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
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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。
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.
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。
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.
引文
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[1]武翠,邵国军,吕文彦,马莲菊,崔鑫福,曹萍,侯秀英.不同发育时期水稻强、弱势粒灌浆速率的遗传分析.中国农业科学,2007,40(6)：1135-1141
[2]姜东,于振文,李永庚,余松烈.高产小麦强势和弱势籽粒淀粉合成相关酶活性的变化.中国农业科学,2002,35(4)：378-383
[3]董明辉,赵步洪,吴翔宙,陈涛,杨建昌.水稻结实期不同粒位籽粒相关内源激素含量和关键酶活性的差异及其与品质的关系,中国农业科学,2008,41(2)：370-380
[4]Xu G-W, Zhang J-H, Lam H-M, Wang Z-Q, Yang J-C. Hormonal changes are related to the poor grain filling in the inferior spikelets of rice cultivated undernon-flooded and mulched condition. Field Crops Research,2007,101:53-61.
[5]梁太波,尹燕枰,蔡瑞国,闫素辉,李文阳,耿庆辉,王平,邬云海,李勇,王振林.不同土壤条件下山农12小麦籽粒HMW-GS积累及GMP粒度分布特征.作物学报,2008,34(12)：2160-2167.
[6]裴雪霞,王姣爱,党建友,王秀斌,张定一.小麦结实粒数、粒重和品质的小穗位和粒位效应.中国农业科学,2008,41(2)：381-390.
[7]梁太波,尹燕枰,蔡瑞国,闫素辉,李文阳,耿庆辉,王平,王振林.大穗型小麦品种强、弱势籽粒淀粉积累和相关酶活性的比较.作物学报,2008,34(1)：150-156.
[8]赵俊晔,于振文.施氮量对小麦强势和弱势籽粒氮素代谢及蛋白质合成的影响.中国农业科学,2005,38(8)：1547-1554.
[9]潘洁,姜东,曹卫星,孙传范.小麦穗籽粒数、单粒重及单粒蛋白质含量的小穗位和粒位效应,2005,31(4)：431-437.
[10]赵全志,吕强,殷春渊,高桐梅,宁慧峰,乔江方,刘辉,陈静蕊.大穗型粳稻籽粒相对充实度的化学调控及其与产量和品质的关系.作物学报,2006,32(10)：1485-1490.
[11]Pirozi M R, B. Margiotta, Lafiandra D, MacRitchie F. Composition of polymeric proteins and bread-making quality of wheat lines with allelic HMW-GS differing in number of cysteines. Journal of Cereal Science,2008,48(1):117-122.
[12]Ohm, J B, Ross A S, Peterson C J, Ong Y L. Relationships of high molecular weight glutenin subunit composition and molecular weight distribution of wheat flour protein with water absorption and color characteristics of noodle dough. Cereal Chemistry,2008,85 (2):123-131
[13]Blechl, A, Lin J, Nguyen S, Chan R, Anderson O D, Dupont F M. Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. Journal of Cereal Science,2007,45 (2):172-183
[14]Wang, Y G, Khan K, Hareland G, Nygard G. Quantitative glutenin composition from gel electrophoresis of glour mill streams and relationship to breadmaking quality. Cereal Chemistry,2006, 83 (3):293
[15]Zhu J-B, Khan K. Characterization of monomeric and glutenin polymeric proteins of hard red spring wheats during grain development by multistacking SDS-PAGE and capillary zone electrophoresis. Cereal Chemistry,1999,76:261-269
[16]Weegel P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and re-polymerisation of wheat glutenin during dough processing:I. Relationships between glutenin macropolymer content and quality parameters. Journal of Cereal Science,1996,23:103-111
[17]孙辉,姚大年,李宝云,刘广田,张树榛.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系.中国粮油学报,1998,13(6)：13-16.
[18]Yue H-W, Jiang D, Dai T-B, Qin X-D, Jing Q, Cao W-X. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45(3):248-256
[19]李硕碧,高翔,单明珠,李必运.小麦高分子量谷蛋白亚基与加工品质.北京：中国农业出版社,2001
[20]周琴,姜东,戴廷波,荆奇,曹卫星.谷氨酰胺对扬麦9号强势和弱势籽粒淀粉及蛋白质积累的调控.麦类作物学报,2005,25(1)：44-49.
