Wheat arabinoxylans: Exploiting variation in amount and composition to develop enhanced varieties

Arabinoxylans (AX) are the major polymers of wheat grain cell walls. The content and the structure of AX polymers show large differences between tissues and between wheat cultivars that affect the end-use properties and nutritional quality of the grain. The development of new wheat cultivars with enhanced quality, therefore, requires methods to exploit this variation and it is essential to understand and modulate the mechanisms controlling the key events of cell-wall polymer synthesis.This paper summarises recent knowledge on the structure and physicochemical properties of AX including variation between cultivars and tissues, methods for analysis and screening, biosynthetic mechanisms and approaches to identifying key genes. This knowledge is essential to understand AX properties and defined possible targets for plant breeding.     

Cell wall deposition and metabolism in wheat grain

There is an amazing diversity of cell wall composition and cell wall polysaccharide structure in wheat grain that is only partly explained by the complexity of plant tissues and cell type functions present in this organ. This review presents the state of the art on the structure and diversity of cell wall polymers in mature wheat grain and recent knowledge on cell wall metabolism with a focus on development-associated changes in wheat grain cell wall polysaccharides and genes controlling their biosynthesis. The diversity of polysaccharide structure observed in endosperm cell walls is tentatively explained on the basis of polysaccharide properties and cell type function.     

Rheological characterisation of aqueous extracts of triticale grains and its relation to dietary fibre characteristics

Rheological properties of triticale extract are important determinants of the grain utility. Extracts of eight triticale cultivars and a wheat reference grown at two different locations i.e. Svalöv and Kölbäck were analysed for their viscoelastic properties. Extracts of triticale cultivars grown at Svalöv were significantly higher in elastic modulus (average 9.0 Pa), viscous modulus (average 12.7 Pa) and complex modulus (average 15.6 Pa) at 2.1 Hz than those grown at Kölbäck (average 3.1, 5.6 and 6.4 Pa, respectively). However phase angle was not affected by location, and all triticale extracts had phase angle >45°. Large inter-cultivar variation was also observed in viscoelastic properties of triticale extracts. Some of the triticale cultivars grown at Kölbäck (Dinaro, Cando, Talentro, and DED 145/02) and Svalöv (DED 145/02) were similar to wheat in their complex modulus values. When evaluating the contribution of arabinoxylan (AX) and β-glucan to rheological properties of triticale extracts by partial least square (PLS) regression, it was evident that β-glucan had little role and most of the contribution to viscoelastic properties came from AX. AX fine structure was more important in explaining variation in complex modulus of triticale extract than content and extractability, or molecular size.     

新疆小麦品种籽粒性状、磨粉品质及其关系的研究

为给新疆冬、春小麦品种籽粒性状与磨粉品质改良提供参考依据,以30个新疆冬小麦品种和36个新疆春小麦品种为试验材料,研究了新疆小麦品种籽粒性状、磨粉品质特性及其两者的关系。结果表明,新疆冬、春小麦品种籽粒性状基本相近,各有优劣,主要差异表现在硬度大小不同,冬小麦品种的硬度（47.1）小于春小麦品种（61.8）;冬小麦品种的出粉率和黄度分别高于和低于春小麦品种,说明冬小麦品种的磨粉品质优于春小麦品种。新疆本地育成冬小麦品种的籽粒硬度（51.8）小于春小麦品种（62.9）,出粉率高于春小麦品种,灰分、红度和黄度也高于春小麦品种。综合来看,新疆本地育成春小麦品种的籽粒性状略优于冬小麦品种,磨粉品质优于冬小麦品种。相关分析表明,新疆冬、春小麦籽粒硬度与破损淀粉率呈极显著正相关,与L*值呈极显著负相关;籽粒蛋白含量与面粉灰分呈显著正相关。说明籽粒硬度和籽粒蛋白含量与新疆小麦磨粉品质关系最为密切。总之,在新疆小麦籽粒性状与磨粉品质改良中,应以籽粒硬度和面粉色泽改良为重点,同时也要提高新疆小麦籽粒蛋白含量和出粉率。     

