Inheritance of Isoproturon Tolerance in Durum Wheat Transferred from Triticum monococcum

The inheritance of isoproturon tolerance exhibited by the selected progenies of durum wheat was investigated. This tolerance to the herbicide isoproturon had been transferred from Triticum monococcum to durum wheat cultivars. It was shown to be conditioned by a single dominant gene     

Prehaustorial resistance to the wheat leaf rust fungus, Puccinia triticina, in Triticum monococcum (s.s.)

Diploid wheat, Triticum monococcum s.l., is a host for the wheat leaf rust fungus, Puccinia triticina. Some accessions have been reported to show a high degree of prehaustorial resistance. This is non-hypersensitivity resistance, which acts before the formation of haustoria by the pathogen. To assess the frequency of prehaustorial resistance 598 accessions of diploid wheat were inoculated with the wheat leaf rust isolate Felix. Most T. monococcum s.s. accessions (84%) were resistant whereas all T. urartu and all but three T. boeoticumaccessions were susceptible. Histological components analysis revealed that a high percentage of prehaustorial resistance to P. triticina was found in only three T. monococcum accessions. No haustoria were observed in such infection units confirming the prehaustorial nature of the resistance. Prehaustorial abortion of certain infection units in an accession always coincided with posthaustorial abortion of the other infection units. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of Cold Hardiness in Triticum aestivum × Synthetic Hexaploid Wheat Crosses

Synthetic hexaploid wheat, produced by combining tetraploid wheat (AB genome) with Triticum tauschii (D genome), was crossed to modern hexaploid wheat (Triticum aestivum ABD genome) in an attempt to introduce new cold hardiness genes into the common hexaploid wheat gene pool. The cold hardiness levels of F) hybrids ranged from similar to parental means to equal to the hardy parent, indicating that cold hardiness was controlled by both additive and dominant genes. As expected when dominant gene action is involved, differences between F₂ and parental means were smaller than comparable differences in the F., Frequency distributions of F₂—derived F₃ lines also suggested that dominant genes were involved in the control of cold hardiness in some crosses. Heritability estimates for cold hardiness ranged from 63 to 70 % indicating that selection for cold hardiness should be effective in populations arising from crosses between common and synthetic hexaploid wheat. However, high selection pressure on the progeny of crosses that included the most hardy T. aestivum, T. durum, and T. tauschii accessions as parents did not identify transgressive segregates for improved cold hardiness. These observations indicate that the close wheat relatives, sharing common genomes with T. aestivum, are not promising sources of new genes to increase the maximum cold hardiness potential of common hexaploid wheat.     

Variation in resistance to the grain aphid, Sitobion avenae (Sternorhynca: Aphididae), among diploid wheat genotypes: Multivariate analysis of agronomic data

A collection of 87 ancient wheat genotypes, 67 Triticum monococcum, 13 Triticum boeoticum, seven Triticum urartu and one cultivar of the modern wheat Triticum aestivum (variety ‘Arminda’) were evaluated for resistance to the cereal aphid Sitobion avenae, the main damaging aphid pest on winter wheat in Europe. The intrinsic rate of natural increase (r_m), which is regarded as a good estimate of the fitness of an aphid population, was used as an indicator for the level of plant resistance. Differentiation of the 88 plant genotypes into four distinct groups was achieved with a cluster analysis of the r_m values. The modern wheat ‘Arminda’ was more susceptible than any of the ancient wheat genotypes tested (r_m= 0.24, i.e. the aphid population doubled every 2.6 days). A second group of 19 plants ranged from relatively susceptible to moderately resistant (0.17 < r_m < 0.21). Fifty–one plants were allocated to a third group and classified as resistant (0.09 < r_m < 0.16). The last group contained 17 genotypes with a high level of resistance where aphid fitness was greatly reduced (0.02 < r_m < 0.09, i.e. the aphid population doubled every 11.4 days or 7.7 days, respectively). Clustering of the accessions into the different phenetic groups did not follow the geographical origin of the wheat genotypes or the species to which they belong. These results show that ancient diploid wheats, all characterized by the genome A, present considerable interest for plant breeding for resistance to S. avenae in modern wheat. The potential use of these strong and partial sources of resistance for introduction of a stable and durable form of resistance to S. avenae in wheat is discussed.     