[21]吴金芝,李友军,郭天财.小麦品种间不同穗粒位籽粒蛋白质及其组分积累规律的研究.河南科技大学学报,2004,24(1)：1-9.
[22]董明辉,桑大志,王朋,王学明,杨建昌.不同施氮水平下水稻穗上不同部位籽粒的蒸煮与营养品质变化.中国水稻科学,2006,20(4)：389-395.
[1]Pirozi M R, B. Margiotta, Lafiandra D, MacRitchie F. Composition of polymeric proteins and bread-making quality of wheat lines with allelic HMW-GS differing in number of cysteines. Journal of Cereal Science,2008,48(1):117-122.
[2]Ohm, J B, Ross A S, Peterson C J, Ong Y L. Relationships of high molecular weight glutenin subunit composition and molecular weight distribution of wheat flour protein with water absorption and color characteristics of noodle dough. Cereal Chemistry,2008,85 (2):123-131.
[3]Zhu J B, Khan K. Characterization of monomeric and glutenin polymeric proteins of hard red spring wheats during grain development by multistacking SDS-PAGE and capillary zone electrophoresis. Cereal Chemistry,1999,76:261-269.
[4]杜金哲,胡尚连,李文雄,刘锦红.不同品质类型春小麦HMW-GS形成时间和积累强度及与品质的关系.作物学报,2003,29(1)：111-118.
[5]Blechl, A, Lin J, Nguyen S, Chan R, Anderson O D, Dupont F M. Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. Journal of Cereal Science,2007,45 (2):172-183
[6]Wang, Y G, Khan K, Hareland G, Nygard G. Quantitative glutenin composition from gel electrophoresis of flour mill streams and relationship to breadmaking quality. Cereal Chemistry, 2006,83 (3):293.
[7]邓志英,田纪春,张永祥,王延训.冬小麦高、低分子量麦谷蛋白亚基形成时间和积累强度及其与沉降值的关系.作物学报,2005,31(3)：308-315.
[8]Vawser M, Cornish G B. Over-expression of HMW glutenin subunit Glu-B1 7x in hexaploid wheat varieties (Triticum aestivum). Australian Journal of Agricultural Research 2004,55 (5):577-588.
[9]Leon E, Marin S, Gimenez M J, Piston F, M Rodriguez-Quijano, Shewry P R, and Barro F. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the lAx1,1Dx5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science,2009,49 (1):148-156.
[10]DuPont F M, Hurkman W J, Vensel W H, Chan R, Lopez R, Tanaka, C K, Altenbach S B. Differential accumulation of sulfur-rich and sulfur-poor wheat flour proteins is affected by temperature and mineral nutrition during grain development. Journal of Cereal Science,2006,44, 101-112.
[11]Yue H W, Jiang D, Dai T B, Qin X D, Jing Q, Cao W X. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45(3):248-256.
[12]Zhu J B, Khan K. Quantitative variation of HMW glutenin subunits from hard red spring wheats grown in different environments. Cereal Chemistry,2002,79:783-786.
[13]岳鸿伟,谭维娜,姜东,戴廷波,荆奇,曹卫星.花后干旱和渍水对小麦籽粒HMW-GS及GMP含量的影响.作物学报,2007,33(11)：1845-1849.
[14]李永庚,于振文,梁晓芳,赵俊晔,邱希宾.小麦产量和品质对灌浆期不同阶段低光照强度的响应.植物生态学报,2005,29(5)：807-813.
[15]王艳君,姜彤,许崇育.长江流域蒸发皿蒸发量及影响因素变化趋势.自然资源学报,2005,20(6)：864-870
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[17]Weegel P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and re-polymerisation of wheat glutenin during dough processing:I. Relationships between glutenin macropolymer content and quality parameters. Journal of Cereal Science,1996,23:103-111
[18]孙辉,姚大年,李宝云,刘广田,张树榛.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系.中国粮油学报,1998,13(6)：13-16
[19]李硕碧,高翔,单明珠,李必运.小麦高分子量谷蛋白亚基与加工品质.北京：中国农业出版社,2001
[20]DuPont F M, Chan R, Lopez R. Molar fractions of high-molecular-weight glutenin subunits are stable when wheat is grown under various mineral nutrition and temperature regimens. Journal of Cereal Science,2007,45(2):134-139.
