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.     

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.     

Location of genes controlling resistance to greenbug (Schizaphis graminum Rond.) in Hordeum chilense

Wheat/Hordeum chilense disomic addition lines have been used to locate genes influencing resistance against greenbug (Schizaphis graminum Rond.) in specific chromosomes of H. chilense. H. chilense is a source of antixenosis, antibiosis and host tolerance to the greenbug, being resistant also to the Russian wheat aphid, the two key pests in wheat. For measuring antixenosis, the numbers of aphids per plant were recorded in a host free choice test; antibiotic resistance was determined by measuring the developmental time, the fecundity and the intrinsic rate of population increase of aphids reared on the different hosts, and host tolerance to aphids was evaluated by the leaf damage and the number of expanded leaves on the hosts after 3 weeks of infestation. The greenbugs belonged to a clone of biotype C. Plant genes with positive effects for antixenosis were located on chromosome 1H^ch. Genes with positive effects for antibiosis were located on three different chromosomes and those that prolonged aphid developmental time were located on chromosomes 5H^ch and 7H^ch while those that reduced the total fecundity were on 4Hch. Chromosome 7H^ch accounted for host tolerance to greenbug.     

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.     

Identification of Aegilops germplasm with multiple aphid resistance

The greenbug, Schizaphis graminum(Rondani), the Russian wheat aphid, Diuraphis noxia (Mordvilko), and the bird cherry oat aphid, Rhopalosiphum padi(L.), annually cause several million dollars worth of wheat production losses in Europe and the United States. In this study, Triticum and Aegilops accessions from the Czech Research Institute of Crop Production and the Kansas State University Wheat Genetic Resources Center were evaluated for resistance to these aphids. Accessions with aphid cross-resistance were examined for expression of the antibiosis, antixenosis, and tolerance categories of resistance. Aegilops neglecta accession 8052 exhibited antibiotic effects toward all three aphids in the form of reduced intrinsic rate of increase (r_m). The r_m of greenbug (biotype I) on Ae. neglecta 8052 was significantly lower than that of greenbugs on plants of the susceptible U. S. variety Thunder bird. The r_m of Russian wheat aphids was significantly lower on foliage of both Ae. neglecta 8052 and T. araraticum accession 168 compared to Thunderbird. The r_m values of bird cherry oat aphids fed both Ae. neglecta 8052 and T. araraticum 168 were also significantly lower than those fed the susceptible accession T. dicoccoides 62. Neither Ae. neglecta 8052 or T. araraticum 168 exhibited tolerance to either greenbug biotype I or Russian wheat aphid. Preliminary data suggest that T. araraticum 168 may also possess tolerance to bird cherry oat aphid. New genes from Ae. neglecta 8052 and T. araraticum 168 expressing aphid antibiosis can be used to develop multiple aphid resistant wheat in the U. S. and Central Europe. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mapping antixenosis genes on chromosome 6A of wheat to greenbug and to a new biotype of Russian wheat aphid

Greenbug and Russian wheat aphid (RWA) are two devastating pests of wheat. The first has a long history of new biotype emergence and recently. RWA resistance has just started to break down. Thus, it is necessary to find new sources of resistance that will broaden the genetic base against these pests in wheat. Seventy‐five doubled haploid recombinant (DHR) lines for chromosome 6A from the F₁ of the cross between “Chinese Spring’ and the “Chinese Spring (Synthetic 6A) (Triticum dicoccoides × Aegilops tauschii)” substitution line were used as a mapping population for testing resistance to greenbug biotype C and to a new strain of RWA that appeared in Argentina in 2003. A quantitative trait locus (QTL) (br antixenosis to greenbug was significantly associated with the marker loci Xgwm1009 and Xgwm1185 located in the centromere region of chromosome 6A. Another QTL which accounted for most of the antixenosis against RWA was associated with the marker loci Xgwm1291 and Xiinni1150. both located on the long arm of chromosome 6A. This is the first report of greenbug and RWA resistance genes located on chromosome 6A. It is also the first report of antixenosis against the new strain of RWA. As most of the RWA resistance genes present in released cultivars have been located in [he D‐ genome, it is highly desirable to find new sources in other genomes to combine the existing resistance genes with new sources.     