分子生物学技术在普通小麦谷蛋白研究中的应用

小麦谷蛋白是面筋的主要成分之一，对小麦食品的加工品质起着重要作用。本文从基因序列、分子结构、多态性、遗传转化、QTL研究和MAS等方面综述了国内外有关麦谷蛋白亚基的研究进展。这些研究结果表明，麦谷蛋白的等位基因变异十分丰富，多态性高．序列之间存在很高的同源性。麦谷蛋白的基因结构分为三部分：无重复结构的N-末端和C-末端以及中部重复区域，等位基因的变异主要由基因中部重复区域的序列大小、重复次数及该区域内DNA序列的插入或缺失所造成；Cys-残基的数目和位置影响麦谷蛋白聚合体内亚基间的相互作用，是影响亚基生化特性的重要因素；应用转基因技术已将HMW-GS基因(1Ax1、1Dx5和1Dy10)导入普通小麦中，有助于进行品质改良和麦谷蛋白结构与功能的深入研究。此外，对面筋强度性状的QTL分析和分子标记辅助育种也进行了阐述。     

Diagnostic DNA markers for quality traits in wheat

The mechanisms underlying some quality traits in wheat are now understood. Examples include the role of high and low molecular weight glutenins in contributing to strength and extensibility of wheat doughs, puroindolines that affect grain texture, and variation in granule-bound starch synthase that produces starches with altered amylose content and physical properties. This knowledge, coupled with the availability of the DNA sequences of various alleles of the genes encoding these proteins and the wide application of the polymerase chain reaction, has enabled the design of diagnostic DNA markers for these quality traits. Such markers are now being used by wheat breeders to select lines with the required quality attributes, without the need for the direct measurement of those traits in early generation screening. DNA markers may be implemented on leaf tissue from individual plants, for a number of independent traits, with results that are independent of environmental variation. The use of a common platform for all marker assays and the potential for multiplexing or parallel analysis of many different markers will further increase the efficiency and speed of the development of improved cultivars in the future. This review provides an overview of diagnostic DNA markers that are currently available for the selection of quality traits in wheat.     

Wheat seed storage proteins: Advances in molecular genetics,diversity and breeding applications

Wheat seed storage proteins, especially glutenins and gliadins, have unique functional properties giving rise to a wide array of food products for human consumption. The wheat seed storage proteins, however, are also the most common cause of food-related allergies and intolerances, and it has become crucially important to understand their composition, variation and functional properties and interface this knowledge with the grain handling industry as well as the breeders. This review focuses on advances in understanding the genetics and function of storage proteins and their application in wheat breeding programs. These include: (1) The development and validation of high-throughput molecular marker systems for defining the composition and variation of low molecular weight glutenin subunits (LMW-GS) genes and a summary of the more than 30 gene-specific markers for rapid screening in wheat breeding programs; (2) The identification of more than 100 alleles of storage proteins in wild species provide candidate genes for future quality improvement; (3) The documentation of quality effects of individual LMW-GS and HMW-GS for improving end-use quality; and (4) The analysis of α-gliadin genes on chromosomes 6A and 6D with non-toxic epitopes as potential targets to develop less toxic cultivars for people with celiac disease. Genomic and proteomic technologies that will continue to provide new tools for understanding variation and function of seed storage proteins in wheat are discussed.     

