水氮互作对小麦谷蛋白亚基以及谷蛋白大聚合体粒度分布的调控
|
摘要
小麦籽粒的高分子量谷蛋白亚基(HMW-GS)和低分子量谷蛋白亚基(LMW-GS)的种类、含量决定着谷蛋白大聚合体(GMP)的粒度,GMP的粒度分布是决定小麦烘焙品质的重要指标。本实验选用了强筋小麦藁城8901(GC8901)、中筋小麦泰山23(TS23)和弱筋小麦山农1391(SN1391)三个小麦品种,探讨了水氮互作栽培措施对小麦籽粒HMW-GS的含量和籽粒GMP粒度分布的影响。在灌溉条件下,随着氮肥量(0,120,240,360 kg hm~(-2))的增加,三个小麦品种均表现为籽粒高、低分子量谷蛋白亚基的含量增加,GMP大颗粒的体积分布变大,产量提高,烘焙品质变优。旱作条件下增施氮肥,提高了三小麦品种籽粒的蛋白质含量,强筋小麦GC8901籽粒的H-、LMW-GS相对含量降低,H/L值升高,大颗粒GMP所占的表面积分数,体积分数降低;而中筋小麦TS23和弱筋小麦SN1391表现出相反的趋势,两者产量显著降低。同时,相互连锁的同一亚基对的相对表达量对外界环境的变化表现出较好的稳定性,且其表达具有协同性,但单位面粉亚基的含量受栽培措施影响显著。增施氮肥能明显改善小麦的谷蛋白品质,主要表现为两条调控途径:一方面,增加GMP大颗粒的数目,另一方面,GMP大颗粒的数目没有明显增加,但其体积和表面积分布增加。水氮互作效应对三品种的产量和品质性状的影响不显著,说明基因型和环境之间的互作是一个复杂的网络,在小麦生产过程中,农民应首先考虑水分对产量和品质的影响,然后通过施肥等栽培措施来进一步改善作物的品质。
三小麦品种的全长序列分析
高分子量谷蛋白亚基的前30个氨基酸是信号肽,决定着高分子量谷蛋白亚基的空间表达。1Ax可读框内仅含有4个半胱氨酸残基,但形成20个潜在的α螺旋结构,1Dx2,1Bx7可读框内含有4个半胱氨酸残基,形成9个潜在的α螺旋结构。1Dx5亚基含有5个半胱氨酸残基可以形成潜在的9个α螺旋。1Bx14可读框内仅含有2个半胱氨酸残基,形成11个潜在的α螺旋结。1Dy10,1By8,1By15,1Dy12可读框内均含有7个半胱氨酸残基,分别形成9,11,11,11个潜在的α螺旋结构。虽然高分子量谷蛋白亚基在进化过程中出现了不同程度的碱基序列的变异,但是其序列的同源性结构决定了它们的蛋白序列和空间结构是类似的,因而在面粉中行使的的功能也是类似的。因此,对形成HMW-GS分子结构有贡献的三个重要氨基酸残基脯氨酸、甘氨酸和谷氨酰胺的含量也是优质面团粘弹性形成的重要参考标准。
亚基含量的HP-HPLC分析
灌溉条件下,强筋小麦GC8901和中筋小麦TS23籽粒的HMW-GS含量和H/L值随着氮肥施用量的增加而增加,LMW-GS的含量在施氮量为120 kg ha~(-1)时达到最大,弱筋小麦品种SN1391则表现出相反的趋势。旱作栽培模式下,中、高氮肥使用量(240 kg N ha~(-1),360 kg N ha~(-1))均提高了三小麦品种的H-、LMW-GS含量,SN1391提高量尤为显著,不施氮和低氮施用量(120 kg ha~(-1))显著降低了三小麦品种的H-、LMW-GS含量。
水氮互作条件下,三小麦品种的1Dx2和1Dx5亚基的相对含量最高,约占总HMW-GS含量的42%和38%。1Bx7和1Dy12亚基相对含量次之,约占31%和26%,1Ax1和1By8亚基的相对含量最小,仅占14%和12%。相互连锁的亚基对的相对含量在不同氮水平上变化不显著,但单个亚基的相对含量对水旱处理的反应不一。1Dy10亚基、1By15亚基在不同氮肥水平条件下占HMW-GS的比值是稳定的,之间无显著性差异;在非灌水条件下,7亚基、8亚基、1亚基的比值在GC8901中是稳定的,氮肥处理之间无显著性差异。与水地相比较,旱作处理使GC8901的1Ax1亚基的相对含量增加了10%,1Bx7亚基的相对含量增加了26.5%,1By8增加了85.7%,然而1Dx5和1Dy10亚基的相对含量保持不变。TS23旱作栽培条件下,1Ax亚基的相对含量下降了25%,1Bx7增加了12.2%而1By8增加了38.7%。SN1391各个亚基的相对含量在水肥料处理间则表现出相对稳定的趋势。亚基对之间相比较,1Bx7+1By8亚基对对环境反应更敏感,在中筋和弱筋小麦籽粒中,亚基对1Dx2+1Dy12往往表达量较高。相互连锁的亚基对的相对表达量表现持一定的互补和协同效应。水、氮栽培措施主要通过增加单位籽粒中的亚基含量来改善小麦的谷蛋白质量。
小麦籽粒GMP含量变化
3个小麦品种的GMP含量的大小依次顺序为:GC8901>TS23>SN1391。在灌水条件下,GMP含量都随着氮肥水平的增加而增加。在非灌水条件下,强筋小麦GC8901GMP含量随氮肥水平的增加呈抛物线状,在施氮量为240 kg ha~(-1)时GMP含量最高。与灌溉相比较,TS23和SN1391的GMP含量在不施氮时最高,分别增加了6-15.4%和30.9-82%,随着施氮量的增加GMP含量下降,中、高氮肥(240 kg N ha~(-1)、360 kg N ha~(-1))处理间无显著差异。在灌水条件下,增施氮肥有利于GMP在小麦籽粒中的积累,旱作栽培模式有利于中筋、和弱筋小麦籽粒的GMP的积累,但不同氮肥水平之间差异不显著。
小麦籽粒GMP的粒度分布
小麦成熟期的GMP粒度分布趋势相似,粒径范围为0.375~256.9μm。小麦GMP颗粒的体积分布为双峰曲线,第一个峰值出现在3.5~4.5μm之间,第二个峰值出现在116~140μm之间,不同品种间及处理间差异较大。GMP颗粒的表面积分布亦呈双峰曲线,数目分布为单峰曲线。小麦GMP数目主要由<10μm颗粒组成(约占99.8%以上),体积分布的47.57%~69%集中在12-100μm,而GMP表面积分布< 12μm,12-110μm和> 110μm百分数分别占72.3-89.4%, 9.5-21.5%和1.1-10.1%。
灌水条件下,强筋小麦GC8901籽粒GMP的12-110μm、< 110μm的表面积分布比例随着施氮量的增加先下降后显著上升。与旱作相比较,强筋小麦GC8901 < 12μm的GMP表面积分布比例显著下降。弱筋小麦SN1391表现出与GC8901完全相反趋势。灌溉条件下,中筋小麦TS23 < 12μm随着施氮量的增加而降低,12-110μm和> 110μm的GMP表面积分布比例升高,而旱作条件下,TS23不施氮水平的GMP表面积分布比例最高。三小麦品种D4.3和D3.2的变化趋势与GMP > 110μm颗粒的表面积分布趋势基本一致。旱作、低氮栽培条件能显著提高强筋小麦和弱筋小麦籽粒的体积分布比例。与旱作相比较,灌溉使TS23和GC8901 < 4.5μm的GMP颗粒的数目分布增加,4.5-12μm和> 12μm的数目分布降低,而增施氮肥使>12μm的GMP颗粒的数目增加。SN1391 <4.5μm的GMP颗粒的数目比例随施氮量而增加,4.5-12μm颗粒的数目比例降低, > 12μm颗粒的数目比例保持不变。
增施氮肥有利于GC8901籽粒HMW-GS的积聚和小粒径GMP的成长为大粒径GMP,调节了弱筋小麦GMP中等粒径颗粒和小粒径GMP颗粒数目的分布,从而形成更大的表面积分布,而不是大粒径的GMP颗粒,水分亏缺可以加速这两个调节过程。
小麦籽粒GMP的电镜观测
通过SEM电镜照片可以看出,GC8901籽粒的GMP颗粒的体积明显大于SN1391和TS23的GMP颗粒的体积,且SN1391GMP多为粒径<1.5μm小颗粒,这与激光粒度分析仪的结果相一致。这应当是由于GC8901(1、5+10、7+8)比SN1391(2+12、14+15)的亚基种类多出了1亚基且所含亚基种类的分子量较大,所以形成的GMP体积也较大的原因导致的。而从GC8901籽粒中的GMP在不同蔗糖浓度梯度下离心、分层后的所得的SDS-PAGE电泳图片看出,各层GMP所含高低分子量谷蛋白亚基种类没有差别,但无法确定其亚基的比例。可见,GMP大颗粒是有许多GMP小颗粒包裹形成,而TEM和SEM图像也表明大的GMP颗粒是由小GMP颗粒聚合而成的,较小的GMP颗粒是由H-和LMW-GS以不同比例组合而成的。
水氮互作对小麦产量和品质的影响
无论是在灌溉还是雨养条件下,增施氮肥均增加了籽粒的蛋白质含量和籽粒的千粒重。在高氮水平下(360kg ha~(-1)),相比灌溉条件,旱作栽培模式下的GC8901的蛋白质含量下降4.8-22.9%,而TS23和S1391则提高了45.6%和36.7%。灌水条件下三个小麦品种的平均产量为6.9-9.9 mt ha~(-1),而旱作模式下三个小麦品种的产量下降了2.0-38.2%,当施氮水平高于240kg N ha~(-1),产量增加不显著。DDT是反映面团的烘烤品质的重要指标,三小麦品种的DDT和籽粒GMP含量变化表现出的趋势,表明DDT和GMP含量之间密切的关系。同时,D3.2和D4.3与谷蛋白含量、HMW-GS含量、LMW-GS含量、DDT、H/L值、GMP含量和淀粉与蛋白质的比值均显著正相关,与单粒重呈极显著负相关,与籽粒淀粉含量相关不显著,但是在旱作栽培条件下D4.3与这些性状没有表现出明显的相关性。因此,H-、LMW-GS的含量和GMP的粒度分布决着面团烘焙品质的好坏。水肥充足的条件下小麦倾向于获得较高的生物量和产量,而在水分亏缺条件下则倾向于提高单粒重和蛋白质含量,以储存较多的能量。
The composition of high molecular weight glutenin subunit (HMW-GS) and low molecular weight glutenin subunit (LMW-GS) in wheat kernel (Triticum aestivum L.) affects the size of glutenin macropolymer (GMP), which is considered the important flour quality traits in wheat. Three wheat cultivars with different end-use quality were used to study the effects of water-nitrogen management (W×N) on relative HMW-GS and GMP contents. Under irrigation, increased N levels promoted the accumulation of H- and LMW-GS, GMP content and the proportion of the larger particle of GMP, with higher yield and better end-use quality. Under rainfed conditions, increased N fertilizer also increased protein content in the three cultivars. As opposed to the Gaocheng8901 (GC8901), the Shannong1391 (SN1391) and Taishan23 (TS23) got higher H- and LMW-GS contents and larger GMP particles, yields of both reduced. Irrigation increased the yield markedly, the N application significantly affected the glutenin, but the W×N interaction was not significant. On one hand, increased N levels improved the dough development time (DDT) by increasing the numbers of larger GMP particles and enhancing volume and surface area percentages on the other. It was suggested that the interaction was a complicated network and some individual effect might be important. When managing for wheat yield and quality, growers should first consider the water factor, then the N fertilizer factor to improve the end-use quality of wheat.
Full-length HMW-GS Sequence Analysis of tthree wheat cultivars
The HMW-GS pairs 1Dx5+1Dy10, 1Bx14+1By15 are recognized to positively correlate with bread-making quality. Both the reading frames (ORFs) of 1Ax and of 1Bx7 contain 4 cysteines, but the ORF of 1Ax forms 20 potentialα-helix structures while 1Bx7 forms 9 potentialα-helix structures. The ORF of 1Dx which forms 9 potentialα-helix structures contains 5 cysteines; the ORF of 1Bx14 which forms 11 potentialα-helix structures contains only 2 cysteines. The ORFs of 1Dy10, 1By8, 1By15 and 1Dy12 all contain 7 cysteines and form respectively 9, 11, 11 and 11 potentialα-helix structures. The DNA sequences showed they are homologous and would had the similar structure and function in wheat kernel. So the cysteine residue which plays important roles in forming the intra- and inter molecular structures of HMW-GS could not be used as the sole evaluation criterion for the merits of subunits. The proline, glycine and glutamine play important roles in forming advanced structure of glutenin, so the three amino acid residues would be the important reference standard for forming the visco-elasticity of quality dough. Meanwhile, the three amino acids have taken a large proportion in x- style HMW-GS. It means that x- style subunits contribute more for dough quality.
Differences in Experimental Factors
Analysis of variance for the contents of subunits, GMP content, the D4.3, D3.2, the single kernel weight, yield, grain protein content and the DDT made it possible to identify the sources of variation. Nitrogen (N) main effect and water×N (W×N) interaction failed to influence those traits except single kernel weight and grain protein content. The Student Newman and Keuls test (p≤0.05) showed that subunits in 1Bx chromosome, HMW-GS content and yield were significantly affected by water treatment. Although the N main effect as well as the W×N were not significant through three wheat cultivars, some individual effect was important to determine the quality and yield. It was suggested the interaction was a complicated network. Variations due to years and in interactions between year and nitrogen (Yr×N), year and water (Yr×W) treatment were not significant (p≤0.05). Since year was not a significant factor in the total reaction, some data from both years were averaged.
RP-HPLC Analysis for Glutenin
Considering the three cultivars, the 1Dx2 and 1Dx5 subunits accumulated to the highest level about 42% and 38% of total high molecular weight glutenin subunit (HMW-GS) content. The 1Bx7 and 1Dy12 subunits were present in a smaller amount, 31% and 26%, and the 1Ax1 and 1By8 subunits were the least abundant and nearly equal, 14% and 12%. The proportions of linked subunit pairs were nearly equal among different N levels, but the proportions of single subunits showed various under the two water treatments. As compared with irrigated regimen, rainfed regimen increased the mean proportion of 1Ax1 subunit by 10%, decreased 1Bx7 by 26.5%, increased 1By8 by 85.7%, whereas, the proportion of 1Dx5 and 1Dy10 remained stable in GC8901. The mean proportion of 1Ax1 in TS23 under irrigated regimen increased by 25%, 1Bx7 increased by 12.2% and 1By8 decreased by 38.7% compared with of under rain-fed regimen. Water treatment showed no significant effects on the subunit pairs in SN1391 with increased N application. Take all the subunit pairs into account, 1Bx7+1By8 seemed more sensitive to the environment and 1Dx2+1Dy12 always high expressed in middle and weak gluten wheat. We observed that under irrigation, both in GC8901 and TS23, the content of HMW-GS and H/L value increased with N increased, and the LMW-GS contents reached the maximums at 120 kg N ha~(-1) (N1). However, the trend was opposite in SN1391 under the same condition. Under rainfed regimen, the contents of H- and LMW-GS increased in three wheat cultivars with higher N application (N2, N3), especially in SN1391. However, the H- and LMW-GS contents of three cultivars decreased markedly under rainfed condition with low N application (N1).