Level and Nature of the Resistance to the Cereal Aphid, Sitobion avenae (F.), in Thirteen Winter Wheat Cultivars

Thirteen winter wheat cultivars were tested under field and greenhouse conditions for resistance to the cereal aphid, Sitobion avenae (F.). Marked differences were recorded between cultivars in nonpreference (anti-xenosis), antibiosis and tolerance to aphids. Nonpreference was significantly (P < 0.001) correlated with antibiosis measured in terms of the biomass of aphids on naturally and nonsignificantly (P > 0.1) on artificially infested ears (criterion for cultivar antibiosis). Over all the cultivars, there was no correlation between nonpreference and tolerance and between antibiosis and tolerance to aphids.
Five cultivars expressed high level of resistance to S. avenae . The high resistance of three cultivars was due to high nonpreference and antibiosis, and only in two cultivars all three mechanisms contributed to resistance.     

Mapping resistance genes conferring tolerance to RWA (Diuraphis noxia) in barley (Hordeum vulgare)

The Russian wheat aphid (RWA) is one of the most aggressive pests of barley and wheat. The outbreak of RWA occurred in Argentina in 2008 caused serious damage to barley cultivars. The most effective and sustainable method of RWA control is to identify new resistance genes. The purpose of the current research was to map RWA resistance genes in a set of double haploid (DH) lines of the Oregon-Wolfe Barley (OWB) mapping population derived from the cross between OWB_DOM and OWB_REC. The DH and both parental lines were screened for antixenosis, tolerance and antibiosis to RWA. There was significant variation among the DH lines in most of the traits studied. However, only tolerance resulted in significant quantitative trait loci (QTLs) associated with the molecular markers. Two main QTLs were identified. These explained 90 and 79 % of the variability of foliar area and chlorophyll content, respectively, of infested and control plants. The initial and final foliar area and the variation in foliar area were associated with the same molecular markers on chromosome 2H (BmAc0125, Vrs1, BmAc0144f and BmAg0113e). The positive alleles were provided by OWB_DOM. The content of chlorophyll was associated with the marker loci WMC1E8, MWG912, ABC261, MWG2028 and Blp on chromosome 1H, with the positive alleles provided by OWB_REC. Both parents contributed to different tolerance traits, with foliar area and chlorophyll content remaining as the plant traits most affected by aphid feeding. The QTLs found in this population are new RWA resistance loci. A sequence homology search was performed to derive the putative function of the genes linked to the QTLs.     

Screening for Greenbug Resistance in Hordeum chilense Roem et Schult.

Schizaphis graminum (Rondani) is a serious pest of cultivated wheat (Triticum aestivum L.) and resistance is only available in other related species such as Hordeum chilense. Amphiploids between H. chilense and Triticum spp. have been obtained, and addition lines of H. chilense in wheat have been developed. Thirty-five accessions of H. chilense were screened to identify greenbug antixenosis, antibiosis and tolerance. Antixenosis was determined in a conventional host free choice test; antibiosis was measured by aphid life cycle and fecundity rate, and tolerance was tested in a conventional infestation test of 4 weeks. Two commercial barley cultivars were used as susceptible and resistant controls. Eight H. chilense accessions showed higher degrees of antixenosis than the resistant check, 19 were similar and the rest were lower. All accessions were more resistant than the susceptible check. Measured by aphid life cycle, 22 H. chilense accessions showed higher antibiosis than the resistant cv., and all exhibited a higher antibiotic effect on fecundity rates than the control. A similar degree of tolerance to that of the resistant control was observed in six accessions, the remaining entries ranged between the controls. The presence of one mechanism did not exclude the existence of other mechanisms in the same entry and therefore, independence of the different mechanisms is proposed. Most of the accessions showed higher variability than both controls for the three mechanisms, and it appears to be genetic variability within entries for the three mechanisms.     