Host plant resistance in some wild wheats to the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae)

A total of 259 accessions of wild Triticum species originating from different countries, along with 91 triticale (6×)× bread wheat true-breeding derivatives, two bread wheat, and three triticale cultivars were screened for resistance to the Russian wheat aphid, a serious insect pest of the wheat crop. Twenty-four entries with low damage ratings on the basis of amount of leaf rolling and leaf chlorosis were retested along with resistant and susceptible controls. On the basis of leaf roll damage ratings, eight entries including four Triticum monococcum var. boeoticum (T. boeoticum), one T. monococcum var. monococcum (T. monococcum), two T. timopheevii var. araraticum (T. araraticum), and one triticale cultivar were significantly superior to ‘Karl’ (susceptible control) wheat. Among these, four accessions — three T. boeoticum and one T. araraticum— were significantly superior to all other entries and were equal to the resistant control (PI 372129) in resistance rating based on leaf rolling and leaf chlorosis (except T. boeoticum TA 202). The leaf chlorosis damage rating of all accessions were significantly lower than that of the susceptible check.     

Intergeneric Transfer (Rye to Wheat) of a Gene(s) for Russian Wheat Aphid Resistance

An octoploid triticale was derived from the F, of a Russian wheat aphid-resistant rye, ‘Turkey 77’, and ‘Chinese Spring’ wheat. The alloploid was crossed to common wheat, and to ‘Imperial’ rye/‘Chinese Spring’ disomic addition lines. F₂, progeny from these crosses were tested for Russian wheat aphid resistance and C-banded. A resistance gene(s) was found to be associated with chromosome arm IRS of the ‘Turkey 77’ rye genome. A monotelosomic IRS (‘Turkey 77’) addition plant was then crossed with the wheat cultivar ‘Gamtoos’, which has the 1BL.1RS ‘Veery’ translocation. Unlike the IRS segment in ‘Gamtoos’, the ‘Turkey 77’-derived 1 RS telosome did not express the rust resistance genes Sr31 and Ar26, which could then be used as markers. From the F, a monotelosomic 1 RS addition plant that was also heterozygous for the 1BL. 1 RS translocation was selected and testerossed with an aphid-susceptible common wheat, ‘Inia 66’ Meiotic pairing between the rye arms resulted in the recovery of five euploid Russian-wheat-aphid-resistant plants. One recombinant also retained Sr31 and Lr26 and was selfed to produce translocation homozygotes.     

Comparative genetics and disease resistance in wheat

The hexaploid wheat genome is too complex for direct map-basedcloning and model genomes have to be used to isolate genes from wheat.Comparative genomic analysis at the genetic map level has shown extensiveconservation of the gene order between the different grass genomes inmany chromosomal regions. However, little is known about the geneorganization in grass genomes at the microlevel. We have investigated themicrocollinearity at Lrk gene loci in the genomes of four grass species:wheat, barley, maize and rice. The Lrk genes, which encodereceptor-like kinases, were found to be consistently associated with anothertype of receptor-like kinase (Tak) on chromosome groups 1 and 3 inTriticeae and on chromosomes homoeologous to Triticeae group 3 in theother grass genomes. On Triticeae chromosome group 1, Tak and Lrk together with genes putatively encoding NBS/LRR proteins form acluster of genes. Comparison of the gene composition at orthologous Lrk loci in wheat, barley and rice revealed a maximal gene density of onegene per 5 kb. We conclude that small and large grass genomes containregions which are highly enriched in genes. Microrearrangements betweendifferent grass genomes have been found and therefore, the choice of agood model genome is critical. We have recently started to work on theT. monococcum model genome and confirmed its usefulness foranalysis of the Lr10 leaf rust disease resistance locus in wheat.     

An approach to DNA polymorphism screening in <Emphasis Type="Italic">SBEIIa</Emphasis> homeologous genes of polyploid wheat (<Emphasis Type="Italic">Triticum</Emphasis> L.)