These results indicated that the ratios of the linked subunit pair’s proportions to the mean proportion of different N levels were nearly identical. The two linked subunit pairs had certain complementary effects, the final proportions showed no significant relationship with W×N interaction and mainly determined by genotypes. Whereas, a significantly correlated was shown among single kernel weight, protein content, GMP content (r = -0.829**; 0.563*; 0.803**) and H/L. It was indicated that the grain kernel production interacts with H- and LMW-GS contents. Rainfed condition could increase the contents of H- and LMW-GS and H/L value in three wheat cultivars.
Content of GMP Particles
The contents of GMP in three wheat cultivars were ordered as follows: GC8901 > TS23 > SN1391. The contents of GMP all increased as N rates increased under irrigated regimen. But under rainfed regimen, the content of GMP in GC8901 showed difference with the N level, and reached the maximum at 240 kg N ha~(-1) (N2); the GMP contents of TS23 and SN1391 both were markedly increased by 6-15.4% and 30.9-82% with no N applied (N0), and tended to decline as increased N application, while the differences of contents between N application of 240 kg N ha~(-1) (N2) and 360 kg N ha~(-1) (N3) were not significant. The results indicated that increasing N level could promote the accumulation of GMP in three wheat cultivars under irrigated regimen. Rainfed condition significantly promoted the accumulation of GMP in middle and weak gluten wheat, but the content remained stable with N rates.
GMP Particle Distribution
The surface area percent of GMP particles at < 12μm, 12-110μm and > 110μm in three cultivars took up 72.3-89.4%, 9.5-21.5% and 1.1-10.1%. Under irrigation, the surface area percentages of GMP particles at 12-110μm and < 110μm in GC8901 decreased first and then markedly increased with N increased, whereas that of < 12μm decreased markedly when compared with those under rainfed regimen. However, those in SN1391 showed a character of opposite extremes. Those at < 12μm in TS23 decreased with N increased, and those of 12-110μm and > 110μm increased under irrigation regimen. While under rainfed regimen, the GMP surface area percentage of TS23 reached the maximum value with no N applied (N0). The irrigation was beneficial to increasing the surface area percent of GMP particles as increased N application, and sight water stress could markedly promote those in weak and middle gluten wheat.
The data in Table 4 indicated that D4.3 and D3.2 were completely in line with the surface area percentages of GMP particles at > 110μm. Dispersed GMP of three cultivars under W×N interaction displayed that rainfed regimen and lower N level could promote the volume of particles in middle and weak gluten wheat. Strong gluten wheat was more adapted to plenteous irrigation and higher N application. And interestingly, the D4.3 under rain-fed regimen of TS23 and SN1391 were bigger than that of in GC8901, but the content of GMP in TS23 and SN1391was less.
Irrigated and rainfed conditions had different influence on percent number of GMP particles in three wheat cultivars. Under irrigation, the number percentage of GC8901 and TS23 at < 4.5μm increased, while those of 4.5-12μm and > 12μm decreased when compared with those under rainfed condition. Increased N level contributed to increase the GMP particles numbers of GC8901 and TS23 at >12μm. The increased N level increased number percentage of GMP particles in SN1391 at <4.5μm, decreased that of 4.5-12μm, whereas that of > 12μm remained the same. The GMP particles number percentage responses of SN1391 to water treatment were similar to those to N application. The number distribution revealed that increased N level could increase the numbers large GMP particles in strong and middle gluten wheat. While in weak gluten wheat, N effect and W×N interaction showed no influence on the numbers of larger GMP particles.
Observations of GMP Particles
To further analyze the effects of W×N interaction on glutenin particle, GMP dispersions of three wheat cultivars were examined for particulate structures obtained. The SEM figures showed that large GMP particle was made from the polymerization of small GMP particles, the smaller GMP particles were made of H- and LMW-GS in various proportions. We also estimated that GC8901 got a highest H/L ratio ranged from 0.36-0.47 and the highest D4.3, TS23 got a middle H/L ratio ranged from 0.26-0.33 and smaller D4.3, SN1391 got the least H/L ratio ranged from 0.20-0.25 and least D4.3. It indicated that D4.3 was mainly determined by genotypes, but there was no evidence to confirm their proportions of H- and LMW-GS in this paper. The D4.3 under irrigated regimen showed strong relation to protein content, GMP content, single kernel weight and H/L value (r=0.822**; 0.755**; -0.806**; 0.698*), but little relation of D4.3 to these traits was found under rainfed regimen. The D4.3 also showed no significantly correlated with above traits (r=0.234; 0.404; -0.189; 0.279) under W×N interaction. Water treatment influenced the accumulation and the size of glutenin particles than any other factors when the genotypes were chosen.
Analysis of Water-Nitrogen Interaction on Single Kernel Weight, Protein content, Yield and DDT
Increasing N fertilizer application increased protein content and average single kernel weight, no matter whether irrigated or not. Compared with irrigated regimens, the protein percentage of GC8901 was decreased by 4.8-22.9%, while in TS23 and SN1391, the flour protein percentage sharply increased by 45.6% and 36.7% at 360 kg N ha~(-1) (N3) under rain-fed regimens. Across the two years, mean grain yields of three cultivars ranged from 6.9-9.9 mt ha~(-1) with irrigation, Rainfed condition led to decline by 2.0-38.2% of the yield. Dough Development Time (DDT) was recognized as an indicator of dough strength and baking quality. we observed that DDT had a similar tendency to GMP content, and there was a strong relation between them (r = 0.739**). Taking the two water treatments together, there was no significant correlation between DDT and D4.3 (r = 0.190), although D3.2 and D4.3 increased under rainfed condition. However, a stronger correlation was observed between DDT and H/L value (r = 0.803**). Apparently, the amounts of H- and LMW-GS were more important than GMP content for DDT.
We suggest that when the genotypes were chosen, the irrigation and N level contributed more to the yield and DDT, when the N application was higher than 240 kg N ha~(-1), the increasing amount of grain yield was not significant, and the effects were often mixed. It might be helpful to explain that wheat tended to earn higher biological mass and yield under well irrigated conditions with sufficient fertilizer, while increasing single kernel weight and keeping more protein in storage under rainfed regimen.
三小麦品种的全长序列分析
高分子量谷蛋白亚基的前30个氨基酸是信号肽,决定着高分子量谷蛋白亚基的空间表达。1Ax可读框内仅含有4个半胱氨酸残基,但形成20个潜在的α螺旋结构,1Dx2,1Bx7可读框内含有4个半胱氨酸残基,形成9个潜在的α螺旋结构。1Dx5亚基含有5个半胱氨酸残基可以形成潜在的9个α螺旋。1Bx14可读框内仅含有2个半胱氨酸残基,形成11个潜在的α螺旋结。1Dy10,1By8,1By15,1Dy12可读框内均含有7个半胱氨酸残基,分别形成9,11,11,11个潜在的α螺旋结构。虽然高分子量谷蛋白亚基在进化过程中出现了不同程度的碱基序列的变异,但是其序列的同源性结构决定了它们的蛋白序列和空间结构是类似的,因而在面粉中行使的的功能也是类似的。因此,对形成HMW-GS分子结构有贡献的三个重要氨基酸残基脯氨酸、甘氨酸和谷氨酰胺的含量也是优质面团粘弹性形成的重要参考标准。
亚基含量的HP-HPLC分析
灌溉条件下,强筋小麦GC8901和中筋小麦TS23籽粒的HMW-GS含量和H/L值随着氮肥施用量的增加而增加,LMW-GS的含量在施氮量为120 kg ha~(-1)时达到最大,弱筋小麦品种SN1391则表现出相反的趋势。旱作栽培模式下,中、高氮肥使用量(240 kg N ha~(-1),360 kg N ha~(-1))均提高了三小麦品种的H-、LMW-GS含量,SN1391提高量尤为显著,不施氮和低氮施用量(120 kg ha~(-1))显著降低了三小麦品种的H-、LMW-GS含量。
水氮互作条件下,三小麦品种的1Dx2和1Dx5亚基的相对含量最高,约占总HMW-GS含量的42%和38%。1Bx7和1Dy12亚基相对含量次之,约占31%和26%,1Ax1和1By8亚基的相对含量最小,仅占14%和12%。相互连锁的亚基对的相对含量在不同氮水平上变化不显著,但单个亚基的相对含量对水旱处理的反应不一。1Dy10亚基、1By15亚基在不同氮肥水平条件下占HMW-GS的比值是稳定的,之间无显著性差异;在非灌水条件下,7亚基、8亚基、1亚基的比值在GC8901中是稳定的,氮肥处理之间无显著性差异。与水地相比较,旱作处理使GC8901的1Ax1亚基的相对含量增加了10%,1Bx7亚基的相对含量增加了26.5%,1By8增加了85.7%,然而1Dx5和1Dy10亚基的相对含量保持不变。TS23旱作栽培条件下,1Ax亚基的相对含量下降了25%,1Bx7增加了12.2%而1By8增加了38.7%。SN1391各个亚基的相对含量在水肥料处理间则表现出相对稳定的趋势。亚基对之间相比较,1Bx7+1By8亚基对对环境反应更敏感,在中筋和弱筋小麦籽粒中,亚基对1Dx2+1Dy12往往表达量较高。相互连锁的亚基对的相对表达量表现持一定的互补和协同效应。水、氮栽培措施主要通过增加单位籽粒中的亚基含量来改善小麦的谷蛋白质量。
小麦籽粒GMP含量变化
3个小麦品种的GMP含量的大小依次顺序为:GC8901>TS23>SN1391。在灌水条件下,GMP含量都随着氮肥水平的增加而增加。在非灌水条件下,强筋小麦GC8901GMP含量随氮肥水平的增加呈抛物线状,在施氮量为240 kg ha~(-1)时GMP含量最高。与灌溉相比较,TS23和SN1391的GMP含量在不施氮时最高,分别增加了6-15.4%和30.9-82%,随着施氮量的增加GMP含量下降,中、高氮肥(240 kg N ha~(-1)、360 kg N ha~(-1))处理间无显著差异。在灌水条件下,增施氮肥有利于GMP在小麦籽粒中的积累,旱作栽培模式有利于中筋、和弱筋小麦籽粒的GMP的积累,但不同氮肥水平之间差异不显著。
小麦籽粒GMP的粒度分布
小麦成熟期的GMP粒度分布趋势相似,粒径范围为0.375~256.9μm。小麦GMP颗粒的体积分布为双峰曲线,第一个峰值出现在3.5~4.5μm之间,第二个峰值出现在116~140μm之间,不同品种间及处理间差异较大。GMP颗粒的表面积分布亦呈双峰曲线,数目分布为单峰曲线。小麦GMP数目主要由<10μm颗粒组成(约占99.8%以上),体积分布的47.57%~69%集中在12-100μm,而GMP表面积分布< 12μm,12-110μm和> 110μm百分数分别占72.3-89.4%, 9.5-21.5%和1.1-10.1%。
灌水条件下,强筋小麦GC8901籽粒GMP的12-110μm、< 110μm的表面积分布比例随着施氮量的增加先下降后显著上升。与旱作相比较,强筋小麦GC8901 < 12μm的GMP表面积分布比例显著下降。弱筋小麦SN1391表现出与GC8901完全相反趋势。灌溉条件下,中筋小麦TS23 < 12μm随着施氮量的增加而降低,12-110μm和> 110μm的GMP表面积分布比例升高,而旱作条件下,TS23不施氮水平的GMP表面积分布比例最高。三小麦品种D4.3和D3.2的变化趋势与GMP > 110μm颗粒的表面积分布趋势基本一致。旱作、低氮栽培条件能显著提高强筋小麦和弱筋小麦籽粒的体积分布比例。与旱作相比较,灌溉使TS23和GC8901 < 4.5μm的GMP颗粒的数目分布增加,4.5-12μm和> 12μm的数目分布降低,而增施氮肥使>12μm的GMP颗粒的数目增加。SN1391 <4.5μm的GMP颗粒的数目比例随施氮量而增加,4.5-12μm颗粒的数目比例降低, > 12μm颗粒的数目比例保持不变。
增施氮肥有利于GC8901籽粒HMW-GS的积聚和小粒径GMP的成长为大粒径GMP,调节了弱筋小麦GMP中等粒径颗粒和小粒径GMP颗粒数目的分布,从而形成更大的表面积分布,而不是大粒径的GMP颗粒,水分亏缺可以加速这两个调节过程。
小麦籽粒GMP的电镜观测
通过SEM电镜照片可以看出,GC8901籽粒的GMP颗粒的体积明显大于SN1391和TS23的GMP颗粒的体积,且SN1391GMP多为粒径<1.5μm小颗粒,这与激光粒度分析仪的结果相一致。这应当是由于GC8901(1、5+10、7+8)比SN1391(2+12、14+15)的亚基种类多出了1亚基且所含亚基种类的分子量较大,所以形成的GMP体积也较大的原因导致的。而从GC8901籽粒中的GMP在不同蔗糖浓度梯度下离心、分层后的所得的SDS-PAGE电泳图片看出,各层GMP所含高低分子量谷蛋白亚基种类没有差别,但无法确定其亚基的比例。可见,GMP大颗粒是有许多GMP小颗粒包裹形成,而TEM和SEM图像也表明大的GMP颗粒是由小GMP颗粒聚合而成的,较小的GMP颗粒是由H-和LMW-GS以不同比例组合而成的。
水氮互作对小麦产量和品质的影响
无论是在灌溉还是雨养条件下,增施氮肥均增加了籽粒的蛋白质含量和籽粒的千粒重。在高氮水平下(360kg ha~(-1)),相比灌溉条件,旱作栽培模式下的GC8901的蛋白质含量下降4.8-22.9%,而TS23和S1391则提高了45.6%和36.7%。灌水条件下三个小麦品种的平均产量为6.9-9.9 mt ha~(-1),而旱作模式下三个小麦品种的产量下降了2.0-38.2%,当施氮水平高于240kg N ha~(-1),产量增加不显著。DDT是反映面团的烘烤品质的重要指标,三小麦品种的DDT和籽粒GMP含量变化表现出的趋势,表明DDT和GMP含量之间密切的关系。同时,D3.2和D4.3与谷蛋白含量、HMW-GS含量、LMW-GS含量、DDT、H/L值、GMP含量和淀粉与蛋白质的比值均显著正相关,与单粒重呈极显著负相关,与籽粒淀粉含量相关不显著,但是在旱作栽培条件下D4.3与这些性状没有表现出明显的相关性。因此,H-、LMW-GS的含量和GMP的粒度分布决着面团烘焙品质的好坏。水肥充足的条件下小麦倾向于获得较高的生物量和产量,而在水分亏缺条件下则倾向于提高单粒重和蛋白质含量,以储存较多的能量。
The composition of high molecular weight glutenin subunit (HMW-GS) and low molecular weight glutenin subunit (LMW-GS) in wheat kernel (Triticum aestivum L.) affects the size of glutenin macropolymer (GMP), which is considered the important flour quality traits in wheat. Three wheat cultivars with different end-use quality were used to study the effects of water-nitrogen management (W×N) on relative HMW-GS and GMP contents. Under irrigation, increased N levels promoted the accumulation of H- and LMW-GS, GMP content and the proportion of the larger particle of GMP, with higher yield and better end-use quality. Under rainfed conditions, increased N fertilizer also increased protein content in the three cultivars. As opposed to the Gaocheng8901 (GC8901), the Shannong1391 (SN1391) and Taishan23 (TS23) got higher H- and LMW-GS contents and larger GMP particles, yields of both reduced. Irrigation increased the yield markedly, the N application significantly affected the glutenin, but the W×N interaction was not significant. On one hand, increased N levels improved the dough development time (DDT) by increasing the numbers of larger GMP particles and enhancing volume and surface area percentages on the other. It was suggested that the interaction was a complicated network and some individual effect might be important. When managing for wheat yield and quality, growers should first consider the water factor, then the N fertilizer factor to improve the end-use quality of wheat.