小麦籽粒淀粉理化特性与品质关系及其生理机制研究进展

淀粉作为籽粒主要组分,是小麦加工品质最重要的决定因素之一。淀粉的加工品质主要取决于直链与支链淀粉含量及其比值、淀粉分子精细结构和淀粉颗粒大小分布等理化特性。明确小麦籽粒不同部位淀粉理化特性、空间分布差异及其形成的调控机制,可深入阐释面粉品质性状差异形成的生理机制。本文从小麦籽粒淀粉理化特性空间分布、淀粉合成关键酶活性及其编码基因时空表达模式等角度综述了小麦籽粒淀粉理化特性空间分布特征形成的生理机制,以期为小麦品质形成机制与调优栽培技术的建立提供理论支持。     

硬质小麦籽粒阿拉伯木聚糖含量的遗传变异研究

为了更好地理解环境和基因型对小麦阿拉伯木聚糖含量的影响,为小麦品质改良提供科学依据,采用比色法测定了美国西部51个硬质小麦品种的阿拉伯木聚糖组分含量,这些小麦品种包括冬小麦和春小麦,分别被种植在3个不同的地点.结果表明,参试冬小麦品种水溶性和总阿拉伯木聚糖含量的变异范围分别为0.39%～0.81%和3.09%～4.04%,春小麦品种这两种木聚糖含量范围分别为0.48%～0.92%和3.94%～4.70%.小麦籽粒阿拉伯木聚糖含量的变异同时受品种、环境及其互作效应的显著影响,环境效应太于品种效应.小麦品种间阿拉伯木聚糖含量的变异主要表现在水溶性组分上.就不同组分而言,基因型对水溶性阿拉伯木聚糖组分的影响相对较大,环境则分别对冬小麦水不溶性、春小麦水溶性阿拉伯木聚糖组分有较大的影响.品种间和环境间阿拉伯木聚糖含量的变异,冬小麦大于春小麦;水溶性阿拉伯木聚糖含量的遗传型方差春小麦大于冬小麦.冬、春小麦的水溶性阿拉伯木聚糖含量的广义遗传力均大于95%,通过遗传育种的方法改良小麦阿拉伯木聚糖含量是可行的.冬小麦与春小麦阿拉伯木聚糖含量的平均值之间无明显差异.     

Wheat endosperm cell walls: Spatial heterogeneity of polysaccharide structure and composition using micro-scale enzymatic fingerprinting and FT-IR microspectroscopy

Micro-scale enzymatic fingerprinting and FT-IR microspectroscopy were used to investigate changes of polysaccharide structure and composition in cell walls from wheat endosperm. These methods were applied to transverse and longitudinal sections of wheat grains harvested at maturity and 270°D. Principal component analysis treatment of the data revealed marked differences in the (1,3)-(1,4)-beta-glucans (BG)/arabinoxylans (AX) ratio and in the structure of AX depending on both cell position in the grain and stage of development. Cell walls close to the germ were enriched in BG in both developing and mature grain. AX in developing grain were characterized by a higher proportion of di-substitution by arabinose as compared to mature grain AX. In addition, AX in mature grains exhibited a different structure depending on prismatic or central origin of cell walls in the grain. These results indicated a high spatial and temporal control of the biosynthesis of AX and BG in wheat endosperm.     

陕西关中小麦品种品质改良现状研究

本项目从１９９２～１９９７ 年,以陕西关中地区近３０ 年来选育、推广的小麦品种、品系为材料,按照小麦品种田间比较试验的要求,对品种的品质性状、新品种的品质改良效果、品质性状的年际变化、品种品质分类等问题,进行了较为系统的研究。结果认为,关中地区选育和推广的小麦品种,容重较高,籽粒硬度适中或偏高,蛋白质含量和湿面筋含量较高,但沉淀值较低;多数品种的面团稳定性差,软化度较大。原因主要是Ｄ组染色体上缺少具有５＋ １０ 带的高分子量谷蛋白亚基。同一品种在同一地点不同年际间某些品质性状表现出较大差异。近年来育成的荔垦２ 号、陕优２２５、小偃６ 号、早优５０４、高优５０３、长武１３４、陕１６０、陕农７８０１ 等品种可以代表陕西关中地区优质小麦品种的品质水平,在全国也有一定的影响。但一些抗病、高产、早熟且推广面积较大的小麦新品种的品质水平有降低趋势。这一问题应引起育种工作者的高度重视。优质小麦生产是一项系统工程,除了遗传育种、栽培问题以外,粮食流通体制、检验标准、分收分贮、加工技术及食品制作等都会影响优质小麦在生产上的发展。     