Genetic variability in antibiotic resistance to the greenbug Schizaphis graminum in Hordeum chilense

Hordeum chilense (Roem. et Schult), a native grass of South America, is a source of antixenotic and antibiotic resistance to the greenbug Schizaphis graminum Rondani. The genetic and environmental components of the variability in antibiotic resistance shown by H. chilense were determined by measuring the developmental time, the length of adult life, fecundity and intrinsic rate of population increase of green-bugs reared on this host. The aphids belonged to a clone of biotype C. Plants were cloned to reduce the incidence of environmental variability. Different plant characters appeared to prolong aphid developmental time and reduce the length of adult life and total fecundity. The broad sense heritability and the genetic variability of these plant characters were different. The plant character that affected aphid development appears to differ from that affecting fecundity.     

Resistance to Brevicoryne brassicae in horticultural brassicas

Summary A wide range of cultivated brassica accessions including broccoli, Brussels sprouts, Chinese cabbage, cauliflower, collard, kale and swede material was tested against the cabbage aphid, Brevicoryne brassicae, at HRI, Wellesbourne in the field and laboratory in both 1992 and 1993. In the field, B. brassicae attack was assessed as the proportion of infested plants and the numbers of aphid colonies present. In the laboratory, aphid performance was measured in terms of the pre-reproductive and reproductive period, population increase, and insect mortality. Interpretation of the data was facilitated by plotting sorted accession means against normal order statistics. This statistical approach indicated the spread of variation amongst the accessions and permitted identification of extremes. Partial levels of antixenosis resistance were discovered in red brassicas. Glossy accessions of cabbage and cauliflower possessed antixenosis and antibiosis resistance that lasted throughout the season of crop growth in the field. Other accessions were shown to withstand aphid attack and therefore possessed tolerance. The laboratory studies provided information on mechanisms of antibiosis resistance. The potential value of the different sources of resistance is discussed.     

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.     

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.     

Cytology, Fertility and Morphology of Amphiploids Hordeum chilense × Tetraploid Wheats (Tritordeum)

With the aim of widening the genetic variability of hexaploid tritorceum through the wheat parents, amphiploids between Hordeum chilense and Triticum turgidum ssp. dicoccoides, ssp. georgicum, and Cody, polonicum have been synthesized. The meiotic behaviour and the fertility were examined in these amphiploids. The morphology of the amphiploids in comparison to their wheat parents was described.     

Expression of barley yellow dwarf virus and Russian wheat aphid resistance genes in and fertility of spring wheat × triticale hybrids and backcross lines

Summary The Barley Yellow Dwarf Virus disease (BYDV) and the Russian wheat aphid (RWA) Diuraphis noxia (Mordvilko) have caused significant losses to wheat and barley in several areas of the world. Important sources of resistance to both BYDV and RWA have been found in Triticale. Different generations of interspecific wheat x Triticale crosses were produced and the progenies were screened for BYDV and RWA tolerance. Plants with equal chromosome numbers showed different levels of fertility. A significant correlation was observed between pollen fertility and seed set in primary florets (r=0.57). In generaL, pollen fertility, seed set and the number of euploid plants (2n=42) increased from one generation to the next. The expression of BYDV tolerance varied from population to population. Additive effects were predominant in F₁ and some backcross populations. A dominant effect of rye tolerance genes was also observed in few populations. A monogenic trait or a quantitative (polygenic) character would not agree with the observed segregation patterns. The heritability of this oligogenic tolerance was quite different between populations and in many populations the tolerance genes were only partially expressed. Some transgressive segregation for tolerance and sensitivity was demonstrated. The genes controlling tolerance to RWA in Triticale lines, Muskox 658 and Nord Kivu were not expressed in advanced lines resistant to BYDV. This indicates that tolerance genes for BYDV and RWA in these lines are located on different chromosomes.     

