Mapping quantitative trait loci in wheat for resistance against greenbug and Russian wheat aphid

Breeding for genetic resistance against greenbug and Russian wheat aphid (RWA) is the most effective way of controlling these widespread pests in wheat. Earlier work had shown that chromosome 7D of a synthetic hexaploid wheat, ‘Synthetic’ (T. dicoccoides × Ae. squarrosa) (AABB × DD) gave resistance when transferred into the genetic background of an aphid‐susceptible cultivar, ‘Chinese Spring’, as the recipient. To map the genes involved, a set of 103 doubled haploid recombinant substitution lines was obtained from crossing the 7D substitution line with the recipient, and used to determine the number and chromosomal location of quantitative trait loci (QTL) controlling antixenosis and antibiosis types of resistance. Antixenosis to RWA was significantly associated with marker loci Xpsr687 on 7DS, and Xgwm437 on 7DL. Antibiosis to greenbug was associated with marker loci Xpsr490, Rc3 (on 7DS), Xgwm44, Xgwm111, Xgwm437, Xgwm121 and D67 (on 7DL). Similarly, antibiosis to RWA was linked to loci Xpsr490, Rc3, Xgwm44, Xgwm437 and Xgwm121. At least two QTL in repulsion phase, one close to the centromere either on the 7DS or 7DL arms, and a second distal on 7DL could explain antibiosis to RWA and, partially, this mechanism against greenbug.     

Resistance in wheat to a new North American–Russian wheat aphid biotype

The Russian wheat aphid (RWA), is a serious threat to wheat production worldwide. The identification of a new RWA biotype in the USA virulent to all commercially grown winter wheats poses new challenges to wheat breeders. Wheat germplasm was evaluated to identify accessions resistant to the new virulent RWA isolate (biotype 2). Eleven biotype 1‐resistant wheats and one susceptible check were challenged with RWA biotype 2. Two resistant wheat entries were identified (one highly resistant and one moderately resistant). This information is useful to wheat breeders searching for sources of resistance to the new RWA biotype to incorporate into their breeding programmes.     

Different types of resistance against greenbug, Schizaphis graminum Rond, and the Russian wheat aphid, Diuraphis noxia Mordvilko, in wheat

A collection of 26 cultivars of wheat Triticum aestivum were screened for resistance against the two main aphid pests of cereals, the greenbug Schizaphis graminum Rond. and the Russian wheat aphid (RWA) Diuraphis noxia Mordvilko. Since genetic variability has been found in Argentinean populations of both aphid species, this work was aimed at determining the response of different types of resistance in wheat cultivars when infested with aphids. Antixenosis, antibiosis and tolerance were evaluated with traditional tests in controlled environmental conditions using a clone of greenbug biotype C and a clone of RWA collected on wheat. Genetic resistance was found against one or both aphid species in several wheats. Most of the highest levels of antixenosis, antibiosis and tolerance against the two aphids occurred in different cultivars; as a consequence the resistance mechanisms for both pests appear to be partly independent. Antibiosis against greenbug or RWA appears to be determined by two different sets of genes, one affecting development time and the other reducing fecundity and longevity. The antibiosis against both aphid species in terms of their development time and the intrinsic rate of population increase resulted in a partial cross effect of these aphid traits against the alternative insect species. Nonetheless, the same cultivars affected the total fertility and the longevity of both aphids. Since the highest plant performance levels and the least plant damage were recorded in different wheats, different patterns of tolerance were displayed against the greenbug and the RWA. Consequently, different genes appear to be involved in several traits of the resistance mechanisms against the two aphids. The genes that independently conferred resistance to aphids could be combined in new cultivars of wheat to broaden their genetic base of resistance against the greenbug and the RWA.     

Resistance against greenbug, Schizuphis graminum Rond., and Russian wheat aphid, Diuraphis noxia Mordvilko, in tritordeum amphiploids

