成,成熟期已趨于穩(wěn)定,且果實的含量遠高于其他組織。研究深化了棗果實中的兩條AsA合成途徑—半乳糖途徑和肌醇途徑:在果實發(fā)育早期半乳糖途徑對AsA的合成起決定作用,并且GMP1、GME1、GGP、GPP和GaLDH是合成AsA的關鍵基因;肌醇通路及半乳糖通路中的GMP2和GME2基因在棗果實成熟期AsA含量的維持起重要作用。AsA再循環(huán)通路中,MDHAR和DHAR對維持棗AsA含量起協(xié)同作用。本研究闡明了棗果實AsA合成關鍵基因和關鍵通路。AsA代謝機制的研究對提高果實抗壞血酸含量、改善果實品質具有重要指導意義。
[Abstract]:Ziziphus jujuba Mill. is the most important economic tree of rhamnarifamily. Its fruit is rich in nutrition and high in sugar content. It is known as "woody grain". The content of fruit sugar and acid affects the quality and commodity value. In this study, the physiological indexes such as sugar, acid and ascorbic acid were measured, combined with genomic data, the transcriptional sequence and fluorescence real-time quantitative P were used. CR analysis, from the molecular level, clarified the genetic and physiological nature of jujube and jujube sugar, acid, ascorbic acid formation and accumulation, and deepened understanding of the accumulation difference between jujube and Zizyphus jujube. At the same time, the whole genome re sequencing technology was used to study the acclimatization of sour jujube to jujube and the key genes of acclimation of sugar and acid. This study was the quality of jujube fruit. The main conclusions are as follows: 1. sucrose is the main form of sugar accumulation in jujube fruit, and the accumulation period begins in the white ripening period. 5 key genes of sucrose synthesis (ZjSPS1, ZjSPS2, ZjSS1, ZjSS2 and ZjSS3) are identified. In sugar metabolism key genes, the invertase genes (vINV, nINV and cINV) are in jujube fruit. The actual low expression is the important reason for the sugar content higher than the fructose and glucose content, and the high expression of the invertase gene and hexose gene (HK3 and HK6) in the sour jujube decreased the sugar accumulation. This study identified the key genes of sugar accumulation during the development of jujube fruit, and revealed the important reason for the fruit sugar to be mainly sucrose. The management of solid sugar accumulation period and the improvement of sugar content provided production guidance.2., 83 sugar transporter genes were identified. Two of them were specifically expressed in the flower and fruit of.ZjSWEET2 and ZjSWEET15. The gene expression pattern revealed that ZjSUC2 and ZjSWEE2 promoted the accumulation of sucrose in fruit; monosaccharide transporter Zj STP12, ZjSTP16, ZjpGlcT3, SWEET15, ZjSWEET20 and ZjTMT2 genes also play an important role in the accumulation of fruit sugar. Compared to their direct homologous genes (grapes and tomatoes), ZjSTP12, ZjSTP16, ZjSUC2 and ZjSWEET2 are highly expressed in jujube fruit, which may be an important reason for the sugar content of jujube fruit higher than that of other fruits. This study deepened the understanding of the biological function of sugar transporter and clarified the mechanism of sugar transport in jujube fruit, and provided a theoretical basis for high quality and new varieties of jujube fruit. The organic acid type of jujube fruit, malic acid and lime, was the main organic acid type of jujube fruit,.3. and lime. Citric acid; the important acid metabolism key gene of the 9 jujube and Zizyphus jujube expressed in this study was the important reason for the acid content of jujube fruit higher than that of jujube. In addition, ZjCS3 promoted the accumulation of citric acid in jujube and Zizyphus jujube, and the specific expression of malic acid synthase ZjMS and malate dehydrogenase ZjMDH12 in fruit to promote the synthesis of malic acid in jujube and Zizyphus jujube, ZjMT3 And ZjMT11 promoted the transport of malic acid. This study revealed the important cause of the wild sour jujube flavor acid, and explored the key genes that lead to the accumulation of organic acids, providing a molecular basis for the elimination of acid flavors in the directional selection process,.4. based on the whole genome re sequencing based on 32 samples, and found 4 key genes for acid metabolism, 13 The selection of key genes, sugar transporter and 3 glycometabolism related genes, reveals that the selection of sugar metabolism genes, especially sugar transporters, plays an important role in the accumulation of high sugar content in jujube fruit, and the selection effect of acid metabolism gene has an important contribution to the inhibition of acid accumulation during the domestication process. The analysis of the acclimatization mechanism of jujube to jujube acid has provided a molecular basis for optimizing the germplasm, raising the sugar content and using the variety of Zizyphus jujuba, and providing a theoretical model for the study of the flavor domestication of other fruit trees,.5. ascorbic acid (AsA), which is mainly synthesized at the early stage of fruit development, and is stable at maturity, and the content of fruit is much higher than that of its fruit. The study deepened two AsA synthesis pathways in jujube fruit - galactose pathway and inositol pathway: in the early stage of fruit development, galactose pathway plays a decisive role in the synthesis of AsA, and GMP1, GME1, GGP, GPP and GaLDH are the key genes for AsA, and the GMP2 and GME2 genes in the inositol pathway and galactose pathway are AsA in date fruit maturity. The maintenance of content plays an important role in the.AsA recirculation pathway. MDHAR and DHAR play a synergistic role in maintaining the AsA content of jujube. This study illustrates that the key genes of AsA synthesis and the mechanism of.AsA metabolism in the key pathway of jujube fruit are of great importance to improve the content of ascorbic acid and improve the quality of fruit.
【學位授予單位】：西北農林科技大學
【學位級別】：博士
【學位授予年份】：2016
【分類號】：S665.1

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