The non-transgenic manipulation of starch properties in common wheat (Triticum aestivum L.) generally implies combining mutant alleles of the particular gene copies in all three subgenomes (A, B and D). The redundancy of the hexaploid wheat chromosome set substantially complicates the identification of recessive mutations and breeding. Nevertheless, naturally occurring or induced genetic polymorphism has already been successfully exploited for the production of waxy (GBSSI-deficient) and elevated amylose (SSIIa-deficient) wheats. However, in order to achieve the amylose content above 50% of wheat endosperm starch, it may be necessary to inactivate the starch branching enzyme (SBEIIa) isoforms, as the RNAi repression results and gene expression data strongly suggest. The identification of null SBEIIa alleles and their combination in a single genotype is therefore a promising approach to the production of non-transgenic high-amylose wheat; however, wheat SBEIIa polymorphism has not been characterized as of yet. In order to develop an approach to SBEIIa mutation screening, we sequenced the SBEIIa central region (exons 9–12) from the three subgenomes of common wheat cv. Chinese Spring and the A genome of diploid einkorn T. monococcum. The genome-specific primers were developed that amplify the exons downstream from intron 11 selectively from each homeologous gene. Using a single-stranded DNA conformation polymorphism (SSCP) approach, we screened 60 wheat cultivars, landraces, and rare species for naturally occurring SNPs in exons 12, 13 and 14 of the three SBEIIa homeologs. In total, 13 SNPs were discovered in the A and B wheat genomes. Two of these SNPs affect the amino acid sequences of SBEIIa isoforms and may change the enzyme functional properties. The presence of restriction site polymorphism at SNP positions enables their easy genotyping with CAPS assays. Our results indicate that the mining for naturally occurring sequence polymorphism in starch biosynthesis genes of wheat can be successfully performed at the DNA level, providing the starting point for a search for SBEIIa mutants at a larger scale.     

Mapping of genes involved in glutathione, carbohydrate and COR14b cold induced protein accumulation during cold hardening in wheat

Using some of the chromosome substitution lines developed from thecrosses of the donor Cheyenne to Chinese Spring we showed that theaccumulation of water soluble carbohydrates during different stages ofhardening was time dependent. Moreover there was a significantcorrelation between the rate of carbohydrate accumulation and the frosttolerance. The expression and regulation of a wheat gene homologous tothe barley cold regulated cor14b gene was compared in frost sensitiveand frost tolerant wheat genotypes at different temperatures. Studies madewith chromosome substitution lines showed that the threshold inductiontemperature polymorphism of the cor14b wheat homologous genewas controlled by loci located on chromosome 5A of wheat, while cor14b gene was mapped, in Triticum monococcum, onto the longarm of chromosome 2A^m. Our study on the effect of cold hardeningon glutathione (GSH) metabolism showed that chromosome 5A of wheathas an influence on the GSH accumulation and on the ratio of reduced andoxidised glutathione as part of a complex regulatory function during coldhardening. In addition, the level of increase in GSH content duringhardening may indicate the degree of the frost tolerance of wheat.     

Reproductive Behavior of Hexaploid/Diploid Wheat Hybrids 1

The bottleneck restricting introgression of useful genes directly from diploid into hexaploid wheats is the low number of BC₁F₁ seeds obtained. In crosses between hexaploid wheat (Triticum aestivum L.; AABBDD) and Aegilops squarrosa L. (DD) or T. urartu Thum. (AA), this bottleneck may be overcome simply by pollinating a sufficient number of F₁ spikes. However, hybrids between hexaploid wheat cultivars (T. aestivum) and T. monococcum L. (AA) generally are highly female-sterile, often having no pistils. One T. monococcum accession, PI 355520, when crossed with T. aestivum, produced hybrids with female fertility in the same range as that of T. aestivum/A. squarrosa or T. aestivum/T. urartu hybrids, ca. 0.5 to 1.0 backcross seed per spike. We found that female fertility was controlled by two duplicate genes in PI 355520, and that this accession can be used as a bridging parent to introgress genes from other T. monococcum accessions into hexaploid wheat. Pairing of homologous chromosomes was less frequent and weaker in such crosses than in T. aestivum/A. squarrosa crosses, but homoeologous bivalents occurred at a rate of almost 0.5 II per cell. Restitution division was detected in crosses involving all three diploid species and was confirmed cytologically in crosses with PI 355520. Chromosome numbers of BC₁F₁ plants ranged from 35 to 67; plants with 49 or more chromosomes occurred at frequencies of 0.09 to 0.21 among progeny of A. squarrosa and T. urartu and 0.29 in progeny of T. aestivum/T. monococcum crosses involving PI 355520. These results are consistent with those of previous studies, demonstrating the potential of direct Hexaploid/diploid crosses for rapidly introgressing useful genes into Hexaploid wheat with minimum disturbance of the background genotype.     