Full-length HMW-GS Sequence Analysis of tthree wheat cultivars
The HMW-GS pairs 1Dx5+1Dy10, 1Bx14+1By15 are recognized to positively correlate with bread-making quality. Both the reading frames (ORFs) of 1Ax and of 1Bx7 contain 4 cysteines, but the ORF of 1Ax forms 20 potentialα-helix structures while 1Bx7 forms 9 potentialα-helix structures. The ORF of 1Dx which forms 9 potentialα-helix structures contains 5 cysteines; the ORF of 1Bx14 which forms 11 potentialα-helix structures contains only 2 cysteines. The ORFs of 1Dy10, 1By8, 1By15 and 1Dy12 all contain 7 cysteines and form respectively 9, 11, 11 and 11 potentialα-helix structures. The DNA sequences showed they are homologous and would had the similar structure and function in wheat kernel. So the cysteine residue which plays important roles in forming the intra- and inter molecular structures of HMW-GS could not be used as the sole evaluation criterion for the merits of subunits. The proline, glycine and glutamine play important roles in forming advanced structure of glutenin, so the three amino acid residues would be the important reference standard for forming the visco-elasticity of quality dough. Meanwhile, the three amino acids have taken a large proportion in x- style HMW-GS. It means that x- style subunits contribute more for dough quality.
Differences in Experimental Factors
Analysis of variance for the contents of subunits, GMP content, the D4.3, D3.2, the single kernel weight, yield, grain protein content and the DDT made it possible to identify the sources of variation. Nitrogen (N) main effect and water×N (W×N) interaction failed to influence those traits except single kernel weight and grain protein content. The Student Newman and Keuls test (p≤0.05) showed that subunits in 1Bx chromosome, HMW-GS content and yield were significantly affected by water treatment. Although the N main effect as well as the W×N were not significant through three wheat cultivars, some individual effect was important to determine the quality and yield. It was suggested the interaction was a complicated network. Variations due to years and in interactions between year and nitrogen (Yr×N), year and water (Yr×W) treatment were not significant (p≤0.05). Since year was not a significant factor in the total reaction, some data from both years were averaged.
RP-HPLC Analysis for Glutenin
Considering the three cultivars, the 1Dx2 and 1Dx5 subunits accumulated to the highest level about 42% and 38% of total high molecular weight glutenin subunit (HMW-GS) content. The 1Bx7 and 1Dy12 subunits were present in a smaller amount, 31% and 26%, and the 1Ax1 and 1By8 subunits were the least abundant and nearly equal, 14% and 12%. The proportions of linked subunit pairs were nearly equal among different N levels, but the proportions of single subunits showed various under the two water treatments. As compared with irrigated regimen, rainfed regimen increased the mean proportion of 1Ax1 subunit by 10%, decreased 1Bx7 by 26.5%, increased 1By8 by 85.7%, whereas, the proportion of 1Dx5 and 1Dy10 remained stable in GC8901. The mean proportion of 1Ax1 in TS23 under irrigated regimen increased by 25%, 1Bx7 increased by 12.2% and 1By8 decreased by 38.7% compared with of under rain-fed regimen. Water treatment showed no significant effects on the subunit pairs in SN1391 with increased N application. Take all the subunit pairs into account, 1Bx7+1By8 seemed more sensitive to the environment and 1Dx2+1Dy12 always high expressed in middle and weak gluten wheat. We observed that under irrigation, both in GC8901 and TS23, the content of HMW-GS and H/L value increased with N increased, and the LMW-GS contents reached the maximums at 120 kg N ha~(-1) (N1). However, the trend was opposite in SN1391 under the same condition. Under rainfed regimen, the contents of H- and LMW-GS increased in three wheat cultivars with higher N application (N2, N3), especially in SN1391. However, the H- and LMW-GS contents of three cultivars decreased markedly under rainfed condition with low N application (N1).
These results indicated that the ratios of the linked subunit pair’s proportions to the mean proportion of different N levels were nearly identical. The two linked subunit pairs had certain complementary effects, the final proportions showed no significant relationship with W×N interaction and mainly determined by genotypes. Whereas, a significantly correlated was shown among single kernel weight, protein content, GMP content (r = -0.829**; 0.563*; 0.803**) and H/L. It was indicated that the grain kernel production interacts with H- and LMW-GS contents. Rainfed condition could increase the contents of H- and LMW-GS and H/L value in three wheat cultivars.
Content of GMP Particles
The contents of GMP in three wheat cultivars were ordered as follows: GC8901 > TS23 > SN1391. The contents of GMP all increased as N rates increased under irrigated regimen. But under rainfed regimen, the content of GMP in GC8901 showed difference with the N level, and reached the maximum at 240 kg N ha~(-1) (N2); the GMP contents of TS23 and SN1391 both were markedly increased by 6-15.4% and 30.9-82% with no N applied (N0), and tended to decline as increased N application, while the differences of contents between N application of 240 kg N ha~(-1) (N2) and 360 kg N ha~(-1) (N3) were not significant. The results indicated that increasing N level could promote the accumulation of GMP in three wheat cultivars under irrigated regimen. Rainfed condition significantly promoted the accumulation of GMP in middle and weak gluten wheat, but the content remained stable with N rates.
GMP Particle Distribution
The surface area percent of GMP particles at < 12μm, 12-110μm and > 110μm in three cultivars took up 72.3-89.4%, 9.5-21.5% and 1.1-10.1%. Under irrigation, the surface area percentages of GMP particles at 12-110μm and < 110μm in GC8901 decreased first and then markedly increased with N increased, whereas that of < 12μm decreased markedly when compared with those under rainfed regimen. However, those in SN1391 showed a character of opposite extremes. Those at < 12μm in TS23 decreased with N increased, and those of 12-110μm and > 110μm increased under irrigation regimen. While under rainfed regimen, the GMP surface area percentage of TS23 reached the maximum value with no N applied (N0). The irrigation was beneficial to increasing the surface area percent of GMP particles as increased N application, and sight water stress could markedly promote those in weak and middle gluten wheat.
The data in Table 4 indicated that D4.3 and D3.2 were completely in line with the surface area percentages of GMP particles at > 110μm. Dispersed GMP of three cultivars under W×N interaction displayed that rainfed regimen and lower N level could promote the volume of particles in middle and weak gluten wheat. Strong gluten wheat was more adapted to plenteous irrigation and higher N application. And interestingly, the D4.3 under rain-fed regimen of TS23 and SN1391 were bigger than that of in GC8901, but the content of GMP in TS23 and SN1391was less.
Irrigated and rainfed conditions had different influence on percent number of GMP particles in three wheat cultivars. Under irrigation, the number percentage of GC8901 and TS23 at < 4.5μm increased, while those of 4.5-12μm and > 12μm decreased when compared with those under rainfed condition. Increased N level contributed to increase the GMP particles numbers of GC8901 and TS23 at >12μm. The increased N level increased number percentage of GMP particles in SN1391 at <4.5μm, decreased that of 4.5-12μm, whereas that of > 12μm remained the same. The GMP particles number percentage responses of SN1391 to water treatment were similar to those to N application. The number distribution revealed that increased N level could increase the numbers large GMP particles in strong and middle gluten wheat. While in weak gluten wheat, N effect and W×N interaction showed no influence on the numbers of larger GMP particles.
Observations of GMP Particles
To further analyze the effects of W×N interaction on glutenin particle, GMP dispersions of three wheat cultivars were examined for particulate structures obtained. The SEM figures showed that large GMP particle was made from the polymerization of small GMP particles, the smaller GMP particles were made of H- and LMW-GS in various proportions. We also estimated that GC8901 got a highest H/L ratio ranged from 0.36-0.47 and the highest D4.3, TS23 got a middle H/L ratio ranged from 0.26-0.33 and smaller D4.3, SN1391 got the least H/L ratio ranged from 0.20-0.25 and least D4.3. It indicated that D4.3 was mainly determined by genotypes, but there was no evidence to confirm their proportions of H- and LMW-GS in this paper. The D4.3 under irrigated regimen showed strong relation to protein content, GMP content, single kernel weight and H/L value (r=0.822**; 0.755**; -0.806**; 0.698*), but little relation of D4.3 to these traits was found under rainfed regimen. The D4.3 also showed no significantly correlated with above traits (r=0.234; 0.404; -0.189; 0.279) under W×N interaction. Water treatment influenced the accumulation and the size of glutenin particles than any other factors when the genotypes were chosen.
Analysis of Water-Nitrogen Interaction on Single Kernel Weight, Protein content, Yield and DDT
Increasing N fertilizer application increased protein content and average single kernel weight, no matter whether irrigated or not. Compared with irrigated regimens, the protein percentage of GC8901 was decreased by 4.8-22.9%, while in TS23 and SN1391, the flour protein percentage sharply increased by 45.6% and 36.7% at 360 kg N ha~(-1) (N3) under rain-fed regimens. Across the two years, mean grain yields of three cultivars ranged from 6.9-9.9 mt ha~(-1) with irrigation, Rainfed condition led to decline by 2.0-38.2% of the yield. Dough Development Time (DDT) was recognized as an indicator of dough strength and baking quality. we observed that DDT had a similar tendency to GMP content, and there was a strong relation between them (r = 0.739**). Taking the two water treatments together, there was no significant correlation between DDT and D4.3 (r = 0.190), although D3.2 and D4.3 increased under rainfed condition. However, a stronger correlation was observed between DDT and H/L value (r = 0.803**). Apparently, the amounts of H- and LMW-GS were more important than GMP content for DDT.
We suggest that when the genotypes were chosen, the irrigation and N level contributed more to the yield and DDT, when the N application was higher than 240 kg N ha~(-1), the increasing amount of grain yield was not significant, and the effects were often mixed. It might be helpful to explain that wheat tended to earn higher biological mass and yield under well irrigated conditions with sufficient fertilizer, while increasing single kernel weight and keeping more protein in storage under rainfed regimen.
引文
曹宏鑫,王世敬,戴晓华.土壤基础肥力和肥水运筹对春小麦产量和品质与植株氮素状况的影响[J].麦类作物学报,2003,23(2):52-56
蔡瑞国,尹燕枰,张敏,戴忠民,严美玲,付国占,贺明荣,王振林.氮素水平对藁城8901和山农1391籽粒品质的调控效应.作物学报, 2007, 33(2): 304-310
邓志英,田纪春,刘现鹏.不同高分子量谷蛋白亚基组合的小麦籽粒蛋白组分及其谷蛋白大聚合体的积累规律[J].作物学报,2004,30(5):481-486
邓志英,田纪春,张永祥,王延训,孙国兴,盛峰.冬小麦高低分子量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): 308-3
何照范.粮油籽粒品质及其分析技术.北京:中国农业出版社, 1985, 144-150
贾光锋,范丽霞,王金水.小麦面筋蛋白结构,功能性及应用.粮食加工, 2004, (2):11-13
姜鸿明,余松烈,于振文,赵倩,邱化蛟,丁晓义.小麦谷蛋白聚合作用对增施氮肥的反应.麦类作物学报, 2003, 23(2): 43-46.
姜东,于振文,李永庚.施氮水平对高产小麦蔗糖含量和光合产物分配及籽粒淀粉积累的影响.中国农业科学, 2002, 35(2): 157-162
李保云,等.小麦高分子量谷蛋白亚基与小麦品质性状关系的研究[J].作物学报.2000, 26(3):322-326
李建敏,王振林,高荣岐,李圣福,蔡瑞国,闫素辉,于安玲,尹燕枰.强、弱筋小麦籽粒形成期蔗糖、淀粉合成相关酶活性及其与氮代谢的关系.作物学报, 2008, 34: 1019-1026.
李青常,王振林,张艳,刘霞,石书兵.施氮水平对小麦面条加工品质的影响.中国农业科学, 2005, 38(2): 420-424
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
刘宝龙,张怀刚.弱筋小麦品质高分子量谷蛋白亚基组成分析.西北农业学报, 2007, 16(2): 19-23.
刘建军,何中虎,赵振东,刘爱峰,宋建, R J Pena.小麦品质与干白面条品质参数关系的研究.作物学报, 2002, 28(6): 738-742
刘振兴,杨振华,邱孝煊,林增泉.肥料增产贡献率及其对土壤有机质的影响.植物营养与肥料学报,1994,(1):19-26
马宗斌,何建国,王小纯,马新明.氮素形态对两个小麦品种籽粒内源激素含量的影响.中国农业科学,2008,41,(1):63-69
马传喜,等.小麦胚乳蛋白质组成及高分子量麦谷蛋白亚基与烘烤品质的关系[J].作物学报.1993,19(16):562-566
马元喜.不同土壤小麦根系生长动态研究.作物学报, 1987,13(1):37-44
盛靖,郭文善,胡宏等.小麦淀粉合成关键酶活性及其与淀粉积累的关系.扬州大学学报(农业与生命科学版),2003,24(3):49-53
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作物学报, 2005, 31(4): 431-437
潘庆民,于振文,王月福等.追氮时期对小麦旗叶中蔗糖合成与籽粒中蔗糖降解的影响.中国农业科学,2002,35(7):771-776
潘庆民,于振文,王月福.小麦开花后旗叶中蔗糖合成与籽粒中蔗糖降解.植物生理与分子生物学学报. 2002b, 28(3):235-240.
师俊玲,魏益民.小麦蛋白质和淀粉与面条品质性状关系的研究进展.郑州粮食学院学报, 1998, 13(3): 1-5
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.
石书兵,马林,石庆华,刘霞,陈乐梅,刘建喜,王振林.不同施氮时期对冬小麦子粒蛋白质组分及其动态变化的影响.植物营养与肥料学报, 2005, 11(4): 456-46
王海波,庞惠玲,刘志宏.小麦谷蛋白研究方法的比较分析[J].中国农学通报. 2006, 22(7): 125 - 129
王瑞,等。一些优质小麦及其杂种后代高分子量谷蛋白亚基组成与面包品质之关系[J].西北农业学报.1995,4(4):25-30
王晓英,贺明荣,李飞,等.水氮耦合对强筋冬小麦子粒蛋白质和淀粉品质的影响.植物营养与肥料学报, 2007, 13(3): 361-367
王旭东,于振文,石玉等.磷对小麦旗叶氮代谢有关酶活性和籽粒蛋白质含量的影响.作物学报,2006,32(3):339-344
王旭东,于振文,王东.钾对小麦茎和叶鞘碳水化合物含量及籽粒淀粉积累的影响.植物营养与肥料学报,2003,9(1):57-62
王月福,于振文,李尚霞.氮素营养水平对小麦开花后碳素同化、运转和产量的影响.麦类作物学报, 2002, 22(2): 55-59
王月福,于振文,李尚霞,等.施氮量对小麦籽粒蛋白质组分含量及加工品质的影响[J].中国农业科学.2002,35(9):1071-1078
闫素辉,王振林,戴忠民,李文阳,付国占,贺明荣,尹燕枰.两个直链淀粉含量不同的小麦品种籽粒淀粉合成酶活性与淀粉积累特征的比较.作物学报, 2007, 33: 84-89.
