Resistance to Karnal Bunt (Tilletia indica Mitra) in Synthetic Hexaploid Wheats Derived from Triticum turgidum×T. tauschii

Synthetic hexaploids (SH) developed at the International Maize and Wheat Improvement Center (CIMMYT), involving four Triticum turgidum and nine T. tauschii parents, were evaluated for resistance to Karnal bunt (KB) (Tilletia indica Mitra) during three crop seasons over three years at Ciudad Obregon, Sonora, Mexico. Ten tillers of each SH at boot stage, taken at random, were injected with a suspension of sporidia in water (10,000 spores/ml of water). At maturity the inoculated spikes were threshed individually and evaluated for the percentage KB-infected grains. Based on the mean KB score of each entry for three seasons, 49 % of the SH were immune (0 % infection) to KB. Highly resistant expressions characterized the SH which appeared to be influenced by the resistance of their T. turgidum and/or T. tauschii parents. The overall mean infection of the SH wheats was 0.24 % compared to 56.14 % in the susceptible bread wheat check cultivat ‘WL711’. Transfer of KB resistance genes from SH wheats into bread wheat is currently underway at CIMMYT.     

Inheritance of threshability in synthetic hexaploid (Triticum turgidum×T. tauschii) by T. aestivum crosses

Triticum tauschii provides breeders with a valuable source of resistance and tolerance genes. Elucidation of the inheritance of traits in this species that hinder its use in breeding programmes is therefore of interest to wheat breeders. Inheritance of threshability was investigated in the crosses of four non-free-threshing (NFT) synthetic hexaploids (Triticum turgidum×T. tauschii) and two free-threshing (FT) T. aestivum cultivars during four crop seasons over 3 years at E1 Batan and Ciudad Obregon, Mexico. The parents, their F₁ Hybrids and individual F₂ plant-derived F₃ progenies of the crosses revealed that ‘Altar 84’/T. tauschii (219), ‘Chen’/T. tauschii (205), ‘Chen’/T. tauschii (224), and ‘Duergand’/T. tauschii (214) have independently segregating loci with two dominant alleles controlling threshability. Intercrosses among the synthetics, except ‘Altar 84’/T. tauschii (219), showed the genes to be allelic to each other. The cross between the FT cultivars showed no segregation in the F₃ generation, indicating common recessive genes. Based on these findings, population sizes of the synthetic-derived breeding materials should be increased to improve the chances of selecting FT desirable plants in the programme.     

CIMMYT新型人工合成小麦Pina和Pinb基因等位变异

六倍体人工合成小麦由硬粒小麦（Triticum turgidum subsp. durum）与粗山羊草（Aegilops tauschii Coss.）杂交产生,是研究小麦进化过程中基因变异的重要材料。以国际玉米小麦改良中心（CIMMYT）提供的57份由野生二粒小麦（T. turgidum subsp. dicoccoides）与粗山羊草杂交产生的新型人工合成六倍体小麦为材料，用单籽粒特性测定仪和Pina、Pinb特异性PCR引物对其籽粒硬度变异以及控制籽粒硬度的主效基因Pina和Pinb的分布情况进行了研究。结果表明，这些材料的SKCS硬度值变异较大，从10.5到42.6，其中15~30的占78%。共有Pina-D1a、Pina-D1c、Pinb-D1h和Pinb-D1j 4种等位变异型，基因型为Pina-D1a/Pinb-D1j的8个，占14%；基因型为Pina-D1c/Pinb-D1h的49个，占86%。方差分析表明，基因型Pina-D1a/Pinb-D1j与Pina-D1c/Pinb-D1h对籽粒硬度的影响差异不显著，但父本粗山羊草和母本野生二粒小麦以及二者间的互作对籽粒硬度有显著影响，说明除Pina和Pinb外，还有其他微效基因影响籽粒硬度的形成。     

Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

麦田杂草节节麦染色体核型分析研究

对麦田杂草二倍体节节麦（Triticum tauschii L．）的植株形态特征及根尖细胞染色体核型作了分析，并和六倍体普通小麦（Triticum aestivum L．）中国春核型作比较。结果表明，节节麦的核型公式为2n=2X=14=10M＋4SM（2SAT），在第4号染色体上有一对随体。节节麦染色体组和普通小麦中国春D组全套染色体类型相同，表明这两组染色体具有较强的同源性；但二者在染色体相对长度和臂比值上表现出一定的差异，表明该地区节节麦未参与普通小麦的起源。     