[21]王东,于振文.不同施氮量下子粒灌浆不同阶段遮光对小麦氮素积累和转移的影响.植物营养与肥料学报,2008,14(4)：615-622.
[22]牟会荣,姜东,戴廷波,荆奇,曹卫星.遮荫对小麦旗叶光合及叶绿素荧光特性的影响.中国农业科学,2008,41(2)；599-606
[23]Caroline V M, Roger C, Anderson, Diane L B. Influence of shading on the growth and leaf photosynthesis of the invasive non-indigenous plant garlic mustard [Alliaria petiolata (M. Bieb) Cavara and Grande] grown under simulated late-winter to mid-spring conditions. Journal of the Torrey Botanical Society,2005,132(1):1-10.
[24]Don C, Mann G, Bekes F, Hamer R J. HMW-GS affect the properties of glutenin particles in GMP and thus flour quality. Journal of Cereal Science,2006,44(2):127-136.
[1]Weegels P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and Re-polymerisation of Wheat Glutenin During Dough Processing. I. Relationships between Glutenin Macropolymer Content and Quality Parameters. Journal of Cereal Science,1996,23 (2):103-111.
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[9]Vawser M, Cornish G B. Over-expression of HMW glutenin subunit Glu-B1 7x in hexaploid wheat varieties (Triticum aestivum). Australian Journal of Agricultural Research,2004,55 (5):577-588.
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[11]Leon E, Marin S, Gimenez M J, Piston F, Rodriguez-Quijano M, Shewry P R, Barro F. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the 1Ax1,1Dx5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science,2009,49 (1):148-156.
[12]Darlington H, Fido R, Tatham A S, Jones H, Salmon S E, Shewry P R. Milling and baking properties of field grown wheat expressing HMW subunit transgenes. Journal of Cereal Science,2003, 38 (3):301-306.
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[14]Yue H, Jiang D, Dai T, Qin X, Jing Q, Cao W. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45 (3):248-256.
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[20]D. Jiang H Y, B. Wollenweber, W. Tan, H. Mu, Y. Bo, T. Dai, Q. Jing, W. Cao,. Effects of post-anthesis drought and waterlogging on accumulation of high-molecular-weight glutenin subunits and glutenin macropolymers content in eheat grain. Journal of Agronomy and Crop Science,2009,195 (2): 89-97.
[21]Altenbach S B, Kothari K M, Lieu D. Environmental conditions during wheat grain development alter temporal regulation of major gluten protein genes. Cereal Chemistry,2002,79 (2):279-285.
[22]Dupont F M, Hurkman W J, Vensel W H, Tanaka C, Kothari K M, Chung O K, Altenbach S B. Protein accumulation and composition in wheat grains:Effects of mineral nutrients and high temperature. European Journal of Agronomy,2006,25 (2):96-107.
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[1]陈佩度,周波,齐莉莉,刘大钧.用分子原位杂交(GISH)鉴定小麦-簇毛麦双倍体、附加系和易位系.遗传学报,1995,22(5)：380-386
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[8]Zhu J B, Khan K. Quantitative variation of HMW glutenin subunits from hard red spring wheats grown in different environments. Cereal Chemistry,2002,79:783-786.