A collection of tritordeum amphiploids (Hordeum chilense × Triticum turgadum) and their wheat parents were screened for resistance against the two main aphid pesis of cereals, the greenhug. Schizaphis graminum Rond. and ihe Russian wheat aphid (RWA) Diuraphis naxia Mord-vilko. Antixenosis. antibiosis and tolerance were evaluated in controlled environmental conditions using a. clone of greenbug biotypc C and a clone of RWA collected on pasta wheat. Tritordeum amphiploids pos-sess genetic resistance against greenbug and RWA; some of the lines tested were more resistant than the parental wheat line. Four principal components explained the resistance against both aphid species. The antixenosis shown against both pests was mainly contributed by their wheat parents. The antibiosis againsl both aphid species was obviously dependent on diflerent plant traits. The highest levels of antibiosis against the two aphids occurred in different amphiploids. Different genes are involved in the antibiotic reaction against the two aphids. The Tritordeum resistance to RWA is based on anlixenosis and ant-biosis since the tolerance trails were not independent of the other types of resistance. The level of tolerance shown to the greenbug was variable and appears to be controlled by differeni mechanisms. The tolerance to aphids shown by H. chilense is expressed in the amphiploids. but with some genomic interaction. Genes conferring resistance to aphids in H. chilensee could be incorporated into new cultivars of wheat to broaden their genetic base of resistance against greenbug and RWA.     

DIBOA in wild Poaceae: Sources of resistance to the Russian wheat aphid (Diuraphis noxia) and the greenbug (Schizaphis graminum)

Hydroxamic acids are secondary metabolites typical of Poaceae that play a role in cereal resistance against pests and pathogens. The content of the hydroxamic acid aglucones DIBOA (2,4-dihydroxy-1,4-benzoxazin-3-one) and DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) was evaluated in wild Poaceae belonging to the tribes Triticeae (genera Hordeum and Elymus) and Aveneae (genera Deschampsia and Phalaris). The concentration of DIBOA in seedling extracts of the wild barleys Hordeum chilense, H. brevisubulatum subsp. violaceum and H. bulbosum was negatively correlated with parameters related to performance of the cereal aphids Schizaphis graminum and Diuraphis noxia. The relevance of the results obtained for breeding programs aimed at increasing cereal resistance against aphids is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of Russian wheat aphid resistance in PI 140207 spring wheat

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), has become a serious, perennial pest of wheat (Triticum aestivum L.) in many areas of the world. This study was initiated to determine the inheritance of RWA resistance in PI 140207 (a RWA-resistant spring wheat) and to determine its allelic relationship with a previously reported RWA resistance gene. Crosses were made between PI 140207 and ‘Pavon’ (a RWA-susceptible spring wheat). Genetic analysis was performed on the parents, F₁, F₂, backcross (BC) population and F₂-derived F₃ families. Analyses of segregation patterns of plants in the F₁, F₂, and BC populations, and F₂-derived F₃ families indicated single dominant gene control of RWA resistance in PI 140207. Results of the allelism test indicated that the resistance gene in PI 140207, while conferring distinctly different seedling reactions to RWA feeding, is the same as Dn 1, the resistance gene in PI 137739.     

The use of low molecular weight glutenin subunits to distinguish between wheat cultivars with and without resistance to the Russian wheat aphid, Diuraphis noxia

A number of resistance sources for the Russian wheat aphid have been reported in the last few years and were used to develop resistant cultivars from current commercial cultivars in various breeding programmes. It can be diffcult to distinguish between the cultivars with and without resistance without actual infestation and so in this study we looked at low molecular weight glutenin subunits (LMW-GS) of the two groups. Distinctly different banding patterns were found for the cultivars tested and their isogenic counterparts. Although the LMW-GS and DN1 and DN5 are coded on different chromosomes, the LMW-GS are highly repeatable and banding profiles of each cultivar can be used for the identification of unknown seed.     

Inheritance and allelism of Russian wheat aphid resistance in several wheat lines

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko) has caused serious reduction in wheat production in 17 Western states of the United States since 1986. Inheritance of resistance to RWA in seven wheat lines and the allelism of the resistance genes in these lines with three known resistance genes Dn4, Dn5, and Dn6 were studied. The seven resistant lines were crossed to a susceptible wheat cultivar ‘Carson’ and three resistant wheats: CORWA1 (Dn4), PI 294994 (Dn5), and PI 243781 (Dn6). Seedlings of the parents, F₁, and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21–28 days after the infestation using a 1–9 scale. The resistance level of all the F₁ hybrids was similar to that of the resistant parent, indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ population, suggesting qualitative, nonadditive gene action, in which the presence of any one of the dominant alleles confers complete resistance to RWA. Resistance in CI 2401 is controlled by two dominant genes. Resistance in CI 6501 and PI 94365 is governed by one dominant gene. Resistance in PI 94355 and PI 151918 may be conditioned by either one dominant gene or one dominant and one recessive gene. No conclusion can be made on how many resistance genes are in AUSVA1-F₃, since the parent population was not a pure line. Allelic analyses showed that one of resistance genes in CI 2401 and PI 151918 was the same allele as Dn4, the resistance gene in CI 6501 was the same allele as Dn6, and AUS-VA1-F₃ had one resistance gene which was the same allele as one of the resistance genes in PI 294994. One non-allelic resistance gene different from the Dn4, Dn5, and Dn6 genes in CI 2401, PI 94355, PI 94365, and PI 222668 was identified and should be very useful in diversifying gene sources in wheat breeding.     