Expression of a Triticum turgidum var. dicoccoides source of Fusarium head blight resistance transferred to synthetic hexaploid wheat

Yield and quality reductions caused by Fusarium head blight (FHB) have spurred spring wheat (Triticum aestivum L.) breeders to identify and develop new sources of host plant resistance. Four wheat synthetic hexaploids (×Aegilotriticum sp.) were developed, each having a quantitative trait locus (QTL), Qfhs.ndsu‐3AS, providing FHB resistance from Triticum turgidum L. var. dicoccoides chromosome 3A. Synthetics were produced by hybridizing a ‘Langdon’‐T. dicoccoides‐ recombinant chromosome 3A substitution line (2n = 4x = 28, AABB with two accessions of T. tauschii (2n= 2x = 14, DD). Synthetics were inoculated and evaluated for FHB resistance in two separate greenhouse seasons. One synthetic, 01NDSWG‐5, exhibited FHB severity ratings of 36% and 32% in the separate seasons, compared with ratings of 9% and 30% for ‘Alsen’, a FHB‐resistant spring cultivar, and ratings of 70% and 96% for ‘McNeal’, a susceptible spring cultivar, respectively. Synthetic × Alsen backcross‐derived lines were produced to initiate combining different sources of FHB resistance.     

Suppression/expression of resistance to stripe rust in synthetic hexaploid wheat (Triticum turgidum×T. tauschii)

Seventy-four hexaploid wheats, synthesized by either crossing resistantTriticum turgidum L. var.durum with susceptible/intermediateT. tauschii or susceptible/intermediateT. turgidum with resistantT. tauschii, and their parents were evaluated as seedlings in the greenhouse and as adult-plants at two field locations in Mexico for resistance to pathotype 14E14 of stripe (or yellow) rust (caused byPuccinia striiformis Westend). The seedlings of different synthetic hexaploids showed high phenotypic diversity for resistance. However, the resistance level of only 15 of the 74 synthetic hexaploid wheats were similar to the low infection types of the respective donor parents. The remaining synthetic wheats displayed either intermediate or high infection types. A similar result was also obtained in field tests, where only 18 synthetic hexaploids were resistant as adult-plants. In general, genotypes with seedling resistance were also resistant as adult-plants. A few synthetic hexaploids, which displayed intermediate or susceptible infection types as seedlings were resistant as adult-plants, indicating that additional genes for adult-plant resistance were also present. The fact that resistance of some donor parents was not expressed, or only partially expressed, in a synthetic hexaploid background suggests the presence of suppressor genes in the both the A or B, and D genomes ofT. turgidum andT. tauschii, respectively. The resistance of a donor parent was expressed in a synthetic hexaploid only if the corresponding suppressor was absent in the second parent. Moreover, the suppressors appeared to be resistance gene specific.     

Genetic Variability for Haploid Production in Crosses Between Tetraploid and Hexaploid Wheats with Maize

Crosses were made between 14 wheat genotypes (11 tetraploid, 3 hexaploid) and a single Fl hybrid of maize that was used as the male parent. The experimental design consisted of randomized blocks with three replications. Plants were grown under controlled greenhouse conditions (day length 16 h and temperature 25 °C/15 °C, day/night). To enhance embryo survival, 2, 4-D treatment (10 mg/1) was applied to spikes 24 h after pollination with maize. Embryos were recovered from all tetraploid and hexaploid wheats at a rate of 2.09 to 26.76 per 100 pollinated florets. Haploid and doubled haploid plants were obtained from all hexaploid genotypes (T. aestivum) and from 5 of 11 tetraploid genotypes (T. turgidum var.). The most important point of these experiments was the ability to produce haploid plants from tetraploid wheat for two reasons: firstly, anther culture cannot be applied in tetraploid wheat (T. turgidum var.) due to the inefficiency of embryo formation and the high proportion of albino plants. Secondly, to date, crosses between tetraploid wheat and maize have resulted in embryo formation, but not in haploid plants.     

Genetic distance between Triticum timopheevi Zhuk., T. aramticum Jakubz. and T. aestivum L.

The genetic distances between two cultivated wheat (Triticum aestivum L.) varieties (Martonvásári 9, Martonvásári 15), a Martonvásári 9 line possessing the crossability gene krl, and 21 accessions of T. timopheevi Zhuk. and T. araraticum Jakubz. were estimated, based on agro-morphological, physiological and biochemical analysis of data. Cluster analysis based on Mahalanobis' D² values was applied. All 21 accessions of T. timopheevi and T. araraticum could be classified into eight clusters. Clusters I and II consisted of all the T. timopheevi, while T. araraticum was located in six clusters. Discriminant analysis was applied to test significant differences between cluster pairs. The genetic distance (GD) based on the electrophoretic data of gliadins indicated two types of electrophoregrams in T. timopheevvi, distinguished as groups A and B. T. araraticum accessions were variable as regards the spectra. Mean, minimum and maximum GD were estimated within and between different wheat groups based on acid polyacrylamide gel electrophoresis.     