[9]邓志英,田纪春,张永祥,王延训.冬小麦高、低分子量麦谷蛋白亚基形成时间和积累强度及其与沉降值的关系.作物学报,2005,31(3)：308-315
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[12]杜金哲,胡尚连,李文雄,刘锦红.不同品质类型春小麦HMW-GS形成时间和积累强度及与品质的关系.作物学报,2003,29(1)：111-118
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[1]武翠,邵国军,吕文彦,马莲菊,崔鑫福,曹萍,侯秀英.不同发育时期水稻强、弱势粒灌浆速率的遗传分析.中国农业科学,2007,40(6)：1135-1141
[2]姜东,于振文,李永庚,余松烈.高产小麦强势和弱势籽粒淀粉合成相关酶活性的变化.中国农业科学,2002,35(4)：378-383
[3]董明辉,赵步洪,吴翔宙,陈涛,杨建昌.水稻结实期不同粒位籽粒相关内源激素含量和关键酶活性的差异及其与品质的关系,中国农业科学,2008,41(2)：370-380
[4]Xu G-W, Zhang J-H, Lam H-M, Wang Z-Q, Yang J-C. Hormonal changes are related to the poor grain filling in the inferior spikelets of rice cultivated undernon-flooded and mulched condition. Field Crops Research,2007,101:53-61.
[5]梁太波,尹燕枰,蔡瑞国,闫素辉,李文阳,耿庆辉,王平,邬云海,李勇,王振林.不同土壤条件下山农12小麦籽粒HMW-GS积累及GMP粒度分布特征.作物学报,2008,34(12)：2160-2167.
[6]裴雪霞,王姣爱,党建友,王秀斌,张定一.小麦结实粒数、粒重和品质的小穗位和粒位效应.中国农业科学,2008,41(2)：381-390.
[7]梁太波,尹燕枰,蔡瑞国,闫素辉,李文阳,耿庆辉,王平,王振林.大穗型小麦品种强、弱势籽粒淀粉积累和相关酶活性的比较.作物学报,2008,34(1)：150-156.
[8]赵俊晔,于振文.施氮量对小麦强势和弱势籽粒氮素代谢及蛋白质合成的影响.中国农业科学,2005,38(8)：1547-1554.
[9]潘洁,姜东,曹卫星,孙传范.小麦穗籽粒数、单粒重及单粒蛋白质含量的小穗位和粒位效应,2005,31(4)：431-437.
[10]赵全志,吕强,殷春渊,高桐梅,宁慧峰,乔江方,刘辉,陈静蕊.大穗型粳稻籽粒相对充实度的化学调控及其与产量和品质的关系.作物学报,2006,32(10)：1485-1490.
[11]Pirozi M R, B. Margiotta, Lafiandra D, MacRitchie F. Composition of polymeric proteins and bread-making quality of wheat lines with allelic HMW-GS differing in number of cysteines. Journal of Cereal Science,2008,48(1):117-122.
[12]Ohm, J B, Ross A S, Peterson C J, Ong Y L. Relationships of high molecular weight glutenin subunit composition and molecular weight distribution of wheat flour protein with water absorption and color characteristics of noodle dough. Cereal Chemistry,2008,85 (2):123-131
[13]Blechl, A, Lin J, Nguyen S, Chan R, Anderson O D, Dupont F M. Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. Journal of Cereal Science,2007,45 (2):172-183
[14]Wang, Y G, Khan K, Hareland G, Nygard G. Quantitative glutenin composition from gel electrophoresis of glour mill streams and relationship to breadmaking quality. Cereal Chemistry,2006, 83 (3):293
[15]Zhu J-B, Khan K. Characterization of monomeric and glutenin polymeric proteins of hard red spring wheats during grain development by multistacking SDS-PAGE and capillary zone electrophoresis. Cereal Chemistry,1999,76:261-269
[16]Weegel P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and re-polymerisation of wheat glutenin during dough processing:I. Relationships between glutenin macropolymer content and quality parameters. Journal of Cereal Science,1996,23:103-111
[17]孙辉,姚大年,李宝云,刘广田,张树榛.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系.中国粮油学报,1998,13(6)：13-16.
[18]Yue H-W, Jiang D, Dai T-B, Qin X-D, Jing Q, Cao W-X. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45(3):248-256
[19]李硕碧,高翔,单明珠,李必运.小麦高分子量谷蛋白亚基与加工品质.北京：中国农业出版社,2001
[20]周琴,姜东,戴廷波,荆奇,曹卫星.谷氨酰胺对扬麦9号强势和弱势籽粒淀粉及蛋白质积累的调控.麦类作物学报,2005,25(1)：44-49.
