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小麦籽粒高分子量谷蛋白亚基(HMW-GS)积累规律及其调控研究

Accumulations of High Molecular Weight Glutenin Subunits and Glutenin Macropolymer and Their Regulatory Approaches in Wheat Grain

【作者】 伯云

【导师】 姜东;

【作者基本信息】 南京农业大学 , 作物栽培与耕作学, 2009, 硕士

【摘要】 谷蛋白小麦籽粒蛋白的重要组成部分,与小麦品质关系密切。其亚基根据分子量大小又可分为高分子量谷蛋白亚基(HMW-GS)和低分子量谷蛋白亚基(LMW-GS)。小麦高分子量谷蛋白亚基与小麦加工品质密切相关。不同小麦品种的麦谷蛋白亚基组成和数目存在较大差异,且受遗传控制并具有品种稳定性,但其含量(表达量)又受环境条件的影响。本文以不同小麦品种为材料,研究RIL群体各家系中小麦高分子量谷蛋白亚基的积累动态,以及其在不同粒位籽粒中的差异,并研究了环境因素对小麦籽粒HMW-GS的影响,以期用不同栽培措施调控小麦HMW-GS积累,为深化小麦籽粒HMW-GS形成机制与栽培措施调控技术研究提供依据。研究共涉及4方面试验。试验1:主要研究重组自交家系中T6VS·6AL染色体对HMW-GS和GMP含量的影响,及不同地点生态环境下的HMW-GS和GMP含量差异。以含T6VS-6AL染色体的普通小麦-簇毛麦品种92R137和辉县红为亲本,通过单粒传法构建的F8重组自交家系为试材。于2005年分别于南京农业大学农场和郑州河南农业大学农场种植。试验2:主要研究了小麦强、弱势粒中HMW-GS和GMP形成和积累动态及其差异。试验于2006-2007年在江苏省农科院进行。选用扬麦158和扬麦11两个品种。以中部6-8小穗的基部第1、2位籽粒为强势粒,对应小穗顶部第1个籽粒为弱势粒。试验3:主要研究遮荫对小麦籽粒HMW-GS和GMP形成和积累动态的影响。试验选用两个小麦品种:扬麦158、扬麦11,于2005-2006和2006-2007两年度在江苏省农科院进行。2005-2006年度试验共设3个处理:不遮荫(对照,S0)、中度遮荫(S2)和重度遮荫(S3),2006-2007年度试验增加了一个轻度遮荫处理(S1)。S1、S2和S3的冠层上部光强分别约为自然光强(S0)的90%、82%和75%。拔节期至成熟实施遮荫处理。试验4:主要研究高温和干旱处理对于小麦籽粒HMW-GS和GMP积累的影响。试验于2007年在丹麦奥尔胡斯大学进行。在两个生长阶段进行水分亏缺和高温胁迫处理:在二棱末期实施对照(CKA)、干旱(WDA)和高温(HTA)3个处理,在开花期时每个处理又分别进行对照(CKB)、干旱(WDB)和高温(HTB),共计9个处理。主要研究结果如下:1.不同生态地点下含T6VS·6AL染色体的RIL群体内各家系HMW-GS和GMP的积累Glu-A1、Glu-B1和Glu-D1三对基因在RIL群体中均完全符合1:1分离,HMW-GS组成受遗传控制,属于质量性状。HMW-GS组成类型对其含量具有影响。T6VS·6AL易位系的存在对于HMW-GS和GMP含量的影响不明显,但不同种植地点生态条件对于HMW-GS和GMP的含量具有显著影响。2.小麦强、弱势粒中HMW-GS和GMP形成和积累动态及其差异强势粒中HMW-GS起始积累早于弱势粒,积累量始终高于弱势粒;HMW-GS含量在灌浆前中期高于弱势粒,但成熟期时却低于弱势粒,这是由于弱势粒干物质积累不足引起粒重下降造成的HMW-GS含量的相对增加。3.遮荫对小麦籽粒HMW-GS和GMP积累动态的影响遮荫使HMW-GS的起始形成时间提早,18%(S2)和25%(S3)重度遮荫处理缩短了亚基的快速积累期,降低灌浆后期积累速度,成熟期单粒总HMW-GS积累量低于对照,但两个品种成熟期籽粒HMW-GS和GMP含量则表现为遮荫(S2、S3)高于对照,且随遮荫程度的增加而增加,表明HMW-GS和GMP含量升高是籽粒干物重减少造成。与之不同,轻度遮荫(10%)延长了籽粒HMW-GS快速积累期,且成熟期亚基的积累量和含量均高于对照。遮荫处理影响小麦籽粒HMW-GS积累,但其效应与遮荫强度有关。4.高温和干旱处理对小麦籽粒HMW-GS和GMP积累动态的影响HMW-GS的积累量和含量变化规律相近。高温和干旱处理均促进了亚基的提早出现,促进了前期积累,尤其是二棱末期的高温处理。所有的处理均不同程度得影响了HMW-GS的积累速度与积累期持续时间,并影响最终积累量。两个阶段的干旱处理均增加了HMW-GS的最终积累量;高温的影响因处理时期不同而异,二棱末期高温处理减少HMW-GS积累而开花期促进HMW-GS的积累。二棱末期的干旱处理强化了开花期各处理对于亚基积累的促进,而高温处理则对开花期处理具有缓解作用。5.小麦籽粒HMW-GS和GMP含量的相关关系本研究表明,在小麦籽粒灌浆期间,不同基因位点(Glu-A1、Glu-B1及Glu-B1)编码的高分子量谷蛋白亚基、亚基对及总HMW-GS含量均与GMP含量呈显著的正相关关系,HMW-GS的组成与含量都与GMP积累关系密切。GMP积累同样受环境影响。