Use of Triticum tauschii to improve yield of wheat in low-yielding environments

Triticum tauschii (Coss.) Schmal. is an ancestor of bread wheat (T. aestivum). This species has been widely used as a source ofsimply-inherited traits, but there are few reports of yield increases due tointrogression of genes from this species. Selections from F₂-derivedlines of backcross derivatives of synthetic hexaploid wheats (T.turgidum / T. tauschii) were evaluated for grain yield in diverseenvironments in southern Australia. Re-selections were made in theF₆ generation and evaluated for grain yield, yield componentsincluding grain weight, and grain growth characters in diverse environmentsin southern Australia and north-western Mexico. Re-selection was effectivein identifying lines which were higher yielding than the recurrent parent,except in full-irrigation environments. Grain yields of the selectedderivatives were highest relative to the recurrent parent in thelowest-yielding environments, which experienced terminal moisture deficitand heat stress during grain filling. The yield advantage of the derivativesin these environments was not due to a change in anthesis date orgrain-filling duration, but was manifest as increased rates of grain-filling andlarger grains, indicating that T. tauschii has outstanding potential forimproving wheat for low-yielding, drought-stressed environments.     

Assessment of genetic diversity in synthetic hexaploid wheats and their Triticum dicoccum and Aegilops tauschii parents using AFLPs and agronomic traits

Synthetic hexaploid wheats are of interest to wheat breeding programs, especially for introducing new genes that confer resistance to biotic and abiotic stresses. A group of 54 synthetic hexaploid wheats derived from crosses between emmer wheat(Triticum dicoccum, source of the A and B genomes) and goat grass (Aegilops tauschii, D genome donor) were investigated for genetic diversity. Using the AFLP technique, dendrograms revealed clear grouping according to geographical origin for the T. dicoccum parents but no clear groups for the Ae. tauschii parents. The geographical clustering of the T. dicoccum parents was also reflected in the dendrogram of their derived synthetic hexaploids. Diversity of the T. dicoccum parents and their derived synthetic hexaploids was further evaluated by measuring 18morphological and agronomic traits on the plants. Clustering based on morphological and agronomic data also reflected geographical origin. However, comparison of genetic distances obtained from AFLP and agronomic data showed no correlation between the two diversity measurements. Nevertheless, similarities among major clusters with the two systems could be identified. Based on percentage of polymorphic markers, the synthetic hexaploids had a considerably higher level of AFLP diversity (39%) than normally observed in cultivated hexaploid wheat (12–21%). This suggests that synthetic hexaploid wheats can be used to introduce new genetic diversity into the bread wheat gene pool. This revised version was published online in July 2006 with corrections to the Cover Date.     

人工合成六倍体小麦后代衍生群体Waxy蛋白亚基的分子标记

四倍体小麦与节节麦杂交培育的人工合成六倍体小麦已广泛应用于国内外小麦品种改良。以引自CIMMYT的Syn768、Syn769和Syn780人工合成六倍体小麦分别与中国四川成都平原主栽普通小麦品种杂交、回交的BC2F2:6后代群体中选育的113份优良高代系为材料,采用SSR特异引物的PAGE凝胶电泳对其Waxy蛋白亚基缺失类型进行了研究。结果表明,在所检测的121份材料中,8份材料缺失Waxy-B1型蛋白亚基,占全部材料的6.6%;没有检测到其他类型的缺失体。从每个人工合成六倍体小麦亲本材料所形成的后代衍生群体来看, Waxy-B1缺失体频率各不相同,说明Waxy蛋白亚基缺失类型在人工合成六倍体小麦后代衍生群体材料中的表现存在着随机性,与亲本的基因型状况关系极大。通过研究人工合成六倍体小麦与普通小麦杂交后代的Waxy蛋白亚基缺失类型,有助于提高分子标记育种效率。     

Agronomic and qualitative traits of T. turgidum ssp. dicoccum genotypes cultivated in Italy