Application of Triticum monococcum for the improvement of triticale resistance to leaf rust (Puccinia triticina)

The aim of this work was to evaluate the leaf rust resistance introduced into introgressive triticale lines with Triticum monococcum genes, and to study the expression of these genes at the hexaploid level. The introgressive lines were developed by incorporating diploid wheat (T. monococcum s.s.) genes into hexaploid triticale LT 522/6 using the synthetic allotetraploid T. monococcum/Secale cereale (A^mA^mRR) as a bridging form. A group of 44 those lines, parental stocks and check cultivars were inoculated at the seedling stage (in a greenhouse) and at the adult‐plant stage (in the field) with four pathotypes of Puccinia triticina. At the seedling stage the assessment of infection type showed that four lines had resistance to all pathotypes as high as in the T. monococcum donor. Adult plant examinations showed some introgressive lines with complete resistance and also lines with partial resistance, expressed in area under the disease progress curve (AUDPC) calculations as slow rusting. Some lines comprise low AUDPC with complete resistance at seedling stage.     

Agronomical performance and breeding value of selected strains of diploid wheat,Triticum monococcum

Summary Twenty-one germplasm accessions and breeding lines of einkorn wheat (Triticum monococcum ssp. monococcum and ssp. sinskajae) were grown at two sites in Italy and evaluated for various field and seed characteristics. Grain yields of germplasm accessions were relatively high (317–3238 kg/ha), but distinctly lower than those of four modern cultivars of tetraploid (T. turgidum ssp. durum) and hexaploid wheat (T. aestivum ssp. Aestivum) included in the experiments as controls. As expected, all Einkorns-including some substantially higher yielding crossbred lines (3415–4362 kg/ha)-were defective for one or more agronomically relevant features. However, a few of the accessions examined were found to contain, as a group, practically all the genes needed to breed monococcums having the main field attributes of a modern wheat cultivar: high yielding capacity, good threshability, large kernel size, earliness, short stature and adequate lodging resistance. Still higher yielding diploid wheats, more responsive to improved growing conditions and of better seed quality, could probably be obtained from crosses with wild monococcums bearing mostly two-seeded florets and with accessions producing less slender-shaped kernels. Some of the Einkorns examined were found to carry minor genes for easy threshing which might enhance the efficacy of the major gene for soft glumes carried by T. monococcum ssp. sinskajae, a partially free-threshing diploid wheat taxon. Seed protein percentage of monococcums was markedly higher than that of durum and bread wheat cultivars even in those cases where their grain yields surpassed those of the polyploid checks. The possibilities offered by diploid wheat in the exploitation of novel endosperm mutants and F₁ hybrid vigour, as well as in the fields of celiac disease, crop diversification and resistance to agro-biological stresses are discussed. Breeding priorities and strategies are also proposed.     

野生二粒小麦导入普通小麦的抗白粉病基因MlWE29分子标记定位

小麦白粉病是严重影响小麦生产的重要病害之一,培育和应用抗病品种是有效控制和减少病害的最经济有效的方法。野生二粒小麦是硬粒小麦和普通小麦的四倍体野生祖先种,是小麦抗病性遗传改良的重要基因资源。本研究利用来自以色列的野生二粒小麦WE29与普通小麦杂交,再用普通小麦连续回交和自交,育成高抗白粉病(Blumeria graminis f. sp. tritici)小麦新品系3D258(系谱为燕大1817/WE29//5*87-1, BC₄F₆)。将3D258和高感小麦白粉病的普通小麦品种薛早配制杂交组合,对其F₁、F₂代分离群体和F₃代家系进行白粉病抗性鉴定和遗传分析。结果表明3D258携带抗白粉病显性单基因,暂命名为MlWE29。利用集群分离分析法(BSA)和分子标记分析,发现6个SSR标记(Xgwm335、Xgwm213、Xgwm639、Xwmc415、Xwmc289和Xwmc75)和5个EST-STS标记(BE494426、BE442763、CD452476、BE445282和BE407068)与抗白粉病基因MlWE29连锁。利用中国春缺体-四体系、双端体系和缺失系将抗白粉病基因MlWE29标记物理定位于5BL染色体的0.59–0.79区域。这一普通小麦抗白粉病种质资源的创制及其连锁分子标记的建立为小麦抗病基因分子标记辅助选择、基因积聚和分子育种提供了新的物质基础。     