Wheat endosperm cell walls: Spatial heterogeneity of polysaccharide structure and composition using micro-scale enzymatic fingerprinting and FT-IR microspectroscopy

Micro-scale enzymatic fingerprinting and FT-IR microspectroscopy were used to investigate changes of polysaccharide structure and composition in cell walls from wheat endosperm. These methods were applied to transverse and longitudinal sections of wheat grains harvested at maturity and 270°D. Principal component analysis treatment of the data revealed marked differences in the (1,3)-(1,4)-beta-glucans (BG)/arabinoxylans (AX) ratio and in the structure of AX depending on both cell position in the grain and stage of development. Cell walls close to the germ were enriched in BG in both developing and mature grain. AX in developing grain were characterized by a higher proportion of di-substitution by arabinose as compared to mature grain AX. In addition, AX in mature grains exhibited a different structure depending on prismatic or central origin of cell walls in the grain. These results indicated a high spatial and temporal control of the biosynthesis of AX and BG in wheat endosperm.     

长江上游小麦新品种(系)品质分析

为了解长江上游冬麦区小麦品种的品质现状及其基因型,以2012-2018年长江上游国家级品种试验109个小麦品种(系)在试验中的品质数据为基础,分析了品种(系)的籽粒分级、品质分析样品的混合点率、粗蛋白质含量、湿面筋含量、吸水量和稳定时间等性状;利用KASP标记检测了这些品种(系)的部分高、低分子量谷蛋白亚基、穗发芽抗性基因、粒重及籽粒大小基因、籽粒硬度基因和小麦-黑麦1BL·1RS易位等20个位点的基因型。发现穗发芽抗性差、蛋白质数量和质量指标不均衡是影响长江上游小麦品质的主要因素,该麦区的品质改良应以提高品种穗发芽抗性和改良稳定时间为主要目标;这些品种(系)在20个位点基因型的差别不足以解释其品质的差异。     

Environmental modification of hard red winter wheat flour protein composition

The intrinsic processing quality of wheat (Triticum aestivum L.) cultivars is modified significantly by cultural conditions and climate. In an attempt to understand the biochemical basis of such variation, environmental modification of flour protein content and composition was measured. Thirty hard red winter wheat cultivars and experimental lines were grown at 17 Nebraska environments during 1990 and 1991. Environmental conditions, including grain filling duration, temperature and relative humidity during grain filling, were monitored. Grain yield and test weight also were determined as environmental indicators. Significant linear correlations between flour protein content, as measured by near-infrared spectroscopy, were observed only with the duration of grain filling. Protein quality, as measured by SDS sedimentation volumes and size-exclusion high-performance liquid chromatography, was highly influenced by the frequency of high temperatures during grain filling and by the relative humidity. Observed ranges in genotypic responses (variance) at locations also were altered by environmental factors. Optimal protein quality, as determined by SDS sedimentation volumes, was observed with exposure to less than 90 h of temperature greater than 32°C during grain filling. Protein quality declined with exposure to a greater number of hours of elevated temperature.     

不同小麦品种抗性淀粉含量及其与其他淀粉品质性状的关系

为了解不同小麦品种抗性淀粉含量的差异及其与其他淀粉品质性状的关系,以30个小麦品种为试验材料,分别在河南省7个地点种植,对面粉中抗性淀粉含量、淀粉组分、淀粉糊化特性、淀粉粒度分布等进行了分析。结果表明,所有被测品质性状均受环境、基因型影响,且多数指标的环境效应大于基因型效应。7个地点间比较,抗性淀粉含量及淀粉品质性状的综合表现在开封点和原阳点均优于其他地区。供试小麦品种的抗性淀粉含量为1.47%～2.28%,平均为1.88%,以赛德麦1号、百农4199、许科316等品种的抗性淀粉含量较高。经相关分析,抗性淀粉含量与直链淀粉含量、峰值粘度、低谷粘度、最后粘度呈显著正相关。以上结果说明,提高小麦抗性淀粉含量,一定程度上可以改善淀粉的糊化特性。     