Summary The increasing popularity of organic agriculture and health food products has led to a renewed interest in hulled wheat species such as emmer (Triticum turgidum ssp. dicoccumSchubler). Knowledge on agronomic and quality traits is required for effective and efficient use of germplasm collections in breeding programs. The objective of this study was to estimate agronomic and grain quality traits of emmer wheat cultivated in Italy. A total of 20 emmer accessions consisting of landraces, breeding lines or cultivars selected from landraces and modern cultivars were examined under low input conditions. The study was conducted for three successive years (2002–2004) at one location of Southern Italy (Foggia). The entries were characterized for agronomic and qualitative traits [grain yield (t ha⁻¹), thousand grain weight (g), test weight (kg hl⁻¹), grain protein content (%), HMWG composition, dry gluten content (%), gluten index and yellow index, alveograph indices and Total Organic Matter (TOM) on cooked pasta]. The results showed a large genetic variability for most of the traits measured and, even if most of the accessions showed inferior bread- and pasta-making performance, modern cultivars exhibited improved quality traits with some potential to perform healthy and tasty food.     

Agronomic performance of chromosomes 1B and T1BL.1RS near-isolines in the spring bread wheat Seri M82

The T1BL.1RS wheat (Triticum aestivum L.) - rye (Secale cereale L.) translocations have been of particular interest and are widely used in bread wheat breeding programs. The objective of this study was to determine the effect of the T1BL.1RS chromosome on grain yield and its components using 20 near-isolines of spring bread wheat cultivar ‘Seri M82’ (10 homozygous for chromosome 1B substitution and 10 homozygous for T1BL.1RS). The test lines have been produced by substituting the 1B chromosome in Seri M82 (T1BL.1RS, T1BL.1RS) through backrossing. Two field experiments were evaluated under optimum (five irrigations) and reduced (one irrigation) moisture conditions for two consecutive production cycles at the Mexican National Agricultural Research Institute, Ciudad Obregon, Sonora, Mexico. The presence of T1BL.1RS had a significant effect on grain yield, harvest index, grains/m², grains/spike, 1000-grain weight, test weight, flowering date and physiological maturity in both moisture conditions. The agronomic advantage of the 1B substitution lines on above-ground biomass yield at maturity, spikes/m²and grain-filling duration was expressed only under the optimum moisture condition. The presence of T1BL.1RS increased grain yield 1.6% and 11.3% for optimum and reduced moisture conditions, respectively. These results encourage further use of T1BL.1RS wheats in improving agronomic traits, especially for reduced irrigation or rainfed environments. This revised version was published online in August 2006 with corrections to the Cover Date.     

四倍体小麦地方品种矮蓝麦矮秆性状的遗传分析

四倍体圆锥小麦(Triticum turgidum L.ssp.turgidum)地方品种矮蓝麦是我国重要的小麦矮秆基因资源,经鉴定其矮秆特性对外源赤霉酸敏感。2012年配制矮蓝麦与2个高秆圆锥小麦的正反交组合,2012—2013年在四川绵阳分别种植F1、F2代和F2:3家系,对株高的遗传分析表明,矮蓝麦的矮秆性状受1对隐性基因控制。利用BSA法构建高秆和矮秆池筛选多态性SSR标记,并对矮蓝麦/青稞麦F2分离群体进行连锁分析,将目标基因定位于7AS染色体上,与标记GWM471的遗传距离为2.5 c M。矮蓝麦与矮秆番麦正反交的F1和F2群体表现非常相似的株高变异特征,初步推测矮蓝麦的矮秆基因是Rht22;进一步用高通量SNP和DAr T标记对两品种进行全基因组扫描,发现二者的遗传相似性高达98.7%~99.3%。因此认为,历史上矮蓝麦和矮秆番麦可能是同一品种,是通过人为交流而传播到不同地方。矮蓝麦携带的矮秆基因在人工合成六倍体小麦遗传背景中降低株高能力中等或较弱,在育种中需要聚合其他矮秆基因而被利用。     

Use of the gene pools of Triticum turgidum ssp. dicoccoides and Aegilops squarrosa for the breeding of common wheat (T. aestivum), through chromosome-doubled hybrids