闫素辉,尹燕枰,李文阳,李勇,等.密穂与疏穂型小麦强、弱势籽粒淀粉积累及库强度的比较.中国农业科学, 2009, 42(8): 2706-2715
闫素辉,尹燕枰,李文阳,李勇,梁太波,等.花后高温对不同耐热性小麦品种籽粒淀粉形成的影响.生态学报, 2008, 28(12): 6138-6147
晏月明,刘广田,Prodanovie S.小麦谷蛋白亚基的凝胶电泳分离及其品种鉴定[J].中国粮油学报.1998,13:1-5
姚大年,李保云,梁荣奇,刘广田.基因型和环境对小麦品种淀粉性状及面条品质的影响.中国农业大学学报,2000, 5(1): 63-68
尹静,胡尚连,肖佳雷,李文雄.不同形态氮肥对春小麦品种籽粒淀粉及其组分的调节效应.作物学报, 2006, 32(9): 1294-1300
于振文.优质专用小麦品种及栽培.北京:中国农业出版社. 2001
赵广才,常旭虹,刘利华,等.施氮量对不同强筋小麦产量和加工品质的影响[J].作物学报.2006,32(5):723—727
赵广才,张艳,刘利华,等.不同施肥处理对冬小麦产量、蛋白组分和加工品质的影响[J].作物学报.1993,19(6):562—567
赵俊晔,于振文.施氮量对小麦强势和弱势子粒氮素代谢及蛋白质合成的影响[J].中国农业科学.2005,38(8):1547—1554
赵友梅,等.高分子量麦谷蛋白亚基的SDS-PAGE图谱在小麦品质研究中的应用[J].作物学报.1990,16(3):208-218
张定一,张永清,杨武德,等.施氮量对不同小麦品种高分子量麦谷蛋白亚基表达量及品质影响的研究[J].植物营养与肥料学报.2008,14(2):235—241
赵和,等.小麦高分子量麦谷蛋白亚基遗传变异及其与品质和其他农艺性状关系的研究[J].作物学报.1994,20(l):67-75
张春庆,李晴祺.影响普通小麦加工馒头质量的主要品质性状的研究.中国农业科学, 1993, 26(2): 39-46
张金锐,刘勇,林刚,何光源.小麦高分子量谷蛋白亚基序列及其结构与加工品质的关系.中国生物工程杂志, 2006, 26(8): 103-110
张军,张洪程,戴其根,等.追施氮肥时期对淮南中筋小麦品质的影响.扬州大学学报(农业与生命科学版), 2003, 24(2): 44-48
张平平,张勇,夏先春,等.小麦贮藏蛋白反相高效液相色谱分析体系研究[J].中国农业科学. 2007,40(5):1002-1009.
Alessandro M, Laura E, Marco M. Post-anthesis accumulation and remobilization of dry matter, nitrogen and phosphorus in durum wheat as affected by soil type. Europ.J.Agronomy, 2007, 26:179-186
Andersm E C. Corn root growth and distribution as influenced by tillage and nitrogen fertilizer. Agronomy Journal, 1987, 79:544-559
Ayoub, M., S. Guertin, J. Fregeau-Reid, and D.L. Smith. 1994. Nitrogen fertilizer effect on breadmaking quality of hard red spring wheat in eastern Canada. Crop Sci. 34:1346–1352.
Ayoub, M., S. Guertin, and D.L. Smith. 1995. Nitrogen fertilizer rate and timing effect on bread wheat protein in eastern Canada. Crop Sci. 174:337–349.
Bae J M, Liu J R. Molecular cloning and characterization of two novel isoforms of the small subunit of ADPglueose pyrophosphorylase from sweet potato. Mol. Gen. Genet, 1997, 254: 179-185
Ball S, Guan H P, James M, et al. From glycogen to amylopectin: a model for the biogenesis of the starch granule. Cell, 1996, 86: 349-352
Batey I L, Curtin B M, Moore S A. Optimization of Rapid Visco Analyser test conditions for predicting Asian noodle quality. Cereal Chem., 1997,74(4): 497-501
Barneix A J, Guitman M R. Leaf regulation of the nitrogen concentration in the grain of wheat plants. Journal of Experimental Botany,1993,44(1):607-612
Bengough A G, Young I M. Root elongation of seeding peas through layered soil of different penetration resistances. Plant and Soil, 1993,149:129-139
Bernd Steingrobe, Harald Schmid, Reinhold Gutser, Norbert Claassen. Root production and root mortality of winter wheat grown on sandy and loamy soils in different farming systems. Biol Fertil Soils, 2001, 33:331-339
Behdaie A E, Hall G D. Farquhar Water-use efficiency and carbon Isotope Discrimination inWheat. Crop Science, 1991, 31: 1282-1288.
Blechl AE, Anderson OD (1996) Expression of a novel highmolecular-weight gluteninsubunit gene in transgenic wheat. Nat Biotechnol 14:875–879
Blechl AE, Le HQ, Anderson OD (1998) Engineering changes in wheat flour by genetic transformation. J Plant Physiol 152:703–707
Blechl A, Lin J, Nguyen S, Chan R, Anderson OD, Dupont FM (2007) Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. J Cereal Sci 45:172–183
Bietz J.A. Analysis of wheat gluten proteins by high-performance liquid chromatography[J]. I. Baker’s Dig. 1984, 58: 15-32
Bietz J.A.Separation of cereal proteins by reversed-phase high-performance liquid chromatography [J]. Journal of Chromatography. 1983, 255: 219-238
Blumenthal C, Bekes F, Gras P W, et al. Identification of wheat genotypes tolerant to the effects of heat stress on grain quality. Cereal Chem., 1995, 72: 539-544
Branlard G.and Dardevet M.Diversity of grain protein and bread wheat quality.II.Correlation between high molecular weight subunits of glutenin and flour quality characteristics [J]. Journal of Cereal Science.1985,3:345-354
Braun H J, Payne T S, Morgounov A I, van Ginkel M, Rajaram S. The challenge: one billion tons of wheat by 2020. In: Slinkard A E (ed) Proceeding of the 9th International Wheat Genetics Symposium. Saskatchewan: University Extension Press, 1998, 33-40
Bushuk W. Wheat breeding for end-product use. Euphytica, 1998, 100(1-3): 137-145
Buttrose B L. Ultrastructure of the developing wheat endosperm. Aust. J. Biol. Sci., 1963, 16: 305-310
Butow, B.J., B.W. Ma, K.R. Gale, G.B. Cornish, L. Rampling, O. Larroque, M.K. Morrell, and F. Bekes. 2003. Molecular discrimination of Bx7 alleles demonstrates that a highly expressed high-molecular-weight glutenin allele has a major impact on wheat flour dough strength. Theor Appl Genet. 107: 1524-1532
Carceller, J.L., and T. Aussenac. 2001. SDS-insoluble glutenin polymer formation in developing grains of hexaploid wheat: the role of the ratio of high to low molecular weight glutenin subunits and drying rate during ripening. Aust. J. Plant Physiol. 28: 193-201
Clyde, D., L. George, N. Hamid, M. Finlay, and J.H. Rob. 2005. Heat stress and genotype affect the glutenin particles of the glutenin macropolymer-gel fraction. J Cereal Sci. 42: 69-80.
Chen Y, Yuan L P, Wang X H, Zhang D Y, Chen J, Deng Q Y, Zhao B R, Xu D Q. Relationship between grain yield and leaf photosynthetic rate in super hybid rice. J. Plant Physiol. Mol. Biol., 2007, 33(3): 235-243
Cressey P J, Campbell W P, Wrigley C W. Statistical correlation between quality attributes and grain protein composition for advanced lines of crossbred wheat. Cereal Chemistry,1987,64(4):299-301
COL ETTE LARRE , YVES NICOLAS ,CLAUDE DESSERME , et al . Preparative Separation of High and Low Molecular Weight Subunits of Glutenin from Wheat [J ] . Journal of Cereal Science ,1997 ,25 :143 - 150.
Crosbie G B. The relationship between starch swelling properties, paste viscosity and boiled noodle quality in wheat flour. Journal of Cereal Science, 1991, 13:145-150
Dai Z M, Yin Y P, Zhang M, Li W Y, Yan S H, Cai R G, Wang Z L. Starch granule size distribution in wheat grains under irrigated and rainfed conditions. Acta Agron. Sin. 2008, 34:Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2003a. Glutenin macropolymer: a gel formed by glutenin particles. J Cereal Sci. 37: 1-7.
Daniel, C., and E.Trib?. 2002. Changes in wheat protein aggregation during grain development: effects of temperatures and water stress. Eur J Agron. 16: 1-12.
Deng, Z.Y., J.C. Tian, R.B. Hu, X.F. Zhou, and Y.X. Zhang. 2006. Effects of genotype and environment on wheat main quality characteristics. Acta Ecologica Sinica. 26(8): 2757-2763. (in Chinese)
Deng, Z., J. Tian, L. Zhao, Y. Zhang, and C. Sun. 2008. High temperature-induced changes in high molecular weight glutenin subunits of Chinese winter wheat and its influences on the texture of Chinese noodles. J Agron Crop Sci. 194: 262-269.
Denyer K. Identification of multiple isoforms of soluble and granule-bound starch synthesis in developing wheat endosperm, Planta, 1995, 196: 256-265
Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2003b. Understanding the link between GMP and dough: from glutenin particles in flour towards developed dough. J Cereal Sci. 38: 157-165.
Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2005. The effect of mixing on glutenin particle properties: aggregation factors that affect gluten function in dough. J Cereal Sci. 41: 69-83.
Don, C., G. Mannb, F. Bekesb, and R.J. Hamer. 2006. HMW-GS effect the properties of glutenin particles in GMP and thus flour quality. J Cereal Sci. 44: 127-136.
Douglas C D, KuoT M, Felker F C. Enzymes of sucrose and hexose metabolism indeveloping kernels of two inbreds of maize. Plant Physiol, 1988, 86: 1013-1019
Dreccer M F, Grashoff C, Rabbinge R. Source-sink ration in barley during grain filling: effects on senescence and grain protein concentration. Field crops research, 1997,49: 269-277
Dupont, F.M., and S.B. Altenbach. 2003. Molecular and biochemical impacts of environmental factors on wheat grain development and protein synthesis. J Cereal Sci. 38: 133-146.
DuPont,F.M.,Chan,R.,Lopez,R.,Vensel,W.H.Sequential extraction and quantitative recovery of gliadins,glutenins,and other proteins from small samples of wheat flour.[J].Journal of Agricultural and Food Chemistry.2005,53:1575-1584.
DuPont,F.M.,Hurkman,W.J.,Hurkman,W.J.,Vensel,W.H.,Chan,R.etal.Differential accumulation of sulfur-rich and sulfur-poor wheat flour proteins is affected by temperature and mineral nutrition during grain development.[J]Journal of Cereal Science. 2006b, 44: 101-112.
DuPont, F.M., R. Chan, and R. Lopez. 2007. Molar fractions of high-molecular-weight glutenin subunits are stable when wheat is grown under various mineral nutrition and temperature regimens. J Cereal Sci. 45: 134-139.
Eliasson A C, Karlsson R. Gelatinization properties of different size classes of wheat starch granules measured with differential scanning calorimetry. Starch, 1983, 35: 130-133.
Endo S, Karibe S, Okada K, Nagao S. Factors affecting gelatinization properties of wheat starch. Nippon Shikuhin Kogyo Gakkaishi, 1988, 35: 7-14
Eugenio L, Coelho, Dan D. Root distribtution and water uptake patterns of corn under surface and subsurface drip irrigation. Plant and soil,1999,206:123-126
Eva Johansson, Maria Luisa prieto-Linde,et al.Effects of Wheat Cultivar and Nitrogen Application on Storage Protein Composition and Breadmaking Quality, Cereal Chemists, 2001, 78(1):19-25.
Evers A D. Scanning electron microscopy of wheat starch.Ⅲ.Granule development in the endosperm. Starch, 1971, 23: 157-160
Evers A.D, Lindley J. The particle-size distribution in wheat endosperm starch, J. Sci. Food Agric., 1977, 28: 98-102
Fernando D, Francisco J C. Characterization of the endoprotease appearing during wheat grain filling. Plant physiol, 1996, 112:1211-1217
Fischer B., Grossmann J., Roth V., Gruissem W., Baginsky S., Buhmann J.M. Semi-supervised LC/MS alignment for differential proteomics[J]. Bioinformatics. 2006,14: 132–140
Fisher D B, Gifford R M. Accumulation and conversion of sugars by developing wheat grains.VI, Gradients along the transport pathway from the peduncle to the endosperm cavity during grain filling[J]. Plant Physiol, 1986, 82: 1024-1030
Fu B X, Kovacs MIP, Wang C. A simple wheat flour swelling test. Cereal Chem, 1998, 75(4):566-567
Gebbing T, and Schnyder H. Pre-anthesis reserve utilization for protein and carbohydrate synthesis in grains of wheat. Plant Physiology, 1999, 121: 871-878
Ceoloni C, Jauhar PP (2006) Chromosome engineering of the durum wheat genome: Strategies and applications of potential breeding value. In: Singh RJ, Jauhar PP (eds) vol 2 Genetic Resources, Chromosome Engineering, and Crop Improvement: Cereals. CRC Press, Taylor & Francis group, pp 27-59
Ceoloni C, Ciaffi M, Lafiandra D, Giorgi B (1995) Chromosome engineering as a means of transferring 1D storage protein genes from common to durum wheat. In: Li ZS, Xin ZY (eds) Proceedings of the 8th International Wheat Genetics Symposium, 20–25 July 1993, Beijing, China. China Agricultural Scientech Press, Beijing, pp 159–163
Ceoloni C, Biagetti M, Ciaffi M, Forte P, Pasquini M (1996) Wheat chromosome engineering at the 4x level: the potential of different alien gene transfers into durum wheat. Euphytica 89:87–97
Goodman A M, Eends A P. The effects of soil bulk density on the morphology and anchorage mechanics of the root systems of sunflower and maize. Annals of Botany, 1999, 83:293-302
Gianibelli M C, Gupta R B, Lafiandra D, Margiotta B, Macritchie F. Polymorphism of high Mr glutenin subunits in Triticum tauschii: characterization by chromatography and electrophoretic methods. J Cereal Sci, 2001, 33(1): 39-52
Graveland, A., P. Bosveld, W.J. Lichtendonk, J.P. Marseille,J.H. E. Moonen, and A. Scheepstra. 1985. A model for the molecular structure of the glutenins from wheat flour. J Cereal Sci. 3(1): 1-16.