【Abstract】 The amount of glutenin macropolymer (GMP) in wheat flour correlates closely with baking quality, and it can be divided into high molecular weight glutenin subunits (HMW-GS) and low molecular weight (LMW-GS) glutenin subunits. So HMW-GS may play key roles in determining the formation and amount of GMP in wheat grain, and its composition and amount is relative to the wheat quality. Composition and amount of HMW-GS in different wheat varieties were different, which present inheritance stabilization. But the content of HMW-GS is affected by environment. In the present study, a RIL population grown in two locations was used to study the effect of T6VS-6AL and environment conditions on HMW-GS content, and different wheat varieties were grown at different cultivation (shading, high temperature and water deficit stress) to study the regulatory efforts of HMW-GS and GMP, and cultivation practices on content and accumulation of HMW-GS and GMP. Besides, the accumulation pattern of HMW-GS and GMP in superior and inferior grains were also studied. The results should help clarify HMW-GS and GMP accumulation mechanism and provide regulatory approaches for super quality wheat. Total experiments were establish as:Experiment 1:The aim of this experiment was to study the effect of wheat-Haynaldia villosa T6VS-6AL and environment conditions of different locations on HMW-GS content. In this experiment, an elite wheat landrace Huixianhong, which had played important role in Chinese wheat breeding, was selected to hybridize with T6VS-6AL translocation line 92R137 and a new F8 RIL population was subsequently constructed from its progenies by single seed descendant (SSD). The populations were grown in two regions, Nanjing, Jiangsu province and Zhengzhou, Henan province. Experiment 2:The aim of this experiment was to study the accumulation pattern in superior and inferior grains, and the differences between them. Two wheat cultivars owning the same HMW-GS type (7+8 and 2+12) as Yangmai 11 and Yangmai158 were planted in a field experiment to study the difference in accumulation of HMW-GS in superior and inferior grains.Experiment 3:The aim of this experiment was to study the effect of shading on accumulation of HMW-GS and GMP. Field experiment was conducted at the experimental station of Jiangsu Academy of Agricultural Sciences for two years. Two wheat varieties of Yangmai158, Yangmai11 were grown. In the first year, there were treatments as CK (SO), 82% light intensity (S2),75% light intensity (S3) from jointing to maturity. And the second year, a weaker shading treatment with 90% light intensity (S1).Experiment 4:The aim of this experiment was to study the effect of high temperature and water deficit on HMW-GS and GMP accumulation. The experiment was conducted in the semifield of Aarhus University, Denmark. Spring wheat (Triticum aestivum L. cv. Vinjett) was planted in a pot experiment. The plants were separately subject to water deficit (WDA), or high temperature (HTA), or control (CKA) conditions at the end of spilelet initiation. At anthesis, the plants of these treatments were separately subjected to water deficit (WDB), or high temperature (HTB), or control (CKB) conditions again. Thus, total nine treatments were set in this experiment, including CKA-CKB, CKA-WDB, CKA-HTB, WDA-CKB, WDA-WDB, WDA-HTB, HTA-CKB, HTA-WDB and HTA-HTB.The main results were as follows:The HMW-GS and GMP content of the RIL population. The results showed that there were 8 HMW-GS composition types in the RIL population and both subpopulations, and the HMW-GS composition was decided by genotype. There were differences between the contents of the 8 types. There was no significant correlation between HMW-GS content and T6VS-6AL, while the HMW-GS content was significantly effected by the environment of different regions.HMW-GS and GMP accumulation in superior and inferior grains. The initial timing of HMW-GS accumulation in the superior grains was earlier with a higher accumulation amount than the inferior ones. The HMW-GS content was lower before 30d after anthesis, while was higher at maturity in inferior grains, compared with the superior grains was, though it was.Shading effects on accumulation of HMW-GS and GMP in wheat grain. In the present experiment, shading showed determinate impact on the formation timing of HMW-GS in wheat grain. Shading between jointing and maturity promoted the initial formation timing of HMW-GS. The severe shading treatments (18% and 25%) shortened the rapid accumulation period of HMW-GS and reduced accumulating rate of total HMW-GS during late grain filling period. Thus, the accumulation amount of total HMW-GS at maturity was lower under these treatments. However, contents of HMW-GS and GMP were higher under severe shading than the control. A low-degree shading treatment (10%) prolonged the rapid accumulation period of HMW-GS, and increased the final accumulation amount and content of total HMW-GS. Shading from jointing to maturity obviously affected the accumulation amounts and contents of total HMW-GS in wheat grain. However, this regulatory effect was related to shading intensity.Effect of high temperature and water deficit on HMW-GS and GMP accumulation in wheat grain. The results showed Water deficit and high temperature at both stages obviously affected accumulations of HMW-GS and GMP. High temperature and water deficit at end of spikelet initiation predated the HMW-GS accumulation, especially for the high temperature treatment. Accumulation rate and duration were affected by these stress events. HMW-GS accumulation amount at maturity reduced under high temperature at spikelet initiation while increased when the heat happened at anthesis. However, water deficit at both stages increased HMW-GS accumulation. In addition, acclimation of high temperature at the end of spikelet initiation benefited accumulation of HMW-GS when the plants were subjected to water deficit or high temperature again at anthesis, while water deficit at the end of spikelet promoted the effect of treatments at anthesis. GMP accumulation showed then same pattern to HMW-GS under each treatment.Relationship between HMW-GS content and GMP content. In the present study, significantly linear relationships were observed between GMP contents and individual encoded by different loci(Glu-A1, Glu-B1, Glu-D1) and total HMW-GS contents.

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