A New Intergeneric Hybrid between Triticum aestivum L. and Agropyron fragile (Roth) Candargy: Variation in A. fragile for Suppression of the Wheat Ph-Locus Activity

New intergeneric hybrids were obtained between Triticum aestivum L. cv. Tukuho’ (2n = 6x = 42, AABBDD) and Agropyron fragile (Roth) Candargy PGR 8097 (2n = 4x = 28, PPPP) at a frequency of 1.06 %, through the use of direct embryo culture and in ovulo embryo culture. Such hybrids could be used to transfer barley yellow dwarf virus (BYDV) resistance and winterhardiness into bread wheat. The somatic chromosome number in all the hybrid plants was 2n = 5x = 35, as expected. Considerable variation in chromosome pairing was observed among the different hybrid plants. Average meiotic chromosome configuration at metaphase I was 17.29 Is + 6.57 rod Us + 1.97 ring Us + 0.18 III + 0.03 IV + 0.002 VI. The high level of chromosome pairing in some F₁ hybrids was attributed to Ph-suppressor gene(s) present in A. fragile. The hybrids could not be backcrossed to wheat, but amphiploid seeds have been obtained by colchicine treatment.     

Detection of Resistance to the Russian Wheat Aphid in Hexaploid Wheat

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), poses a serious threat to wheat (Triticum aestivum L.) production in many parts of the world. This research was initiated to evaluate wheat accessions for detection of resistance to the RWA. Over 12,000 wheat cultivars and plant introductions (PIs) from the USDA-ARS National Small Grains Collection were evaluated for reaction to RWA feeding damage. Twenty-nine PIs from Iran, Afghanistan, and the former Soviet Union, of various agronomic backgrounds were identified as having moderate to high levels of RWA resistance. This information is useful to wheat breeders searching for sources of resistance to the RWA to incorporate into their breeding programmes.     

Wheat streak mosaic virus resistance in eight wheat germplasm lines

Wheat streak mosaic virus (WSMV) is a destructive pathogen in wheat (Triticum aestivum L.). Host resistance is the most effective way to control this virus. To date, Wsm2 is the only wheat resistance gene that is genetically mapped. The objective of this study was to identify germplasm lines that might carry resistance genes different from Wsm2. Eight newly reported resistant germplasm lines were examined by allelic tests. To validate the allelic test results, five of them were further analysed for the inheritance of WSMV resistance. A Wsm2‐linked marker was also genotyped on populations developed for the inheritance study. Our results suggested that the WSMV resistance in lines CItr9358, PI225288, PI243652, PI245439, PI245526 and PI478095 was controlled by either Wsm2 or a gene very closely linked to Wsm2. The resistance in PI243753 and PI321730, however, is likely controlled by a gene different from, but linked to Wsm2. The resistance in PI321730 might also involve some minor genes. This study provided useful information for breeders to select appropriate resistant lines to improve WSMV resistance in wheat.     

Molecular mapping of cereal cyst nematode resistance in <Emphasis Type="Italic">Triticum monococcum</Emphasis> L. and its transfer to the genetic background of cultivated wheat