Gupta R B, Khan K, MacRitchie F. Biochemical basis of flour properties in bread wheatsⅠ. Effects of variation in the quantity and size distribution of polymeric protein. J Cereal Sci, 1993, 18(1): 23-41
Gupta, R.B., K. Khan, and F. Macritchie. 1993. Biochemical basis of flour properties in bread wheats. I. Effects of variation in the quantity and size distribution of polymeric protein. J Cereal Sci. 18(1): 23-41.
Gupta, R.B., Y. Popineau, J. Lefebvre, M. Cornec, J.G. Lawrence, and F. Macritchie. 1995. Biochemical basis of flour properties in bread wheats. II. Changes in polymeric protein formation and dough/gluten properties associated with the loss of low Mr or high Mr glutenin subunits. J Cereal Sci. 21: 103-116.
Herbert Wieser,Susanne Antes,Werner Seilmeier.Quantitative Determination of Gluten Protein Types in Wheat Flour by Reversed-Phase High-Performance Liquid Chromatography [J]. Cereal Chem.1998,75(5):644-650
Hou G.,Yamamoto Y.and Ng P.K.W.Relationship of glutenin subunits of selected U.S.soft wheat flours to rheological and baking properties[J].Cereal Chem.1996,73:358-363
Huang D.Y.and Khan K.Quantitative determination of high molecular weight glutenin subunit of hard red spring wheat by SDS-PAGE.II.Quantitative effects of individual subunits on breadmaking quality characteristics[J].Cereal Chem.1997b,74:786-790
Holm J, Bjorck I. Bioavailablity of starch in various wheat-based bread products: evaluation of metabolic responses in healthy subjects and rate and extent of in vitro starch digestion [J].American Journal of Clinical Nutrition,1992, 55(2):420-429
Hurkman W. J, McCue K F, S. Altenbach B, Effect of temperature on expression of genes encoding enzymes for starch biosynthesis in developing wheat endosperm, Plant Science 2003,164: 873-881
James M G, Robertson M G, Myers A M. Characterisation of the maize gene sugary, a determinant of starch composition in kernels. Plant Cell, 1995, 7: 417-429
James M G, Denyer K, Myers A M. Starch synthesis in the cereal endosperm. Current Opinion in Plant Biology, 2003, 6: 215-222
Jenner C F. Effects of exposure wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars. Immediate responses. Aust.J.Pant Physiol,1991,18:165-177
Johnson V V. Genetic advances in wheat protein quantity and composition, Pron.4th
Intern.Wheat Genet.Symp.Columbia, 1973.
Jiang, H.M., S.L. Yu, Z.W. Yu, Q. Zhao, H.J. Qiu, and X.Y. Ding. 2003. Response of wheat glutenin polymerization to increased N fertilization. Journal of Triticeae Crops. 23(2): 43-46. (in Chinese)
Jiang D, Cao W X, Dai T B, Jing Q. Activities of key enzymes for starch synthesis in relation to growth of superior and inferior grains on winter wheat spike. Plant Growth Regul., 2003, 41, 247-257
Jiang, D., H. Yue, B. Wollenweber, W. Tan, H. Mu, Y. Bo, T. Dai, Q. Jing, and W. Cao. 2009.Effects of post-anthesis drought and water logging on Aaccumulation of high-molecular-weight glutenin subunits and glutenin macropolymers content in wheat grain. J Agron Crop Sci. 195: 89-97.
K. Dong, C.Y. Hao, A.L. Wang, M.H. Cai, and Y.M. Yan. Characterization of HMW Glutenin Subunits in Bread and Tetraploid Wheats by Reversed-Phase High-Performance Liquid Chromatography[J]. Cereal Research Communications. 2009, 37(1):65–72
Konarev, V. G., Gavrilyuk, I. P., Gubareva, N. K., and Peneva, T. I. (1979). Seed proteins in genome analysis, cultivar identification, and documentation of cereal genetic resources: A review. Cereal Chem. 56(4):272.
Khelifi D.and Branlard G.The effect of HMW and nological quality of progeny from four crosses between poor breadmaking quality and strong wheat cultivars[J].Journal of Cereal Science.1992,16:195-209
Kim H.R., Bushuk W. Changes in some physicochemical properties of flour proteins due to partial reduction with dithiothreitol[J].Cereal Chem.1995, 72(5): 450-456
Kolster P., Krechting C.F., Van Gelder W.M.J. Quantification of individual high molecular weight glutenin subunits using SDS-PAGE and scanning densitometry[J]. Journal of Cereal Science.1992, 15:49-61
Konik C M, Mikkelsen L M, Moss R, and Gore P J. Relationships between physical starch properties and yellow alkaline noodle quality, Starch / staerke, 1994, 46: 292-299
Laemmli U.K.Cleavage of structural proteins during the assembly of the head of bacteriophage T4 [J].Natrue.1970, 227:680-685
Lafiandra D, Margiotta B, Colaprico G, Masci S, Roth MR, MacRitchie F (2000) Introduction of the D-genome related high- and low-Mr glutenin subunits into durum wheat and their effect on technological properties. In: Shewry PR, Tatham AS (eds) Wheat Gluten: Proceedings of the 7th International Workshop Gluten, 2–6 April 2000, Bristol, UK. The Royal Society of Chemistry, Cambridge, pp 51–54
Lawrence G.J.,MacRitchie F.and Wrigley C.W. Dough and baking quality of wheat lines deficient in glutenin subunits controlled by the Glu-A1,Glu-B1 and Glu-D1 loci[J].Journal of Cereal Science.1988,7:109-112
Lawrence G. J. The high-molecula-weight glutenin subunit composition of Australia wheat cultivars [J]. Aust. J. Agri. Res.1986,37:127-133
Larroque O.R., Bekes F., Wrigley C.W., Rathmell W.G. Analysis Luo C, Branlard G, Griffin B, et al. The effect of nitrogen and sulphur fertilization and their interaction with genotype on wheat glutenins and quality parameters[J].Journal of Cereal cience,2000,31:185-194.
Ley, I., 2005. Particle size analysis—evaluating laser diffraction systems in the light of ISO 13320-1, publication T-403, Beckman Coulter, High Wycombe,UK.
Lefebvre, J., Y. Popineau, G. Deshayes, and L. Lavenant. 2000. Temperature-Induced changes in the dynamic rheological behavior and size distribution of polymeric proteins for glutens from wheat near-isogenic lines differing in HMW glutenin subunit composition. Cereal Chem. 77(2): 193-201.
Liu, B.L., and H.G. Zhang. 2007. Analysis on the HMW-GS components of weak gluten wheat varieties. Acta Agriculturae Boreali-occidentalis Sinica. 16(2): 19-23. (in Chinese)
Liang, T.B., Y.P. Yin, R.G. Cai, S.H. Yan, W.Y. Li, Q.H. Geng, P. Wang, Y.H. Wu, Y. Li, and Z.L. Wang. 2008. HMW-GS accumulation and GMP size distribution in grains of shannong 12 grown in different soil conditions. Acta Agronomica Sinica. 34(12): 2160-2167. (in Chinese)
Lucow O.M.,Payne P.I.and Thachuk R.The HMW glutenin subunits compositions in Canadian wheat cultivars and their association with bread making quality[J].Journal of the Sciecne of Food and Agriculture.1989,46:451-460
Lookart G.Bean S.Ultrafast CE analysis of cereal storage proteins and its application to protein characterization and cultivar differentiation [J]. Agric.Food Chem. 2000, 48: 344-353
Mackay A D, Barber S A. Effect of nitrogen on root growth of two corn genotypes in the field. Agronomy Journal, 1986, 78: 699-703
Masc S, Egorov T A,Ronchi C,et al. Evidence for the presence of only one cysteine residue in theD-type low molecular weight subunits of wheat subunits of wheat glutenin[J].Journal of Cereal Science,1999, 29:17-25.
Masci, S., La?andra, D., Porceddu, E., Lew, E.J.-L., Tao, H.P. and Kasarda, D.D. (1993) D-glutenin Subunits: N-terminal Sequences and Evidence for the Presence of Cysteine' in Cereal Chem. 70, 581±585
Masci, S., Lew, E.J.L., La?andra, D., Porceddu, E. and Kasarda,D.D. (1995) Characterization of Low Molecular Weight Sub-units in Durum Wheat by Reversed-phase High-performance Liquid Chromatography and N-terminal Sequencing' in Cereal Chem. 72, 100±104
D'Ovidio, R., Simeone, M., Masci, S., Porceddu, E. and Kasarda,D.D. (1995) `Nucleotide Sequence of a g-type Glutenin Gene from a Durum Wheat: Correlation with a g-type Glutenin
Subunit From the Same Biotype' in Cereal Chem. 72, 443±449
Masoni A, Ercoli L, Mariotti M, Arduini I. Post-anthesis accumulation and remobilization of dry matter, nitrogen and phosphorus in durum wheat as affected by soil type. Eur J Agron, 2007, 26: 179-186
Materechera S A. Penetration of very strong soil by seeding roots of different species. Plant and soil, 1991, 135:31-41
Metakovsky EV (1991) Gliadin allele identification in common wheat. II. Catalogue of gliadin alleles in common wheat. J Genet Breed 45:325-343
Metakovsky EV, Akhmedov MG, Sozinov AA (1986) Genetic analysis of gliadin-encoding genes reveals gene clusters as well as single remote genes. Theor Appl Genet 73:278-285
Metakovsky EV, Ng PKW, Chernakov VM, Pogna NE, Bushuk W (1992) Gliadin alleles in Canadian Western Red Spring (CWRS) class of wheat eultivars: use of the two procedures of A-PAGE for gliadin separation. Genome (in press) Ng PKW, Bushu
Melas V ,Morel M H ,Autran J C. Simple and rapid methed for purifying low molecular weight glutenin from wheat [J ] . Cereal Chemistry ,1994 ,71 :234 - 237.
Mouille G, Maddelein M L, Ball S. Preamylopectin processing: A mandatory step for starch biosynthesis in plants. Plant Cell, 1996, 8: 1353-1366
Moonen C, Masoni A, Ercoli L, Mariotti M, Bonari E. Long-term changes in rainfall and temperature in Pisa, Italy. Agr. Med, 2001, 130: 11-22
Morel M.H.Acid polyaerylamide gel eleclrophoresis of wheat gluteins:a new tool of the separation of high and low molecularweight subunits[J].Cereal Chem.1994,71:238-242
Mdiaz Zorita, Grosso G A.. Effect of soil texture, organic carbon and water retention on the compactability of soils from the Argentinean pampas.Soil & Tillage Research, 2000, (54): 121-126
Nakamura Y. Some properties of the starch debranching enzymes and their possible role in amylopectin biosynthesis. Plant Sci, 1996, 121: 1-18
Nakamura T, Vrinten P, Hayakawa K, Ikeda J. Characterization of a Granule-Bound starch synthase isoform found in the pericarp of wheat. Plant Physiol, 1998, 118:451-459
Nakamura Y, Umemoto T, Takahata Y, Komae K, Amano E, Satoh H. Changes in structure of starch and enzyme activities affected by sugary mutations in developing rice endosperm: possible role of starch debranching enzyme(R-enzyme) in amylopectin biosynthesis.Physiol Plant,1996b,97:491-498
Nakamura Y, Yuki K, Park S Y. Carbohydrate metabolism in the developing endosperm of rice grains. Plant Cell Physiol, 1989, 30: 833-839
Nicolas, M E, Turner, M C. The use of chemical dessiccants and senescing agents to selectwheat lines maintaining stable grain size during postanthesis drought. Field Crops Res, 1993.31, 155-171
Orth R.A.and Bushuk W.A comparative study of the proteins of wheats of diverse baking qualities [J]. Cereal Chemistry.1972,49:268-275
Osborne T.B. The Proteins of the Wheat Kernel [M].Carnegie Inst Washington Publ.1907, 84-119
Payne, P.I, and K.G. Corfield. 1979. Subunit composition of wheat glutenin proteins Iisolated by gel filtration in a dissociating medium. Planta. 145: 83-88.
Payne P.I.,Law C.N.and Mudd E.E.Control by homologous group 1 chromosomes of the high-molecular-weight subunits of glutenin,a major protein of wheat endosperm[J].Theoretical and Applied Genetics.1980,58:113-120
Payne P.I.,Corfield K.G.,Holt L.M.,etal.Correlations between the heritance of certain high-molecular-weight subunits of glutenin and bread making quality in progenies of six crosses of bread wheat[J].Journal of the Sciecne of Food and Agriculture.1981,32:51-60
Payne P.I.,Holt L.M.and Worland A.G.Strueture and genetical studies on the High-molecular-weight subunits of glutenin:III.Telocentric mapping of the subunit genes on the long arm of the homoeologous group 1 chromosomes[J].Thero,Appl.Genet.,1982,63:129-138Panozzo J F, Eagles H A, Wootton M. Changes in protein composition during grain development in wheat. Australian Journal of Agricultural Research, 2001, 52:485-493.
Payne P.I.,Seekings J.A.,Worland A.J.,Jarvis M.G.and Holt L.M.Allelic variation of glutenin subunits and gliadins and its effect on breadmaking quality in wheat:analysis of F5 progeny from Chinese Spring×ChineseSpring(Hope 1A) [J]. Journal of Cereal Science. 1987,6:103-118
Pan D, Nelson O E A debranching enzyme deficiency in endosperm of the sugary-1mutants of maize.Plant Physiol,984, 74: 324-29
Panozzo J F, Eagles H A, Wootton M. Changes in protein composition during grain development in wheat. Australian Journal of Agricultural Research, 2001, 52:485-493. Paul W U, Thomas C K. Soil compaction and root growth: A Review. Agronomy Journal, 1994, 86:759-766
Papakosta, D K. Phosphorus accumulation and translocation in wheat as affected by cultivar and nitrogen fertilization. J. Agron. Crop Sci. 1994, 173: 260-270
Pechanek, U., A. Karger, S. Gr?ger, B. Charvat, G. Sch?ggl, and T. Lelley. 1997. Effect of N fertilization on quantity of flour protein components, dough properties and breadmakingquality of wheat. Cereal Chem. 74(6): 800-805.
Peterson D G, Fulcher R G. Variation in Minnesota HRS wheats: starch granule size distribution. Food Research International, 2001, 34(4):357-363
P L de Fraitas, Zobel R W, Snyder V A. Corn root growth in soil columns with artificially constructed aggregates. Crop Science, 1999, 39: 735-730
Pritchard P E. The glutenin fraction(gel-protein) of wheat protein-a new tool in the prediction of baking quality. Aspects of applied biology,1993,36:75-84
Przulj N, Momcilovic V. Genetic variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley I. Dry matter translocation. Eur. J. Agron, 2001, 15: 241-254.
Raun W R, John G V. Soil-plant buffering of inorganic nitrogen in continous winter wheat[J]. Agron.J.1995, 87: 827-834.
Rogers W J, Payne, P I, Seekings J A, Sayers E J. Effect of bread-making quality of x-type high molecular weight subunits of glutenin. J.Cereal Sci,1991,14;209-221
Sun, H., D.N. Yao, B.Y. Li, G.T. Liu, and S.Z. Zhang. 1998. Correlation between content of glutenin macropolymer (GMP) in wheat and baking quality. Journal of the Chinese Cereals and Oils Association. 13(6): 13-16. (in Chinese)
Southan, M., and F. MacRitchie. 1999. Molecular weight distribution of wheat proteins. Cereal Chem. 76: 827-836.