Identification of rye genotypes resistant to biotypes B,C, E,and F of the greenbug

Summary The greenbug, Schizaphis graminum (Rondani), is a serious pest of wheat, Triticum aestivum L., and other small grains. Cultivar resistance would be an efficient means of control. Unfortunately, a paucity of greenbug resistance in wheat germplasm and occurrence of new virulent biotypes of the greenbug have made development of resistant cultivars difficult. Therefore, resistance genes are sought in species related to and crossable with wheat. Our objective was to evaluate, in greenhouse seedling tests, 11 rye (Secale cereale L.) accessions for their reaction to greenbug biotypes B, C, E, and F. Two ryes, CI 187 and PI 240675, segregated for resistance to all four biotypes. It may be possible to transfer this resistance to wheat. These resistance sources may also be of importance in rye and triticale (X Triticosecale Wittmack) breeding.     

Genetic mapping and inheritance of Russian wheat aphid resistance gene in accession IG 100695

The Russian wheat aphid (RWA), Diuraphis noxia (Kurdjumov), is an important pest of small‐grain cereals, particularly wheat, worldwide. The most efficient strategy against the RWA is to identify sources of resistance and to introduce them into susceptible wheat genotypes. This study was conducted to determine the mode of inheritance of the RWA resistance found in ICARDA accession IG 100695, to identify wheat microsatellite markers closely linked to the gene and to map the chromosomal location of the gene. Simple sequence repeat (SSR) marker scores were identified in a mapping population of 190 F₂ individuals and compared, while phenotypic screening for resistance was performed in F_2 : 3 families derived from a cross between ‘Basribey’ (susceptible) and IG 100695 (resistant). Phenotypic segregation of leaf chlorosis and rolling displayed the effect of a single dominant gene, temporarily denoted Dn100695, in IG 100695. Dn100695 was mapped on the short arm of chromosome 7D with four linked SSR markers, Xgwm44, Xcfd14, Xcfd46 and Xbarc126. Dn100695 and linked SSR markers may be useful for improving resistance for RWA in wheat breeding.     

Genetic control of Russian wheat aphid (Diuraphis noxia) resistance in wheat accession PI 47545

The Russian wheat aphid, Diuraphis noxia (Mordvilko), is a major pest of cereal crops in many areas of the world, causing serious reduction in grain yield in wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.). Incorporating genetic resistance to D. noxia into wheat cultivars is paramount to effectively reduce damage inflicted by this pest. Genetic resistance to D. noxia has been identified in wheat, barley and rye germplasm, and several resistance genes are available for use for cultivar improvement. In the United States of America, only a few Russian wheat aphid (RWA) resistant winter wheat cultivars are currently available, and these cultivars contain only one of the six known RWA resistance genes. The objective of this study was to determine the inheritance of RWA resistance in wheat accession PI 47545, using a screening method based on differences in the leaf morphology of resistant and susceptible types following insect challenge. PI 47545 was selected for study, since it displayed high levels of resistance in a white-grained wheat background, the predominant wheat class produced in the Pacific Northwest of the USA. Segregation analysis was conducted on an F₂ population developed by cross-hybridizing the susceptible soft white winter wheat cultivar ‘Daws’ to the resistant accession PI 47545. Russian wheat aphid screening data from this population indicated that the resistance in PI 47545 is controlled by a single, dominant gene (χ² = 1.72; p ≤ 0.189). This revised version was published online in August 2006 with corrections to the Cover Date.     

Russian wheat aphid-induced protein alterations in spring wheat

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), has become a perennial, serious pest of wheat (Triticum aestivum L.) in the western United States. Current methodologies used to enhance RWA resistance in wheat germplasm could benefit from an understanding of the biochemical mechanisms underlying resistance to RWA. This study was initiated to identify specific polypeptides induced by RWA feeding that may be associated with RWA resistance. The effects of RWA feeding on PI 140207 (a RWA-resistant spring wheat) and Pavon (a RWA-susceptible spring wheat) were examined by visualizing, silver-stained denatured leaf proteins separated by two-dimensional polyacrylamide gel electrophoresis. Comparisons of protein profiles of noninfested and RWA-infested Pavon and PI 140207 revealed a 24-kilodalton-protein complex selectively inhibited in Pavon that persisted in PI 140207during RWA attack. No other significant qualitative or quantitative differences were detected in RWA-induced alterations of protein profiles. These results suggest that RWA feeding selectively inhibit synthesis and accumulation of proteins necessary for normal metabolic functions in susceptible plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.