Summary Triticum turgidum ssp. dicoccoides (wild emmer wheat, AABB, 2n=28) and Aegilops squarrosa (goat grass, DD, 2n=14) comprise a rich reservoir of valuable genetic material, which could be useful for the breeding of common wheat (T. aestivum, AABBDD, 2n=42). Many accessions of both wild species, most of them selected for resistance to stripe rust, were used to make amphiploids. Two strategies were applied: (1) the production of autopolyploid cytotypes of the wild species, followed by hybridisation, and (2) the production of allotriploid interspecific hybrids, followed by doubling of the number of chromosomes. The first route was unsuccessful because of failure of the crosses between the autopolyploid cytotypes, possibly due to incongruity between the two species and to reduced fertility in the autopolyploid cytotypes. The second route yielded the desired synthetic hexaploids. However, the rate of success of the crosses was low and there were great differences between years, and within years between crosses. Embryo rescue was applied to obtain the primary hybrids (2n=21), which were highly sterile and had on average 0.3 bivalents and 20.4 univalents per pollen mother cell. Various abnormalities were recorded. Doubling of the number of chromosomes sometimes occurred spontaneously or was brought about by colchicine treatment. The large scale of the interspecific hybridisation programme ensured that one-third of the female and one-sixth of the male accessions were represented in the synthetic hexaploids.     

Further Studies on the Restoration of Fertility in Triticum durum×T. timopheevi Hybrids

Twenty one F₁ hybrids involving three Triticum durum cultivars and eleven forms of T. timopheevi were backcrossed to their respective durum parents. Backcross fertility (BC₁ grain set) of these sterile hybrids improved with the rise in temperature at the time of pollination. A mean temperature of less than 20 °C results in poor seed set while higher mean temperatures around 24 °C result in increased seed set. It is suggested that a large number of backcross seeds can be produced by pollinating these hybrids late in the season, i.e., after 20th of March in northern India.     

Intra- and inter-population variations in grain protein percentage in wild tetraploid wheat,Triticum turgidum var. dicoccoides

Summary Grain protein percentage (GPP) was studied in 910 accessions of the wild tetraploid wheat, Triticum turgidum var. dicoccoides, collected from 22 populations representing different ecogeographical conditions in Israel. High values of GPP were found, ranging from 19.7% to 28.0% for population means, and from 14.1% to 35.1% for single accessions. Marginal populations had usually lower GPP and smaller intra-population variation than central ones. Repeated sampling of some central populations for four consecutive years revealed relatively large intra-population fluctuations. A high and significant genetic component of variation was found within and between populations by a nested analysis of variance in two nurseries. However, the regression coefficients of parents vs. offsprings were relatively low, indicating a smaller genetic component of variation which may be accounted for by a significant genotype × environment interaction. No correlation was found between GPP and ecological factors, except for soil type: accessions growing on terra-rossa had higher GPP than those growing on basaltic soil. Accessions with black glumes, or with glabrous auricles, or with large grains exhibited high GPP values. A strategy for collecting accessions with high GPP is presented, and the potential use of high GPP genotypes in breeding programs is discussed.     

Polymorphism of two waxy proteins in the emmer group of tetraploid wheat, Triticum dicoccoides, T. dicoccum, and T. durum

Gel-electrophoretic analyses detected polymorphism of two waxy (Wx) proteins, Wx-Al and Wx-Bl, in 334 accessions of the emmer group of tetraploid wheat, Triticum dicoccoides, T. dicoccum and T. durum. The null allele for the Wx-Al protein (Wx-Alb) was found in one accession of T. dicoccoides and seven of T. dicoccum, but it was not present in those T. durum accessions analysed. The null allele for the Wx-Bl protein (Wx-Blb) did not occur in three emmer-wheat species. Wx-Al and Wx-Bl proteins showing alteration in mobility in SDS-PAGE gel or in isoelectric points were found in five accessions and considered to be the products of new alleles, Wx-Ald, Wx-Ale and/or Wx-Bld. Densitometric analysis of Wx protein bands revealed that Wx-Al was present in smaller amounts than Wx-B1 in almost all accessions.     

New durum wheat with Hessian fly resistance from Triticum araraticum and T. carthlicum in Morocco

Hessian fly is an important pest of wheat on the North American continent and the temperate Mediterranean drylands. Yield losses caused by this insect in Morocco are the heaviest in the Mediterranean region and are estimated to be 36% on average. Genetic resistance to Mediterranean Hessian fly biotypes has not been found in durum wheat, although large numbers of durum accessions were screened. Genes for resistance were found in common wheat; some of which are transferable to durum. However, there is a need to broaden the genetic base for resistance in durum wheat. The objective of this work was to introgress resistance from selected Triticum araraticum and T. carthlicum accessions using multiple backcross methodology. The experimental recipient durum wheat included numerous adapted and high‐yielding lines. Testing for Hessian fly resistance under controlled conditions and field yield data showed that this programme yields Hessian fly‐resistant durum lines with good yields and adaptation.