Song, J.M., A.F. Liu, X.Y. Wu, J.J. Liu, Z.D. Zhao, and G.T. Liu. 2003. Composition and content of high-molecular-glutenin subunits and their relations with wheat quality. Scientia Agricultura Sinica. 36 (2): 128-133. (in Chinese)
Shewry P R, Halford N G, Tatham A S. High molecular weight subunits of wheat glutenin. J.Cereal Sci,1992,15:105-120
Singh H, MacRitchi F. Application of polymer scienceto properties of gluten. Journal of cereal science, 2001, 33:231-243
Singh N K, Shepherd K W, Cornish G B. A simplified SDS-PAGE procedure for separating LMWsubunits of glutenin. J.Cereal Sci,1991,14:203-208
Stephen P. Baenziger R. Clements L, McIntosh M S, et al. Effect of Cultivar,Environment, and Their Interaction and Stability Analyses on Milling and Baking Quality of Soft Red Winter Wheat[J]. Crop Sci, 1985, 25:5-8
Santiveri F, Royo C, Romagosa I. Growth and yield responses of spring and winter triticale cultivated under Mediterranean conditions. Eur.J. Agron, 2004, 20: 281-292.
Schnyder H. The role of carbohydrate storage and redistribution in the source-sink relations ofwheat and barley during grain filling-a review.New Phytol. 1993, 123: 233-245.
Shewry P R, Halford N G, Tatham A S, Popineau Y, Lafiandra D, Belton P S. The high molecular weight subunits of wheat glutenin and their role in determining wheat processing properties. Advances in Food and Nutrition Research, 2003, 45: 219-302.
Skerrit J H, Hac L, Bekes F. Depolymerization of the glutenin macropolymer during dough mixing. I. Changes in levels, molecular weight distribution, and overall composition. Cereal Chemistry, 1999, 76, 395-401.
Sutton K H, Larsen N G. Morgenstern M P, et al. Differing effects of mechanical dough development and sheeting development methods on aggregated glutenin proteins.Cereal Chemistry, 2003, 80: 707-711.
Shewry P R, Halford N G, Tatham A S. The classification and nomenclature of wheat gluten protenins[J ] . Journal of Cereal Science ,1986 ,9 :97 - 106.
Veraverbeke, W.S., and J.A. Delcour. 2002. Wheat protein composition and properties of wheat glutenin in relation to breadmaking functionality. Crit Rev Food Sci. 42: 179-208.
Wieser, H., S. Antes, and W. Seilmeier. 1998. Quantitative determination of gluten protein types in wheat flour by reversed-phase high-performance liquid chromatography. Cereal Chem. 75(5): 644-650.
Wieser, H., and G. Zimmermann. 2000. Importance of amounts and proportions of high molecular weight subunits of glutenin for wheat quality. Eur Food Res Technol. 210: 324-330.
Weegels, P.L., A.M. Pijpekamp, A. Graveland, R.J. Hamer, and J.D. Schofield. 1996. Depolymerisation and re-polymerisation of wheat glutenin during dough processing. I. Relationships between glutenin macropolymer content and quality parameters. J Cereal Sci. 23: 103-111.
Weegels, P.L., R.J. Hamer, and J.D. Schofield. 1995. RP–HPLC and capillary electrophoresis of subunits from glutenin isolated by SDS and osborne fractionation. J Cereal Sci. 22: 211-224.
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 under non-flooded and mulched condition. Field Crops Research, 2007, 101: 53-61
Yamamori M, Nakamura T, Kuroda A. Variations in the content of starch-granule bound protein among several Japanese cultivars of common wheat (Triticum aestivum L.). Euphytica, 1992, 64: 215-219
Zeeman S C, Tiessen A, Pilling E, Kato K L, Donald A M, Simth A M. Starch synthesis inArabidopsis. granule synthesis, composition and structure. Plant Physiology, 2002, 129: 516-529
Zeng Z R, King R W. Regulation of grain number in wheat: changes in endogenous levels of abscisic acid. Aust. J. Plant Physiol, 1986,13:347-352
Zhao X C, Shap P J. An improved 1D SDS-PAGE method for the identification of three bread wheat waxy protein. J, of Cereal Science,1996,23:191-193
Yuwei Qian,Ken Preston, Oleg Krokhin,Jean Mellish, and Werner Ens.Characterization of Wheat Gluten Proteins by HPLC and MALDI TOF Mass Spectrometry[J].Journal of American Society for Mass Spectrometry.2008,19:1542–1550
Yue, H.W., W.N. Tan, D. Jiang, T.B. Dai, Q. Jing, and W.X. Cao. 2007. Effects of post anthesis drought and waterlogging on contents of high molecular weight glutenin subunits and glutenin macropolymer in wheat grain. Acta Agronomica Sinica. 33(11): 1845-1849. (in Chinese)
Zhu, J., K. Khan, S. Huang, and O’Brien L. 1999. Allelic variation at Glu-D1 locus for high molecular weight (HMW) glutenin subunits: Quantification by multistacking SDS-PAGE of wheat grown under N fertilization. Cereal Chem. 76(6): 915-919.
Zhang, J.R., Y. Liu, G. Lin, and G.Y. He. 2006. Structure of wheat high molecular weight glutenin subunits and their role in determining processing properties. China Biotechnology, 26(8): 103-110. (in Chinese)
蔡瑞国,尹燕枰,张敏,戴忠民,严美玲,付国占,贺明荣,王振林.氮素水平对藁城8901和山农1391籽粒品质的调控效应.作物学报, 2007, 33(2): 304-310
邓志英,田纪春,刘现鹏.不同高分子量谷蛋白亚基组合的小麦籽粒蛋白组分及其谷蛋白大聚合体的积累规律[J].作物学报,2004,30(5):481-486
邓志英,田纪春,张永祥,王延训,孙国兴,盛峰.冬小麦高低分子量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): 308-3
何照范.粮油籽粒品质及其分析技术.北京:中国农业出版社, 1985, 144-150
贾光锋,范丽霞,王金水.小麦面筋蛋白结构,功能性及应用.粮食加工, 2004, (2):11-13
姜鸿明,余松烈,于振文,赵倩,邱化蛟,丁晓义.小麦谷蛋白聚合作用对增施氮肥的反应.麦类作物学报, 2003, 23(2): 43-46.
姜东,于振文,李永庚.施氮水平对高产小麦蔗糖含量和光合产物分配及籽粒淀粉积累的影响.中国农业科学, 2002, 35(2): 157-162
李保云,等.小麦高分子量谷蛋白亚基与小麦品质性状关系的研究[J].作物学报.2000, 26(3):322-326
李建敏,王振林,高荣岐,李圣福,蔡瑞国,闫素辉,于安玲,尹燕枰.强、弱筋小麦籽粒形成期蔗糖、淀粉合成相关酶活性及其与氮代谢的关系.作物学报, 2008, 34: 1019-1026.
李青常,王振林,张艳,刘霞,石书兵.施氮水平对小麦面条加工品质的影响.中国农业科学, 2005, 38(2): 420-424
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
刘宝龙,张怀刚.弱筋小麦品质高分子量谷蛋白亚基组成分析.西北农业学报, 2007, 16(2): 19-23.
刘建军,何中虎,赵振东,刘爱峰,宋建, R J Pena.小麦品质与干白面条品质参数关系的研究.作物学报, 2002, 28(6): 738-742
刘振兴,杨振华,邱孝煊,林增泉.肥料增产贡献率及其对土壤有机质的影响.植物营养与肥料学报,1994,(1):19-26
马宗斌,何建国,王小纯,马新明.氮素形态对两个小麦品种籽粒内源激素含量的影响.中国农业科学,2008,41,(1):63-69
马传喜,等.小麦胚乳蛋白质组成及高分子量麦谷蛋白亚基与烘烤品质的关系[J].作物学报.1993,19(16):562-566
马元喜.不同土壤小麦根系生长动态研究.作物学报, 1987,13(1):37-44
盛靖,郭文善,胡宏等.小麦淀粉合成关键酶活性及其与淀粉积累的关系.扬州大学学报(农业与生命科学版),2003,24(3):49-53
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作物学报, 2005, 31(4): 431-437
潘庆民,于振文,王月福等.追氮时期对小麦旗叶中蔗糖合成与籽粒中蔗糖降解的影响.中国农业科学,2002,35(7):771-776
潘庆民,于振文,王月福.小麦开花后旗叶中蔗糖合成与籽粒中蔗糖降解.植物生理与分子生物学学报. 2002b, 28(3):235-240.
师俊玲,魏益民.小麦蛋白质和淀粉与面条品质性状关系的研究进展.郑州粮食学院学报, 1998, 13(3): 1-5
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.
石书兵,马林,石庆华,刘霞,陈乐梅,刘建喜,王振林.不同施氮时期对冬小麦子粒蛋白质组分及其动态变化的影响.植物营养与肥料学报, 2005, 11(4): 456-46
王海波,庞惠玲,刘志宏.小麦谷蛋白研究方法的比较分析[J].中国农学通报. 2006, 22(7): 125 - 129
王瑞,等。一些优质小麦及其杂种后代高分子量谷蛋白亚基组成与面包品质之关系[J].西北农业学报.1995,4(4):25-30
王晓英,贺明荣,李飞,等.水氮耦合对强筋冬小麦子粒蛋白质和淀粉品质的影响.植物营养与肥料学报, 2007, 13(3): 361-367
王旭东,于振文,石玉等.磷对小麦旗叶氮代谢有关酶活性和籽粒蛋白质含量的影响.作物学报,2006,32(3):339-344
王旭东,于振文,王东.钾对小麦茎和叶鞘碳水化合物含量及籽粒淀粉积累的影响.植物营养与肥料学报,2003,9(1):57-62
王月福,于振文,李尚霞.氮素营养水平对小麦开花后碳素同化、运转和产量的影响.麦类作物学报, 2002, 22(2): 55-59
王月福,于振文,李尚霞,等.施氮量对小麦籽粒蛋白质组分含量及加工品质的影响[J].中国农业科学.2002,35(9):1071-1078
闫素辉,王振林,戴忠民,李文阳,付国占,贺明荣,尹燕枰.两个直链淀粉含量不同的小麦品种籽粒淀粉合成酶活性与淀粉积累特征的比较.作物学报, 2007, 33: 84-89.
闫素辉,尹燕枰,李文阳,李勇,等.密穂与疏穂型小麦强、弱势籽粒淀粉积累及库强度的比较.中国农业科学, 2009, 42(8): 2706-2715
闫素辉,尹燕枰,李文阳,李勇,梁太波,等.花后高温对不同耐热性小麦品种籽粒淀粉形成的影响.生态学报, 2008, 28(12): 6138-6147
晏月明,刘广田,Prodanovie S.小麦谷蛋白亚基的凝胶电泳分离及其品种鉴定[J].中国粮油学报.1998,13:1-5
姚大年,李保云,梁荣奇,刘广田.基因型和环境对小麦品种淀粉性状及面条品质的影响.中国农业大学学报,2000, 5(1): 63-68
尹静,胡尚连,肖佳雷,李文雄.不同形态氮肥对春小麦品种籽粒淀粉及其组分的调节效应.作物学报, 2006, 32(9): 1294-1300
于振文.优质专用小麦品种及栽培.北京:中国农业出版社. 2001
赵广才,常旭虹,刘利华,等.施氮量对不同强筋小麦产量和加工品质的影响[J].作物学报.2006,32(5):723—727
赵广才,张艳,刘利华,等.不同施肥处理对冬小麦产量、蛋白组分和加工品质的影响[J].作物学报.1993,19(6):562—567
赵俊晔,于振文.施氮量对小麦强势和弱势子粒氮素代谢及蛋白质合成的影响[J].中国农业科学.2005,38(8):1547—1554
赵友梅,等.高分子量麦谷蛋白亚基的SDS-PAGE图谱在小麦品质研究中的应用[J].作物学报.1990,16(3):208-218
张定一,张永清,杨武德,等.施氮量对不同小麦品种高分子量麦谷蛋白亚基表达量及品质影响的研究[J].植物营养与肥料学报.2008,14(2):235—241
赵和,等.小麦高分子量麦谷蛋白亚基遗传变异及其与品质和其他农艺性状关系的研究[J].作物学报.1994,20(l):67-75
张春庆,李晴祺.影响普通小麦加工馒头质量的主要品质性状的研究.中国农业科学, 1993, 26(2): 39-46
张金锐,刘勇,林刚,何光源.小麦高分子量谷蛋白亚基序列及其结构与加工品质的关系.中国生物工程杂志, 2006, 26(8): 103-110
张军,张洪程,戴其根,等.追施氮肥时期对淮南中筋小麦品质的影响.扬州大学学报(农业与生命科学版), 2003, 24(2): 44-48
张平平,张勇,夏先春,等.小麦贮藏蛋白反相高效液相色谱分析体系研究[J].中国农业科学. 2007,40(5):1002-1009.
Alessandro M, Laura E, Marco M. Post-anthesis accumulation and remobilization of dry matter, nitrogen and phosphorus in durum wheat as affected by soil type. Europ.J.Agronomy, 2007, 26:179-186
Andersm E C. Corn root growth and distribution as influenced by tillage and nitrogen fertilizer. Agronomy Journal, 1987, 79:544-559
Ayoub, M., S. Guertin, J. Fregeau-Reid, and D.L. Smith. 1994. Nitrogen fertilizer effect on breadmaking quality of hard red spring wheat in eastern Canada. Crop Sci. 34:1346–1352.
Ayoub, M., S. Guertin, and D.L. Smith. 1995. Nitrogen fertilizer rate and timing effect on bread wheat protein in eastern Canada. Crop Sci. 174:337–349.
Bae J M, Liu J R. Molecular cloning and characterization of two novel isoforms of the small subunit of ADPglueose pyrophosphorylase from sweet potato. Mol. Gen. Genet, 1997, 254: 179-185
Ball S, Guan H P, James M, et al. From glycogen to amylopectin: a model for the biogenesis of the starch granule. Cell, 1996, 86: 349-352
Batey I L, Curtin B M, Moore S A. Optimization of Rapid Visco Analyser test conditions for predicting Asian noodle quality. Cereal Chem., 1997,74(4): 497-501
Barneix A J, Guitman M R. Leaf regulation of the nitrogen concentration in the grain of wheat plants. Journal of Experimental Botany,1993,44(1):607-612
Bengough A G, Young I M. Root elongation of seeding peas through layered soil of different penetration resistances. Plant and Soil, 1993,149:129-139
Bernd Steingrobe, Harald Schmid, Reinhold Gutser, Norbert Claassen. Root production and root mortality of winter wheat grown on sandy and loamy soils in different farming systems. Biol Fertil Soils, 2001, 33:331-339
Behdaie A E, Hall G D. Farquhar Water-use efficiency and carbon Isotope Discrimination inWheat. Crop Science, 1991, 31: 1282-1288.