Triticum monococcum, the diploid A genome species, harbours enormous variability for resistance to biotic stresses. A spring type T. monococcum acc. 14087 was found to be resistant to Heterodera avenae (cereal cyst nematode, CCN). A recombinant inbred line population (RIL) developed by crossing this accession with a CCN susceptible T. boeoticum acc. 5088 was used for studying the inheritance and map location of the CCN resistance. Based on composite interval mapping two QTL, one each on chromosome 1AS and 2AS, were detected. The QTL on 1A, designated as Qcre.pau-1A, appeared to be a major gene with 26% contribution to the overall phenotypic variance whereas the QTL on 2A designated as Qcre.pau-2A contributed 13% to total phenotypic variation. Qcre.pau-1A is novel, being the only CCN resistance gene mapped in any ‘A’ genome species and none of the other known genes have been mapped on chromosome 1A. The QTL Qcre.pau-2A might be allelic to Cre5, a CCN resistance gene transferred from Ae. ventricosa and mapped on 2AS. The Qcre.pau-1A was transferred to cultivated wheat using T. durum cv. PBW114 as the bridging species. Selected CCN resistant F₈ lines showed introgression for the molecular markers identified to be linked with CCN resistance locus Qcre.pau-1A. Thus, this gene alone could impart complete resistance against CCN. These introgression lines can be used for marker-assisted transfer of Qcre.pau-1A to elite wheat cultivars.     

The influence of Elymus sibiricus L. genome on the diploidization system of wheat

Summary The meiotic behaviour of five tetraploid wheat strains x Elytricum fertile (2n=42 chromosomes, AABBD(SH) genomes) F₁ hybrids has been analysed. Multivalent associations were observed in the hybrids which could be attributed to Elymus sibiricus L. gene (s) somewhat suppressing the activity of the wheat homoelogous pairing control system. This interaction depends on the wheat genotype. The effect was particularly notable when Triticum turgidum var. salomonis was the wheat parent. The possibility of gene transfer from Elymus to wheat is discussed.     

野生二粒小麦导入普通小麦的抗白粉病基因MlWE29分子标记定位

小麦白粉病是严重影响小麦生产的重要病害之一,培育和应用抗病品种是有效控制和减少病害的最经济有效的方法。野生二粒小麦是硬粒小麦和普通小麦的四倍体野生祖先种,是小麦抗病性遗传改良的重要基因资源。本研究利用来自以色列的野生二粒小麦WE29与普通小麦杂交,再用普通小麦连续回交和自交,育成高抗白粉病(Blumeria graminis f. sp. tritici)小麦新品系3D258(系谱为燕大1817/WE29//5*87-1, BC₄F₆)。将3D258和高感小麦白粉病的普通小麦品种薛早配制杂交组合,对其F₁、F₂代分离群体和F₃代家系进行白粉病抗性鉴定和遗传分析。结果表明3D258携带抗白粉病显性单基因,暂命名为MlWE29。利用集群分离分析法(BSA)和分子标记分析,发现6个SSR标记(Xgwm335、Xgwm213、Xgwm639、Xwmc415、Xwmc289和Xwmc75)和5个EST-STS标记(BE494426、BE442763、CD452476、BE445282和BE407068)与抗白粉病基因MlWE29连锁。利用中国春缺体-四体系、双端体系和缺失系将抗白粉病基因MlWE29标记物理定位于5BL染色体的0.59–0.79区域。这一普通小麦抗白粉病种质资源的创制及其连锁分子标记的建立为小麦抗病基因分子标记辅助选择、基因积聚和分子育种提供了新的物质基础。     

Resistance to Septoria nodorum blotch in several Triticum species

Summary Resistance to septoria nodorum blotch in Triticum monococcum, T. tauschii, T. timopheevii, T. dicoccum and T. durum was evaluated on plants at the three-leaf stage in greenhouse tests. A high frequency of resistant genotypes was found in T. monococcum, T. tauschii and T. timopheevii, but not in T. dicoccum and T. durum. The resistance of F₁ plants of crosses of resistant T. monococcum (PI 289599) and T. timopheevii (PI 290518) accessions with susceptible common wheat cv. Park and durum wheat cv. Wakooma, respectively, was evaluated on the basis of percentage leaf necrosis, lesion number, lesion size and incubation period. No dominance was found for long incubation period, but various dominance relationships occurred for low percentage leaf necrosis, low lesion number and small lesion size, depending on the cross. Multiple regression analysis showed that lesion number contributed more to percentage leaf necrosis than lesion size or incubation period. Resistance to septoria nodorum blotch was transferred successfully from T. timopheevii to cultivated durum wheat. Resistant BC₁F₇ lines, recovered from the T. timopheevii (PI 290518) × Wakooma cross, showed normal chromosome behaviour at meiosis (14 bivalents) and were self-fertile. However, an effective level of resistance was not recovered in lines derived from the other interspecific crosses.