[21]吴金芝,李友军,郭天财.小麦品种间不同穗粒位籽粒蛋白质及其组分积累规律的研究.河南科技大学学报,2004,24(1)：1-9.
[22]董明辉,桑大志,王朋,王学明,杨建昌.不同施氮水平下水稻穗上不同部位籽粒的蒸煮与营养品质变化.中国水稻科学,2006,20(4)：389-395.
[1]Pirozi M R, B. Margiotta, Lafiandra D, MacRitchie F. Composition of polymeric proteins and bread-making quality of wheat lines with allelic HMW-GS differing in number of cysteines. Journal of Cereal Science,2008,48(1):117-122.
[2]Ohm, J B, Ross A S, Peterson C J, Ong Y L. Relationships of high molecular weight glutenin subunit composition and molecular weight distribution of wheat flour protein with water absorption and color characteristics of noodle dough. Cereal Chemistry,2008,85 (2):123-131.
[3]Zhu J B, Khan K. Characterization of monomeric and glutenin polymeric proteins of hard red spring wheats during grain development by multistacking SDS-PAGE and capillary zone electrophoresis. Cereal Chemistry,1999,76:261-269.
[4]杜金哲,胡尚连,李文雄,刘锦红.不同品质类型春小麦HMW-GS形成时间和积累强度及与品质的关系.作物学报,2003,29(1)：111-118.
[5]Blechl, A, Lin J, Nguyen S, Chan R, Anderson O D, Dupont F M. Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. Journal of Cereal Science,2007,45 (2):172-183
[6]Wang, Y G, Khan K, Hareland G, Nygard G. Quantitative glutenin composition from gel electrophoresis of flour mill streams and relationship to breadmaking quality. Cereal Chemistry, 2006,83 (3):293.
[7]邓志英,田纪春,张永祥,王延训.冬小麦高、低分子量麦谷蛋白亚基形成时间和积累强度及其与沉降值的关系.作物学报,2005,31(3)：308-315.
[8]Vawser M, Cornish G B. Over-expression of HMW glutenin subunit Glu-B1 7x in hexaploid wheat varieties (Triticum aestivum). Australian Journal of Agricultural Research 2004,55 (5):577-588.
[9]Leon E, Marin S, Gimenez M J, Piston F, M Rodriguez-Quijano, Shewry P R, and Barro F. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the lAx1,1Dx5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science,2009,49 (1):148-156.
[10]DuPont F M, Hurkman W J, Vensel W H, Chan R, Lopez R, Tanaka, C K, Altenbach S B. Differential accumulation of sulfur-rich and sulfur-poor wheat flour proteins is affected by temperature and mineral nutrition during grain development. Journal of Cereal Science,2006,44, 101-112.
[11]Yue H W, Jiang D, Dai T B, Qin X D, Jing Q, Cao W X. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45(3):248-256.
[12]Zhu J B, Khan K. Quantitative variation of HMW glutenin subunits from hard red spring wheats grown in different environments. Cereal Chemistry,2002,79:783-786.
[13]岳鸿伟,谭维娜,姜东,戴廷波,荆奇,曹卫星.花后干旱和渍水对小麦籽粒HMW-GS及GMP含量的影响.作物学报,2007,33(11)：1845-1849.
[14]李永庚,于振文,梁晓芳,赵俊晔,邱希宾.小麦产量和品质对灌浆期不同阶段低光照强度的响应.植物生态学报,2005,29(5)：807-813.
[15]王艳君,姜彤,许崇育.长江流域蒸发皿蒸发量及影响因素变化趋势.自然资源学报,2005,20(6)：864-870
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[17]Weegel P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and re-polymerisation of wheat glutenin during dough processing:I. Relationships between glutenin macropolymer content and quality parameters. Journal of Cereal Science,1996,23:103-111
[18]孙辉,姚大年,李宝云,刘广田,张树榛.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系.中国粮油学报,1998,13(6)：13-16
[19]李硕碧,高翔,单明珠,李必运.小麦高分子量谷蛋白亚基与加工品质.北京：中国农业出版社,2001
[20]DuPont F M, Chan R, Lopez R. Molar fractions of high-molecular-weight glutenin subunits are stable when wheat is grown under various mineral nutrition and temperature regimens. Journal of Cereal Science,2007,45(2):134-139.