The impact of Septoria tritici blotch in bread making quality among argentinean wheat cultivars

Septoria tritici blotch (STB), caused by Zymoseptoria tritici is a relevant foliar wheat disease worldwide. Several reports show the importance of STB on grain yield, their components and grain protein while little is known about its effect on the rheological properties of the wheat flour. The scarce literature found, only mentions the effect of the complex of foliar diseases on wheat quality, without individualizing the effect of the different pathogens separately. This study analyze the influence of increasing doses of inoculum of Zymoseptoria tritici, on the bread making quality of ten Argentinean wheat cultivars and its possible variation according to their quality group. The increase of inoculum concentration augmented the area under disease progress curve, decreased green flag leaf area duration and green leaf area duration. Cultivars K. Flecha and B.75 Aniversario had the lowest green flag leaf area duration causing higher reduction in grain filling period and higher reductions in P, indicating a lower gliadin/glutenin ratio. STB decreased P/L and E while L, W, D, SV and bread volume increased. Cultivars differed in rheological parameters according to their quality group. Gluten/protein relationship was significant in quality group 1 and non-significant in cultivars belonging to quality group 2 and 3.     

新疆冬小麦品种品质性状与面包、馒头、面条加工品质的关系

为了给优质面包、馒头、面条专用品种选育提供品质辅助选择指标,以30份新疆冬小麦品种(包括自育品种和引进品种)为材料,分析了新疆冬小麦品种品质性状与面包、馒头、面条加工品质的关系.回归分析结果表明,小麦籽粒性状、面粉品质、面团特性、淀粉糊化特性以及面粉色泽对新疆冬小麦品种面包、馒头和面条加工品质均有显著影响;面粉灰分、湿面筋、稀懈值、亮度和红度是影响面包、馒头和面条加工品质的共同品质性状;形成时间、稳定时间、延展度是影响面包和面条加工品质的共同品质性状;而籽粒性状仅对新疆冬小麦品种馒头加工品质有显著影响.相关分析结果表明,千粒重、形成时间、稳定时间、拉伸面积、最大拉伸阻力、红度和黄度与面包总分呈显著相关关系,相关系数分别为0.460、0.516、0.537、0.719、0.707、0.534和-0.403;籽粒蛋白质含量、面粉蛋白含量和湿面筋含量与馒头总分呈显著相关关系,相关系数分别为-0.397、-0.458和-0.552,面团延展度、稀懈值与面条总分呈显著相关关系,相关系数分别为0.438和0.432.从以上结果可以看出,面包与面团流变学特性,馒头与蛋白质和面筋数量,面条与面团流变学特性及淀粉糊化特性的关系更为密切,这些品质性状可以作为新疆冬小麦品种面包、馒头、面条加工品质改良时的辅助选择指标.     

The impact of emmer genetic diversity on grain protein content and test weight of hexaploid wheat under high temperature stress