Blechl AE, Anderson OD (1996) Expression of a novel highmolecular-weight gluteninsubunit gene in transgenic wheat. Nat Biotechnol 14:875–879
Blechl AE, Le HQ, Anderson OD (1998) Engineering changes in wheat flour by genetic transformation. J Plant Physiol 152:703–707
Blechl A, Lin J, Nguyen S, Chan R, Anderson OD, Dupont FM (2007) Transgenic wheats with elevated levels of Dx5 and/or Dy10 high-molecular-weight glutenin subunits yield doughs with increased mixing strength and tolerance. J Cereal Sci 45:172–183
Bietz J.A. Analysis of wheat gluten proteins by high-performance liquid chromatography[J]. I. Baker’s Dig. 1984, 58: 15-32
Bietz J.A.Separation of cereal proteins by reversed-phase high-performance liquid chromatography [J]. Journal of Chromatography. 1983, 255: 219-238
Blumenthal C, Bekes F, Gras P W, et al. Identification of wheat genotypes tolerant to the effects of heat stress on grain quality. Cereal Chem., 1995, 72: 539-544
Branlard G.and Dardevet M.Diversity of grain protein and bread wheat quality.II.Correlation between high molecular weight subunits of glutenin and flour quality characteristics [J]. Journal of Cereal Science.1985,3:345-354
Braun H J, Payne T S, Morgounov A I, van Ginkel M, Rajaram S. The challenge: one billion tons of wheat by 2020. In: Slinkard A E (ed) Proceeding of the 9th International Wheat Genetics Symposium. Saskatchewan: University Extension Press, 1998, 33-40
Bushuk W. Wheat breeding for end-product use. Euphytica, 1998, 100(1-3): 137-145
Buttrose B L. Ultrastructure of the developing wheat endosperm. Aust. J. Biol. Sci., 1963, 16: 305-310
Butow, B.J., B.W. Ma, K.R. Gale, G.B. Cornish, L. Rampling, O. Larroque, M.K. Morrell, and F. Bekes. 2003. Molecular discrimination of Bx7 alleles demonstrates that a highly expressed high-molecular-weight glutenin allele has a major impact on wheat flour dough strength. Theor Appl Genet. 107: 1524-1532
Carceller, J.L., and T. Aussenac. 2001. SDS-insoluble glutenin polymer formation in developing grains of hexaploid wheat: the role of the ratio of high to low molecular weight glutenin subunits and drying rate during ripening. Aust. J. Plant Physiol. 28: 193-201
Clyde, D., L. George, N. Hamid, M. Finlay, and J.H. Rob. 2005. Heat stress and genotype affect the glutenin particles of the glutenin macropolymer-gel fraction. J Cereal Sci. 42: 69-80.
Chen Y, Yuan L P, Wang X H, Zhang D Y, Chen J, Deng Q Y, Zhao B R, Xu D Q. Relationship between grain yield and leaf photosynthetic rate in super hybid rice. J. Plant Physiol. Mol. Biol., 2007, 33(3): 235-243
Cressey P J, Campbell W P, Wrigley C W. Statistical correlation between quality attributes and grain protein composition for advanced lines of crossbred wheat. Cereal Chemistry,1987,64(4):299-301
COL ETTE LARRE , YVES NICOLAS ,CLAUDE DESSERME , et al . Preparative Separation of High and Low Molecular Weight Subunits of Glutenin from Wheat [J ] . Journal of Cereal Science ,1997 ,25 :143 - 150.
Crosbie G B. The relationship between starch swelling properties, paste viscosity and boiled noodle quality in wheat flour. Journal of Cereal Science, 1991, 13:145-150
Dai Z M, Yin Y P, Zhang M, Li W Y, Yan S H, Cai R G, Wang Z L. Starch granule size distribution in wheat grains under irrigated and rainfed conditions. Acta Agron. Sin. 2008, 34:Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2003a. Glutenin macropolymer: a gel formed by glutenin particles. J Cereal Sci. 37: 1-7.
Daniel, C., and E.Trib?. 2002. Changes in wheat protein aggregation during grain development: effects of temperatures and water stress. Eur J Agron. 16: 1-12.
Deng, Z.Y., J.C. Tian, R.B. Hu, X.F. Zhou, and Y.X. Zhang. 2006. Effects of genotype and environment on wheat main quality characteristics. Acta Ecologica Sinica. 26(8): 2757-2763. (in Chinese)
Deng, Z., J. Tian, L. Zhao, Y. Zhang, and C. Sun. 2008. High temperature-induced changes in high molecular weight glutenin subunits of Chinese winter wheat and its influences on the texture of Chinese noodles. J Agron Crop Sci. 194: 262-269.
Denyer K. Identification of multiple isoforms of soluble and granule-bound starch synthesis in developing wheat endosperm, Planta, 1995, 196: 256-265
Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2003b. Understanding the link between GMP and dough: from glutenin particles in flour towards developed dough. J Cereal Sci. 38: 157-165.
Don, C., W. Lichtendonk, J.J. Plijter, and R.J. Hamer. 2005. The effect of mixing on glutenin particle properties: aggregation factors that affect gluten function in dough. J Cereal Sci. 41: 69-83.
Don, C., G. Mannb, F. Bekesb, and R.J. Hamer. 2006. HMW-GS effect the properties of glutenin particles in GMP and thus flour quality. J Cereal Sci. 44: 127-136.
Douglas C D, KuoT M, Felker F C. Enzymes of sucrose and hexose metabolism indeveloping kernels of two inbreds of maize. Plant Physiol, 1988, 86: 1013-1019
Dreccer M F, Grashoff C, Rabbinge R. Source-sink ration in barley during grain filling: effects on senescence and grain protein concentration. Field crops research, 1997,49: 269-277
Dupont, F.M., and S.B. Altenbach. 2003. Molecular and biochemical impacts of environmental factors on wheat grain development and protein synthesis. J Cereal Sci. 38: 133-146.
DuPont,F.M.,Chan,R.,Lopez,R.,Vensel,W.H.Sequential extraction and quantitative recovery of gliadins,glutenins,and other proteins from small samples of wheat flour.[J].Journal of Agricultural and Food Chemistry.2005,53:1575-1584.
DuPont,F.M.,Hurkman,W.J.,Hurkman,W.J.,Vensel,W.H.,Chan,R.etal.Differential accumulation of sulfur-rich and sulfur-poor wheat flour proteins is affected by temperature and mineral nutrition during grain development.[J]Journal of Cereal Science. 2006b, 44: 101-112.
DuPont, F.M., R. Chan, and R. Lopez. 2007. Molar fractions of high-molecular-weight glutenin subunits are stable when wheat is grown under various mineral nutrition and temperature regimens. J Cereal Sci. 45: 134-139.
Eliasson A C, Karlsson R. Gelatinization properties of different size classes of wheat starch granules measured with differential scanning calorimetry. Starch, 1983, 35: 130-133.
Endo S, Karibe S, Okada K, Nagao S. Factors affecting gelatinization properties of wheat starch. Nippon Shikuhin Kogyo Gakkaishi, 1988, 35: 7-14
Eugenio L, Coelho, Dan D. Root distribtution and water uptake patterns of corn under surface and subsurface drip irrigation. Plant and soil,1999,206:123-126
Eva Johansson, Maria Luisa prieto-Linde,et al.Effects of Wheat Cultivar and Nitrogen Application on Storage Protein Composition and Breadmaking Quality, Cereal Chemists, 2001, 78(1):19-25.
Evers A D. Scanning electron microscopy of wheat starch.Ⅲ.Granule development in the endosperm. Starch, 1971, 23: 157-160
Evers A.D, Lindley J. The particle-size distribution in wheat endosperm starch, J. Sci. Food Agric., 1977, 28: 98-102
Fernando D, Francisco J C. Characterization of the endoprotease appearing during wheat grain filling. Plant physiol, 1996, 112:1211-1217
Fischer B., Grossmann J., Roth V., Gruissem W., Baginsky S., Buhmann J.M. Semi-supervised LC/MS alignment for differential proteomics[J]. Bioinformatics. 2006,14: 132–140
Fisher D B, Gifford R M. Accumulation and conversion of sugars by developing wheat grains.VI, Gradients along the transport pathway from the peduncle to the endosperm cavity during grain filling[J]. Plant Physiol, 1986, 82: 1024-1030
Fu B X, Kovacs MIP, Wang C. A simple wheat flour swelling test. Cereal Chem, 1998, 75(4):566-567
Gebbing T, and Schnyder H. Pre-anthesis reserve utilization for protein and carbohydrate synthesis in grains of wheat. Plant Physiology, 1999, 121: 871-878
Ceoloni C, Jauhar PP (2006) Chromosome engineering of the durum wheat genome: Strategies and applications of potential breeding value. In: Singh RJ, Jauhar PP (eds) vol 2 Genetic Resources, Chromosome Engineering, and Crop Improvement: Cereals. CRC Press, Taylor & Francis group, pp 27-59
Ceoloni C, Ciaffi M, Lafiandra D, Giorgi B (1995) Chromosome engineering as a means of transferring 1D storage protein genes from common to durum wheat. In: Li ZS, Xin ZY (eds) Proceedings of the 8th International Wheat Genetics Symposium, 20–25 July 1993, Beijing, China. China Agricultural Scientech Press, Beijing, pp 159–163
Ceoloni C, Biagetti M, Ciaffi M, Forte P, Pasquini M (1996) Wheat chromosome engineering at the 4x level: the potential of different alien gene transfers into durum wheat. Euphytica 89:87–97
Goodman A M, Eends A P. The effects of soil bulk density on the morphology and anchorage mechanics of the root systems of sunflower and maize. Annals of Botany, 1999, 83:293-302
Gianibelli M C, Gupta R B, Lafiandra D, Margiotta B, Macritchie F. Polymorphism of high Mr glutenin subunits in Triticum tauschii: characterization by chromatography and electrophoretic methods. J Cereal Sci, 2001, 33(1): 39-52
Graveland, A., P. Bosveld, W.J. Lichtendonk, J.P. Marseille,J.H. E. Moonen, and A. Scheepstra. 1985. A model for the molecular structure of the glutenins from wheat flour. J Cereal Sci. 3(1): 1-16.
Gupta R B, Khan K, MacRitchie F. Biochemical basis of flour properties in bread wheatsⅠ. Effects of variation in the quantity and size distribution of polymeric protein. J Cereal Sci, 1993, 18(1): 23-41
Gupta, R.B., K. Khan, and F. Macritchie. 1993. Biochemical basis of flour properties in bread wheats. I. Effects of variation in the quantity and size distribution of polymeric protein. J Cereal Sci. 18(1): 23-41.
Gupta, R.B., Y. Popineau, J. Lefebvre, M. Cornec, J.G. Lawrence, and F. Macritchie. 1995. Biochemical basis of flour properties in bread wheats. II. Changes in polymeric protein formation and dough/gluten properties associated with the loss of low Mr or high Mr glutenin subunits. J Cereal Sci. 21: 103-116.
Herbert Wieser,Susanne Antes,Werner Seilmeier.Quantitative Determination of Gluten Protein Types in Wheat Flour by Reversed-Phase High-Performance Liquid Chromatography [J]. Cereal Chem.1998,75(5):644-650
Hou G.,Yamamoto Y.and Ng P.K.W.Relationship of glutenin subunits of selected U.S.soft wheat flours to rheological and baking properties[J].Cereal Chem.1996,73:358-363
Huang D.Y.and Khan K.Quantitative determination of high molecular weight glutenin subunit of hard red spring wheat by SDS-PAGE.II.Quantitative effects of individual subunits on breadmaking quality characteristics[J].Cereal Chem.1997b,74:786-790
Holm J, Bjorck I. Bioavailablity of starch in various wheat-based bread products: evaluation of metabolic responses in healthy subjects and rate and extent of in vitro starch digestion [J].American Journal of Clinical Nutrition,1992, 55(2):420-429
Hurkman W. J, McCue K F, S. Altenbach B, Effect of temperature on expression of genes encoding enzymes for starch biosynthesis in developing wheat endosperm, Plant Science 2003,164: 873-881
James M G, Robertson M G, Myers A M. Characterisation of the maize gene sugary, a determinant of starch composition in kernels. Plant Cell, 1995, 7: 417-429
James M G, Denyer K, Myers A M. Starch synthesis in the cereal endosperm. Current Opinion in Plant Biology, 2003, 6: 215-222
Jenner C F. Effects of exposure wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars. Immediate responses. Aust.J.Pant Physiol,1991,18:165-177
Johnson V V. Genetic advances in wheat protein quantity and composition, Pron.4th
Intern.Wheat Genet.Symp.Columbia, 1973.
Jiang, H.M., S.L. Yu, Z.W. Yu, Q. Zhao, H.J. Qiu, and X.Y. Ding. 2003. Response of wheat glutenin polymerization to increased N fertilization. Journal of Triticeae Crops. 23(2): 43-46. (in Chinese)
Jiang D, Cao W X, Dai T B, Jing Q. Activities of key enzymes for starch synthesis in relation to growth of superior and inferior grains on winter wheat spike. Plant Growth Regul., 2003, 41, 247-257
Jiang, D., H. Yue, B. Wollenweber, W. Tan, H. Mu, Y. Bo, T. Dai, Q. Jing, and W. Cao. 2009.Effects of post-anthesis drought and water logging on Aaccumulation of high-molecular-weight glutenin subunits and glutenin macropolymers content in wheat grain. J Agron Crop Sci. 195: 89-97.
K. Dong, C.Y. Hao, A.L. Wang, M.H. Cai, and Y.M. Yan. Characterization of HMW Glutenin Subunits in Bread and Tetraploid Wheats by Reversed-Phase High-Performance Liquid Chromatography[J]. Cereal Research Communications. 2009, 37(1):65–72
Konarev, V. G., Gavrilyuk, I. P., Gubareva, N. K., and Peneva, T. I. (1979). Seed proteins in genome analysis, cultivar identification, and documentation of cereal genetic resources: A review. Cereal Chem. 56(4):272.
Khelifi D.and Branlard G.The effect of HMW and nological quality of progeny from four crosses between poor breadmaking quality and strong wheat cultivars[J].Journal of Cereal Science.1992,16:195-209
Kim H.R., Bushuk W. Changes in some physicochemical properties of flour proteins due to partial reduction with dithiothreitol[J].Cereal Chem.1995, 72(5): 450-456
Kolster P., Krechting C.F., Van Gelder W.M.J. Quantification of individual high molecular weight glutenin subunits using SDS-PAGE and scanning densitometry[J]. Journal of Cereal Science.1992, 15:49-61
Konik C M, Mikkelsen L M, Moss R, and Gore P J. Relationships between physical starch properties and yellow alkaline noodle quality, Starch / staerke, 1994, 46: 292-299
Laemmli U.K.Cleavage of structural proteins during the assembly of the head of bacteriophage T4 [J].Natrue.1970, 227:680-685
Lafiandra D, Margiotta B, Colaprico G, Masci S, Roth MR, MacRitchie F (2000) Introduction of the D-genome related high- and low-Mr glutenin subunits into durum wheat and their effect on technological properties. In: Shewry PR, Tatham AS (eds) Wheat Gluten: Proceedings of the 7th International Workshop Gluten, 2–6 April 2000, Bristol, UK. The Royal Society of Chemistry, Cambridge, pp 51–54
Lawrence G.J.,MacRitchie F.and Wrigley C.W. Dough and baking quality of wheat lines deficient in glutenin subunits controlled by the Glu-A1,Glu-B1 and Glu-D1 loci[J].Journal of Cereal Science.1988,7:109-112
Lawrence G. J. The high-molecula-weight glutenin subunit composition of Australia wheat cultivars [J]. Aust. J. Agri. Res.1986,37:127-133
Larroque O.R., Bekes F., Wrigley C.W., Rathmell W.G. Analysis Luo C, Branlard G, Griffin B, et al. The effect of nitrogen and sulphur fertilization and their interaction with genotype on wheat glutenins and quality parameters[J].Journal of Cereal cience,2000,31:185-194.