[21]王东,于振文.不同施氮量下子粒灌浆不同阶段遮光对小麦氮素积累和转移的影响.植物营养与肥料学报,2008,14(4)：615-622.
[22]牟会荣,姜东,戴廷波,荆奇,曹卫星.遮荫对小麦旗叶光合及叶绿素荧光特性的影响.中国农业科学,2008,41(2)；599-606
[23]Caroline V M, Roger C, Anderson, Diane L B. Influence of shading on the growth and leaf photosynthesis of the invasive non-indigenous plant garlic mustard [Alliaria petiolata (M. Bieb) Cavara and Grande] grown under simulated late-winter to mid-spring conditions. Journal of the Torrey Botanical Society,2005,132(1):1-10.
[24]Don C, Mann G, Bekes F, Hamer R J. HMW-GS affect the properties of glutenin particles in GMP and thus flour quality. Journal of Cereal Science,2006,44(2):127-136.
[1]Weegels P L, van de Pijpekamp A M, Graveland A, Hamer R J, Schofield J D. Depolymerisation and Re-polymerisation of Wheat Glutenin During Dough Processing. I. Relationships between Glutenin Macropolymer Content and Quality Parameters. Journal of Cereal Science,1996,23 (2):103-111.
[2]Labuschagne M T, Koen E, Dessalegn T. Use of Size-Exclusion High-Performance Liquid Chromatography for Wheat Quality Prediction in Ethiopia. Cereal Chemistry,2004,81 (4):533-537.
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[5]Pirozi M R, Margiotta B, Lafiandra D, MacRitchie F. Composition of polymeric proteins and bread-making quality of wheat lines with allelic HMW-GS differing in number of cysteines Journal of Cereal Science,2008,48 (1):117-122.
[6]Wang Y G, Khan K, Hareland G, Nygard G Quantitative Glutenin Composition from Gel Electrophoresis of Flour Mill Streams and Relationship to Breadmaking Quality. Cereal Chemistry,2006, 83 (3):293.
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[9]Vawser M, Cornish G B. Over-expression of HMW glutenin subunit Glu-B1 7x in hexaploid wheat varieties (Triticum aestivum). Australian Journal of Agricultural Research,2004,55 (5):577-588.
[10]Zhu J B, Liu G T, Zhang S Z, Sun H. High and low molecular subunits of glutenin and their relationships with wheat quality. Scientia Agrcultural Sinica in Chinese 1996,29:34-39.
[11]Leon E, Marin S, Gimenez M J, Piston F, Rodriguez-Quijano M, Shewry P R, Barro F. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the 1Ax1,1Dx5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science,2009,49 (1):148-156.
[12]Darlington H, Fido R, Tatham A S, Jones H, Salmon S E, Shewry P R. Milling and baking properties of field grown wheat expressing HMW subunit transgenes. Journal of Cereal Science,2003, 38 (3):301-306.
[13]Popineau Y, Deshayes G, Lefebvre J, Fido R, Tatham A S, Shewry P R. Prolamin aggregation, gluten viscoelasticity, and mixing properties of transgenic wheat lines expressing 1Ax and 1Dx high molecular weight glutenin subunit transgenes. Journal of Agricultural and Food Chemistry,2001,49 (1): 395-401.
[14]Yue H, Jiang D, Dai T, Qin X, Jing Q, Cao W. Effect of nitrogen application rate on content of glutenin macropolymer and high molecular weight glutenin subunits in grains of two winter wheat cultivars. Journal of Cereal Science,2007,45 (3):248-256.
[15]Z. Deng J T, L. Zhao, Y. Zhang, C. Sun,. High temperature-induced changes in high molecular weight glutenin subunits of Chinese winter wheat and its influences on the texture of Chinese noodles. Journal of Agronomy and Crop Science,2008,194 (4):262-269.
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