Multigenic inheritance of biotype-E greenbug resistance in wheat

Relatively little is known about host-plant insect-resistance genes. Near-isogenic lines (NILs) can be a useful tool in evaluating such genes and their mechanisms of action. Through backcrossing and single-seed selection, NILs have been produced in wheat (Triticum aestivum L.) for resistance/susceptibility to biotype-E greenbug, Schizaphis graminum (Rondani). Crosses among several resistant and susceptible NILs produced segregation patterns in the F₂ generation that were inconsistent with simple monogenic inheritance. The simplest genetic hypothesis consistent with the data would involve two independently segregating loci, requiring complementary expression of multiple resistance alleles. F₃ segregation data were also consistent with the proposed genetic hypothesis. Multiple alleles are required to account for the observed variation, although the source of the multiple alleles is unknown. Direct comparison of specific resistant/susceptible pairs of NIL's, as well as bulked segregant analysis, should permit elucidation of resistance mechanisms. Based on the segregation data presented, new designations for the resistance loci studied are proposed.     

Inheritance of resistance to the Russian wheat aphid in an Iranian durum wheat line

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), is a major economic pest of small grains in many countries. An experiment was therefore conducted to determine the inheritance of gene(s) controlling resistance to RWA in a resistant tetraploid durum wheat line. This resistant line,‘1881′, was crossed to a susceptible line, ‘Orejy‐e‐Kazeroon’, and then F₁ F₂ and BCF₁ (backcross to susceptible line) seedlings were screened in a greenhouse for RWA resistance following artificial infection. Resistance in ‘1881’ was apparently controlled by one dominant gene. Since Dnl, Dn2, dn3, Dn4 and Dn5 have been reported to be located on genome D, it was reasoned that the resistance gene in ‘1881’ is not allelic to them.     

Amplified fragment length polymorphism- and simple sequence repeat-based molecular tagging and mapping of greenbug resistance gene Gb3 in wheat

The greenbug, Schizaphis graminum (Rondani), is the most economically damaging aphid pest of wheat in the southern Great Plains of the USA. In this study, the single, dominant greenbug resistance gene, Gb3, was molecularly tagged and genetically mapped using amplified fragment length polymorphism (AFLP) and simple sequence repeat(SSR) markers. Three AFLP loci were associated with the Gb3 locus in linkage analysis with 75 F_2:3 families from the cross between two near‐isogenic lines (NILs) for Gb3,‘TXGBE273’ and ‘TXGBE281′. Two of these loci, XMgcc Pagg and Xmagg Patg cosegregate with Gb3 in the population analysed. Further analysis indicated that XMgcc Pagg and Xmagg Patg are specific for the Gb3 locus in diverse genetic backgrounds. Two SSR markers, Xgwm111 and Xgwm428 previously mapped in wheat chromosome 7D, were shown to be linked with Gb3, 22.5 cM and 33.1 cM from Gb3, respectively, in an F₂ population of ‘Largo’בTAM 107’, suggesting that Gb3 is located in the long arm of chromosome 7D. The two AFLP markers cosegregating with Gb3 are valuable tools in developing molecular markers for marker‐assisted selection of greenbug resistance in wheat breeding.     

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.     

A Monosomic Analysis of Russian Wheat Aphid Resistance in the Common Wheat PI 294994

Chromosome 7D of PI 294994 was indicated as carrying a single dominant gene for resistance to the Russian wheat aphid. The symbol Dn5 is proposed to designate the gene.     

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.     

QTL mapping and marker-assisted selection for Fusarium head blight resistance in wheat: a review

During the past decade, numerous studies have been published on molecular mapping of Fusarium head blight (FHB) resistance in wheat. We summarize the relevant findings from 52 quantitative trait loci (QTL) mapping studies, nine research articles on marker-assisted selection and seven on marker-assisted germplasm evaluation. QTL for FHB resistance were found on all wheat chromosomes except chromosome 7D. Some QTL were found in several independent mapping studies indicating that such QTL are stable and therefore useful in breeding programmes. We summarize and update current knowledge on the genetics of FHB resistance in wheat resulting from QTL mapping investigations and review and suggest FHB breeding strategies based on the available information and DNA markers.     