Ley, I., 2005. Particle size analysis—evaluating laser diffraction systems in the light of ISO 13320-1, publication T-403, Beckman Coulter, High Wycombe,UK.
Lefebvre, J., Y. Popineau, G. Deshayes, and L. Lavenant. 2000. Temperature-Induced changes in the dynamic rheological behavior and size distribution of polymeric proteins for glutens from wheat near-isogenic lines differing in HMW glutenin subunit composition. Cereal Chem. 77(2): 193-201.
Liu, B.L., and H.G. Zhang. 2007. Analysis on the HMW-GS components of weak gluten wheat varieties. Acta Agriculturae Boreali-occidentalis Sinica. 16(2): 19-23. (in Chinese)
Liang, T.B., Y.P. Yin, R.G. Cai, S.H. Yan, W.Y. Li, Q.H. Geng, P. Wang, Y.H. Wu, Y. Li, and Z.L. Wang. 2008. HMW-GS accumulation and GMP size distribution in grains of shannong 12 grown in different soil conditions. Acta Agronomica Sinica. 34(12): 2160-2167. (in Chinese)
Lucow O.M.,Payne P.I.and Thachuk R.The HMW glutenin subunits compositions in Canadian wheat cultivars and their association with bread making quality[J].Journal of the Sciecne of Food and Agriculture.1989,46:451-460
Lookart G.Bean S.Ultrafast CE analysis of cereal storage proteins and its application to protein characterization and cultivar differentiation [J]. Agric.Food Chem. 2000, 48: 344-353
Mackay A D, Barber S A. Effect of nitrogen on root growth of two corn genotypes in the field. Agronomy Journal, 1986, 78: 699-703
Masc S, Egorov T A,Ronchi C,et al. Evidence for the presence of only one cysteine residue in theD-type low molecular weight subunits of wheat subunits of wheat glutenin[J].Journal of Cereal Science,1999, 29:17-25.
Masci, S., La?andra, D., Porceddu, E., Lew, E.J.-L., Tao, H.P. and Kasarda, D.D. (1993) D-glutenin Subunits: N-terminal Sequences and Evidence for the Presence of Cysteine' in Cereal Chem. 70, 581±585
Masci, S., Lew, E.J.L., La?andra, D., Porceddu, E. and Kasarda,D.D. (1995) Characterization of Low Molecular Weight Sub-units in Durum Wheat by Reversed-phase High-performance Liquid Chromatography and N-terminal Sequencing' in Cereal Chem. 72, 100±104
D'Ovidio, R., Simeone, M., Masci, S., Porceddu, E. and Kasarda,D.D. (1995) `Nucleotide Sequence of a g-type Glutenin Gene from a Durum Wheat: Correlation with a g-type Glutenin
Subunit From the Same Biotype' in Cereal Chem. 72, 443±449
Masoni A, Ercoli L, Mariotti M, Arduini I. Post-anthesis accumulation and remobilization of dry matter, nitrogen and phosphorus in durum wheat as affected by soil type. Eur J Agron, 2007, 26: 179-186
Materechera S A. Penetration of very strong soil by seeding roots of different species. Plant and soil, 1991, 135:31-41
Metakovsky EV (1991) Gliadin allele identification in common wheat. II. Catalogue of gliadin alleles in common wheat. J Genet Breed 45:325-343
Metakovsky EV, Akhmedov MG, Sozinov AA (1986) Genetic analysis of gliadin-encoding genes reveals gene clusters as well as single remote genes. Theor Appl Genet 73:278-285
Metakovsky EV, Ng PKW, Chernakov VM, Pogna NE, Bushuk W (1992) Gliadin alleles in Canadian Western Red Spring (CWRS) class of wheat eultivars: use of the two procedures of A-PAGE for gliadin separation. Genome (in press) Ng PKW, Bushu
Melas V ,Morel M H ,Autran J C. Simple and rapid methed for purifying low molecular weight glutenin from wheat [J ] . Cereal Chemistry ,1994 ,71 :234 - 237.
Mouille G, Maddelein M L, Ball S. Preamylopectin processing: A mandatory step for starch biosynthesis in plants. Plant Cell, 1996, 8: 1353-1366
Moonen C, Masoni A, Ercoli L, Mariotti M, Bonari E. Long-term changes in rainfall and temperature in Pisa, Italy. Agr. Med, 2001, 130: 11-22
Morel M.H.Acid polyaerylamide gel eleclrophoresis of wheat gluteins:a new tool of the separation of high and low molecularweight subunits[J].Cereal Chem.1994,71:238-242
Mdiaz Zorita, Grosso G A.. Effect of soil texture, organic carbon and water retention on the compactability of soils from the Argentinean pampas.Soil & Tillage Research, 2000, (54): 121-126
Nakamura Y. Some properties of the starch debranching enzymes and their possible role in amylopectin biosynthesis. Plant Sci, 1996, 121: 1-18
Nakamura T, Vrinten P, Hayakawa K, Ikeda J. Characterization of a Granule-Bound starch synthase isoform found in the pericarp of wheat. Plant Physiol, 1998, 118:451-459
Nakamura Y, Umemoto T, Takahata Y, Komae K, Amano E, Satoh H. Changes in structure of starch and enzyme activities affected by sugary mutations in developing rice endosperm: possible role of starch debranching enzyme(R-enzyme) in amylopectin biosynthesis.Physiol Plant,1996b,97:491-498
Nakamura Y, Yuki K, Park S Y. Carbohydrate metabolism in the developing endosperm of rice grains. Plant Cell Physiol, 1989, 30: 833-839
Nicolas, M E, Turner, M C. The use of chemical dessiccants and senescing agents to selectwheat lines maintaining stable grain size during postanthesis drought. Field Crops Res, 1993.31, 155-171
Orth R.A.and Bushuk W.A comparative study of the proteins of wheats of diverse baking qualities [J]. Cereal Chemistry.1972,49:268-275
Osborne T.B. The Proteins of the Wheat Kernel [M].Carnegie Inst Washington Publ.1907, 84-119
Payne, P.I, and K.G. Corfield. 1979. Subunit composition of wheat glutenin proteins Iisolated by gel filtration in a dissociating medium. Planta. 145: 83-88.
Payne P.I.,Law C.N.and Mudd E.E.Control by homologous group 1 chromosomes of the high-molecular-weight subunits of glutenin,a major protein of wheat endosperm[J].Theoretical and Applied Genetics.1980,58:113-120
Payne P.I.,Corfield K.G.,Holt L.M.,etal.Correlations between the heritance of certain high-molecular-weight subunits of glutenin and bread making quality in progenies of six crosses of bread wheat[J].Journal of the Sciecne of Food and Agriculture.1981,32:51-60
Payne P.I.,Holt L.M.and Worland A.G.Strueture and genetical studies on the High-molecular-weight subunits of glutenin:III.Telocentric mapping of the subunit genes on the long arm of the homoeologous group 1 chromosomes[J].Thero,Appl.Genet.,1982,63:129-138Panozzo J F, Eagles H A, Wootton M. Changes in protein composition during grain development in wheat. Australian Journal of Agricultural Research, 2001, 52:485-493.
Payne P.I.,Seekings J.A.,Worland A.J.,Jarvis M.G.and Holt L.M.Allelic variation of glutenin subunits and gliadins and its effect on breadmaking quality in wheat:analysis of F5 progeny from Chinese Spring×ChineseSpring(Hope 1A) [J]. Journal of Cereal Science. 1987,6:103-118
Pan D, Nelson O E A debranching enzyme deficiency in endosperm of the sugary-1mutants of maize.Plant Physiol,984, 74: 324-29
Panozzo J F, Eagles H A, Wootton M. Changes in protein composition during grain development in wheat. Australian Journal of Agricultural Research, 2001, 52:485-493. Paul W U, Thomas C K. Soil compaction and root growth: A Review. Agronomy Journal, 1994, 86:759-766
Papakosta, D K. Phosphorus accumulation and translocation in wheat as affected by cultivar and nitrogen fertilization. J. Agron. Crop Sci. 1994, 173: 260-270
Pechanek, U., A. Karger, S. Gr?ger, B. Charvat, G. Sch?ggl, and T. Lelley. 1997. Effect of N fertilization on quantity of flour protein components, dough properties and breadmakingquality of wheat. Cereal Chem. 74(6): 800-805.
Peterson D G, Fulcher R G. Variation in Minnesota HRS wheats: starch granule size distribution. Food Research International, 2001, 34(4):357-363
P L de Fraitas, Zobel R W, Snyder V A. Corn root growth in soil columns with artificially constructed aggregates. Crop Science, 1999, 39: 735-730
Pritchard P E. The glutenin fraction(gel-protein) of wheat protein-a new tool in the prediction of baking quality. Aspects of applied biology,1993,36:75-84
Przulj N, Momcilovic V. Genetic variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley I. Dry matter translocation. Eur. J. Agron, 2001, 15: 241-254.
Raun W R, John G V. Soil-plant buffering of inorganic nitrogen in continous winter wheat[J]. Agron.J.1995, 87: 827-834.
Rogers W J, Payne, P I, Seekings J A, Sayers E J. Effect of bread-making quality of x-type high molecular weight subunits of glutenin. J.Cereal Sci,1991,14;209-221
Sun, H., D.N. Yao, B.Y. Li, G.T. Liu, and S.Z. Zhang. 1998. Correlation between content of glutenin macropolymer (GMP) in wheat and baking quality. Journal of the Chinese Cereals and Oils Association. 13(6): 13-16. (in Chinese)
Southan, M., and F. MacRitchie. 1999. Molecular weight distribution of wheat proteins. Cereal Chem. 76: 827-836.
Song, J.M., A.F. Liu, X.Y. Wu, J.J. Liu, Z.D. Zhao, and G.T. Liu. 2003. Composition and content of high-molecular-glutenin subunits and their relations with wheat quality. Scientia Agricultura Sinica. 36 (2): 128-133. (in Chinese)
Shewry P R, Halford N G, Tatham A S. High molecular weight subunits of wheat glutenin. J.Cereal Sci,1992,15:105-120
Singh H, MacRitchi F. Application of polymer scienceto properties of gluten. Journal of cereal science, 2001, 33:231-243
Singh N K, Shepherd K W, Cornish G B. A simplified SDS-PAGE procedure for separating LMWsubunits of glutenin. J.Cereal Sci,1991,14:203-208
Stephen P. Baenziger R. Clements L, McIntosh M S, et al. Effect of Cultivar,Environment, and Their Interaction and Stability Analyses on Milling and Baking Quality of Soft Red Winter Wheat[J]. Crop Sci, 1985, 25:5-8
Santiveri F, Royo C, Romagosa I. Growth and yield responses of spring and winter triticale cultivated under Mediterranean conditions. Eur.J. Agron, 2004, 20: 281-292.
Schnyder H. The role of carbohydrate storage and redistribution in the source-sink relations ofwheat and barley during grain filling-a review.New Phytol. 1993, 123: 233-245.
Shewry P R, Halford N G, Tatham A S, Popineau Y, Lafiandra D, Belton P S. The high molecular weight subunits of wheat glutenin and their role in determining wheat processing properties. Advances in Food and Nutrition Research, 2003, 45: 219-302.
Skerrit J H, Hac L, Bekes F. Depolymerization of the glutenin macropolymer during dough mixing. I. Changes in levels, molecular weight distribution, and overall composition. Cereal Chemistry, 1999, 76, 395-401.
Sutton K H, Larsen N G. Morgenstern M P, et al. Differing effects of mechanical dough development and sheeting development methods on aggregated glutenin proteins.Cereal Chemistry, 2003, 80: 707-711.
Shewry P R, Halford N G, Tatham A S. The classification and nomenclature of wheat gluten protenins[J ] . Journal of Cereal Science ,1986 ,9 :97 - 106.
Veraverbeke, W.S., and J.A. Delcour. 2002. Wheat protein composition and properties of wheat glutenin in relation to breadmaking functionality. Crit Rev Food Sci. 42: 179-208.
Wieser, H., S. Antes, and W. Seilmeier. 1998. Quantitative determination of gluten protein types in wheat flour by reversed-phase high-performance liquid chromatography. Cereal Chem. 75(5): 644-650.
Wieser, H., and G. Zimmermann. 2000. Importance of amounts and proportions of high molecular weight subunits of glutenin for wheat quality. Eur Food Res Technol. 210: 324-330.
Weegels, P.L., A.M. Pijpekamp, A. Graveland, R.J. Hamer, and J.D. Schofield. 1996. Depolymerisation and re-polymerisation of wheat glutenin during dough processing. I. Relationships between glutenin macropolymer content and quality parameters. J Cereal Sci. 23: 103-111.
Weegels, P.L., R.J. Hamer, and J.D. Schofield. 1995. RP–HPLC and capillary electrophoresis of subunits from glutenin isolated by SDS and osborne fractionation. J Cereal Sci. 22: 211-224.
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 under non-flooded and mulched condition. Field Crops Research, 2007, 101: 53-61
Yamamori M, Nakamura T, Kuroda A. Variations in the content of starch-granule bound protein among several Japanese cultivars of common wheat (Triticum aestivum L.). Euphytica, 1992, 64: 215-219
Zeeman S C, Tiessen A, Pilling E, Kato K L, Donald A M, Simth A M. Starch synthesis inArabidopsis. granule synthesis, composition and structure. Plant Physiology, 2002, 129: 516-529
Zeng Z R, King R W. Regulation of grain number in wheat: changes in endogenous levels of abscisic acid. Aust. J. Plant Physiol, 1986,13:347-352
Zhao X C, Shap P J. An improved 1D SDS-PAGE method for the identification of three bread wheat waxy protein. J, of Cereal Science,1996,23:191-193
Yuwei Qian,Ken Preston, Oleg Krokhin,Jean Mellish, and Werner Ens.Characterization of Wheat Gluten Proteins by HPLC and MALDI TOF Mass Spectrometry[J].Journal of American Society for Mass Spectrometry.2008,19:1542–1550
Yue, H.W., W.N. Tan, D. Jiang, T.B. Dai, Q. Jing, and W.X. Cao. 2007. Effects of post anthesis drought and waterlogging on contents of high molecular weight glutenin subunits and glutenin macropolymer in wheat grain. Acta Agronomica Sinica. 33(11): 1845-1849. (in Chinese)
Zhu, J., K. Khan, S. Huang, and O’Brien L. 1999. Allelic variation at Glu-D1 locus for high molecular weight (HMW) glutenin subunits: Quantification by multistacking SDS-PAGE of wheat grown under N fertilization. Cereal Chem. 76(6): 915-919.
Zhang, J.R., Y. Liu, G. Lin, and G.Y. He. 2006. Structure of wheat high molecular weight glutenin subunits and their role in determining processing properties. China Biotechnology, 26(8): 103-110. (in Chinese)