High temperature has a negative impact on wheat grain quality and reduces market value. Emmer wheat (Triticum dicoccon Schrank), one of the earliest domesticated wheat species, is a source of genetic diversity for the improvement of heat and drought tolerance in modern wheat. However, the potential of emmer wheat for the improvement of grain physical quality under high temperature stress is little studied. A diverse set of 184 emmer-based hexaploid lines was developed by crossing emmer wheat with hexaploid wheat and backcrossing once to hexaploid wheat. These materials, seven hexaploid recurrent parents and seven commercial cultivars, were evaluated at two times of sowing (E1 and E2) in the field, in 2015–2016. The materials were genotyped using a 90 K SNP platform and these data were used to estimate the contribution of emmer wheat to the progeny. Significant phenotypic and genetic variation for grain physical quality traits including protein content and test weight was observed. High temperature significantly increased protein content and decreased test weight. Large scale field phenotyping identified emmer progenies with improved grain characteristic compared to their respective parents and commercial cultivars in both environments. A few families consistently produced higher trait means across environments compared to their recurrent parents. The emmer wheat parent contributed between 1 and 37% of the genome in emmer-based genotypes. Selected emmer derived lines with superior protein content and test weight, tended to have a greater genetic contribution from the emmer parent, ranging from 12 to 37% and 7–37% in E1 and E2, respectively. It was concluded that new genetic variation for seed traits, such as protein content and test weight, can be introduced to hexaploid wheat from emmer wheat. The newly developed emmer derivatives identified with enhanced grain quality under high temperature stress can potentially be used to improve grain quality through breeding.     

Agronomic characteristics, grain quality and flour rheology of 372 bread wheats in a worldwide core collection

A core collection of 372 accessions representative of worldwide hexaploid bread wheat diversity [Balfourier, F., Roussel, V., Strelchenko, P., Exbrayat-Vinson, F., Sourdille, P., Boutet, G., Koenig, J., Ravel, C., Mitrofanova, O., Beckert, M., Charmet, G., 2007. A worldwide bread wheat core collection arrayed in a 384-well plate. Theoretical and Applied Genetics 114, 1265–1275] was used to evaluate the available genetic diversity of agronomic and quality characteristics. The traits assessed during the vegetative period were date of ear-emergence, date of flowering, lodging, disease susceptibility and pre-harvest sprouting. Thousand kernel weight, test weight, grain hardness, grain protein content, pentosan viscosity and grain colour were also measured. The rheological properties of the derived white flours were estimated using mixograph and alveograph tests. For most of the traits, a wide phenotypic variation was observed across all the accessions. Several parameters (mixograph width parameters before and after peak time, alveograph dough tenacity and extensibility, near infrared measurements, like those for protein content, and absorbance measurements of palmitic acid and linoleic acid content) made it easier to discriminate between the cultivars. The largest ranges of variation were found in landraces and old cultivars rather than in more recent varieties. This is evidence that there is sufficient variability available for rare alleles, which have been eliminated in breeding modern varieties to be detected. Such a core collection will therefore be a useful resource for future genetic studies on wheat quality.     

Agronomic characteristics,grain quality and flour rheology of 372 bread wheats in a worldwide core collection

A core collection of 372 accessions representative of worldwide hexaploid bread wheat diversity [Balfourier, F., Roussel, V., Strelchenko, P., Exbrayat-Vinson, F., Sourdille, P., Boutet, G., Koenig, J., Ravel, C., Mitrofanova, O., Beckert, M., Charmet, G., 2007. A worldwide bread wheat core collection arrayed in a 384-well plate. Theoretical and Applied Genetics 114, 1265–1275] was used to evaluate the available genetic diversity of agronomic and quality characteristics. The traits assessed during the vegetative period were date of ear-emergence, date of flowering, lodging, disease susceptibility and pre-harvest sprouting. Thousand kernel weight, test weight, grain hardness, grain protein content, pentosan viscosity and grain colour were also measured. The rheological properties of the derived white flours were estimated using mixograph and alveograph tests. For most of the traits, a wide phenotypic variation was observed across all the accessions. Several parameters (mixograph width parameters before and after peak time, alveograph dough tenacity and extensibility, near infrared measurements, like those for protein content, and absorbance measurements of palmitic acid and linoleic acid content) made it easier to discriminate between the cultivars. The largest ranges of variation were found in landraces and old cultivars rather than in more recent varieties. This is evidence that there is sufficient variability available for rare alleles, which have been eliminated in breeding modern varieties to be detected. Such a core collection will therefore be a useful resource for future genetic studies on wheat quality.