Molecular marker analysis of kernel size and shape in bread wheat

The economic value of wheat grain is determined by the kernel morphology which is an important parameter for manufacturing different food products requiring specific grain characteristics. Although kernel size and shape have emerged as important breeding objectives, not much information is available about the number or location of associated gene(s)/quantitative trait loci. In the present study, a recombinant inbred line population of 106 plants (F₇) was phenotyped for four traits, namely kernel length, width, weight and factor form density (FFD) and genotyped with different polymerase chain reaction‐based markers. Transgressive segregants were observed for all the traits and genetic correlation studies showed positive correlations between the majority of the traits. The number of markers associated with each trait ranged from two to nine and the phenotypic contribution by an individual marker ranged from 3.3 to 16.6%. Many of the markers showed linkage to more than one trait. Strategies for improving the wheat grain quality traits and the utility of such markers in marker‐assisted selection (MAS) efforts are discussed.     

The Transfer of Resistance to the Russian Wheat Aphid from Triticum monococcum L. to Common Wheat

Two Triticum monococcum accessions were found to be highly resistant to the Russian wheat aphid. An attempt was made to transfer the resistance to common wheat through direct hybridization or by using bridge species. In each of the four crossing strategies tried, a gradual loss of resistance occurred as hybrids with higher ploidy levels were obtained. It appeared that the level of resistance observed was directly proportional to the ratio of the Triticum monococcum genome relative to other genomes. This would indicate suppression of the resistance gene(s) by the added genomes or dilution of its product(s) by those of homoeoloci. The degree of protection afforded by the gene(s) at the hexaploid level may prove to be small. Plants suspected to be homozygous for the resistance gene(s) were identified; however, further backcrossing to common wheat will be required to improve their agronomic types and meiotic regularities.     

Inheritance and allelism of resistances to the Russian wheat aphid in seven wheat lines

Summary Studies were conducted to determine the inheritance and allelic relationships of genes controlling resistance to the Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), in seven wheat germplasm lines previously identified as resistant to RWA. The seven resistant lines were crossed to a susceptible wheat cultivar Carson, and three resistant wheats, CORWA1, PI294994 and PI243781, lines carrying the resistance genes Dn4, Dn5 and Dn6, respectively. Seedlings of the parents, F₁ and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21 to 28 days after the infestation using a 1 to 9 scale. All the F₁ hybrids had equal or near equal levels of resistance to the resistant parent indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ segregation suggesting major gene actions. The resistance in PI225262 was controlled by two dominant genes. Resistance in all other lines was controlled by a single dominant gene. KS92WGRC24 appeared to have the same resistance gene as PI243781 and STARS-9302W-sib had a common allele with PI294994. The other lines had genes different from the three known genes.     

Identification of wheat chromosomes involved with different types of resistance against greenbug (Schizaphis graminum, Rond.) and the Russian wheat aphid (Diuraphis noxia, Mordvilko)

Two sets of intervarietal chromosome substitution lines in the recipient,susceptible cultivar ‘Chinese Spring’ were screened to identify the wheat chromosomes involved with antixenosis, antibiosis and tolerance resistance to greenbug and Russian wheat aphid. The amphiploid ‘Synthetic’ and the cultivar ‘Hope’ were the donor parents. Antixenosis, antibiosis and tolerance were evaluated with conventional tests in controlled environmental conditions using a clone of greenbug biotype C and a clone of RWA collected on wheat. Antixenosis against greenbug was accounted for by several chromosomes in both sets of substitution lines with chromosome 2B contributing the highest level of this type of resistance. The highest levels of antixenosis against RWA were associated with the group of chromosomes 7 of the substitutions CS/Syn set and the chromosome substitutions 2B, 6A and 7D of the CS/Hope set. Antibiosis against both aphids species was accounted for by several different chromosomes. The highest levels of antibiosis for most of RWA resistance traits were recorded from the 1B substitution line of the CS/Hope set. More than one gene appears to determine antibiosis. Tolerance to both greenbug and the RWA was significantly associated with chromosomes 1A,1D, and 6D in the CS/Syn set of substitutions. These lines showed enhanced plant growth under aphid infestation. The highest levels of antixenosis, antibiosis and tolerance against the two aphid species occurred mostly in different substitution lines. Consequently, the different types of resistance for both pests seem to be partially independent. Since different genes seem to be involved in at least several traits of the resistance categories against the two aphid species, such genes could be combined in new cultivars of wheat to broaden their genetic base of resistance against the greenbug and the RWA. This revised version was published online in July 2006 with corrections to the Cover Date.