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.     

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.     

Growth dependency of wild,primitive and modern cultivated wheat lines on vesicular-arbuscular mycorrhiza fungi

Summary Mycorrhizal colonization and growth dependency were studied at a single rate of phosphorous application in wild and cultivated primitive and modern wheats, inoculated with Glomus intraradices Schenck & Smith. Mycorrhizal colonization found in Triticum timopheevii var. araraticum (AAGG) was higher than that found in the other tetraploid wheats (AABB). Mycorrhizal dependency was higher in representatives of the D genome donor — Aegilops squarrosa, compared with representatives of the A and possible B genome donors T. monococcum and Ae. sharonensis, Ae. longissima and Ae. speltoides, respectively. The nature of response to VAM in hexaploid wheat was controlled by factors of the A and B genomes which are epistatic over those located in the D genome. The high mycorrhizal colonization and dependency which was found in T. timopheevii var. araraticum may indicate special genomic affinity possessed by the G genome of wheat in VAM interaction. Based on the 27 wheat lines and species tested in this study only low correlation between G. intraradices colonization and its contribution to plant growth can be suggested.     

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.     

Monosomic analysis of Russian wheat aphid (Diuraphis noxia) resistance in Triticum aestivum line PI137739

Summary The Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae), has become an important pest of wheat (Triticum aestivum L.) in the United States. The aphid causes a phytotoxemic reaction in wheat evidenced by local and systemic chlorosis and rolling of infested leaves. Developing resistance in wheat cultivars to D. noxia is an essential factor in controlling the damage caused by this pest. Several sources of genetic resistance to D. noxia have been identified in wheat germplasm. Monosomic analysis of the monogenic resistant T. aestivum accession PI137739 has shown that the gene (Dn1) for resistance is carried on chromosome 7D. It appears that chromosome 7B may carry a second resistance gene for D. noxia that might be a source of minor or complementary gene action for resistance.     

Aphid acceptance of Hordeum genotypes is affected by plant volatile exposure and is correlated with aphid growth

Nineteen genotypes of Hordeum vulgare (L.) were screened and characterized as partially resistant or susceptible regarding growth of the bird cherry—oat aphid (Rhopalosiphum padi L.). In a separate test, these same genotypes were treated with volatiles from undamaged plants of barley cultivar Alva. As a result of this treatment, aphid host acceptance (AHA) was significantly affected in seven genotypes, and the magnitude of the effect was positively correlated with aphid growth (AG) in the independent resistance screening test. Changes in AHA induced by volatiles from the same genotype as the volatile receiver were also positively correlated with AG. All the 19 genotypes were also tested as inducers with cultivar Kara as the receiver of volatiles. Five genotypes induced significant reductions in AHA of Kara. The results show that aphids are able to detect changes in responding plants induced by volatiles from another plant. Plant volatile interactions may thus be a component of induced resistance to aphids. These interactions could influence the results of experiments used to select for insect-resistant plants in plant breeding programmes, where normally plant genotypes are mixed at testing.     

Genetic diversity of wheat wild relatives in the Near East detected by AFLP

In order to reveal the molecular genetic diversity of wheat wild relatives, an AFLP analysis was conducted with 16 accessions of five Triticum andAegilops species originating from the Near East. Variation within population was studied with at least seven individuals per accession. Four primer combinations were used for selective amplification. Based on the scored bands, we estimated percentage of polymorphic bands, 1 – proportion of shared bands (1-psb) and nucleotide diversity (π). Of the five species used in this study, Ae. speltoides had the highest level of `within population' variation. This species had also the highest value of the variation among populations. As for Triticum species, the level of variation within population was low in diploid species (T. urartu and T. boeoticum),whereas two tetraploid species (T. dicoccoides and T. araraticum) had relatively high levels of variation within population. While the two diploid Triticum indicated a clear interspecific divergence, the two tetraploid wild wheats were not clearly divergent in this study. The variance portioning analysis indicated that the variation detected for diploid Triticum species was mainly composed of `between species' variation, on the other hand that for tetraploid Triticum was mostly composed of `within population' variation. In conclusion, AFLP analysis reveals molecular variation in all accessions used in this study, suggesting a potential genetic diversity of the wheat wild relatives in natural populations. These results have implications for the design of strategies to maintain genetic diversity within genebank collections. This revised version was published online in August 2006 with corrections to the Cover Date.     

Transfer of rust resistance from Aegilops ovata into bread wheat (Triticum aestivum L.) and molecular characterisation of resistant derivatives

An interspecific cross was made to transfer leaf rust and stripe rust resistance from an accession of Aegilops ovata (UUMM) to susceptible Triticum aestivum (AABBDD) cv. WL711. The F₁was backcrossed to the recurrent wheat parent, and after two to three backcrosses and selfing, rust resistant progenies were selected. The C-banding study in a uniformly leaf rust and stripe rust resistant derivative showed a substitution of the 5M chromosome of Ae. ovata for 5D of wheat. Analysis of rust resistant derivatives with mapped wheat microsatellite makers confirmed the substitution of 5M for 5D. Some of these derivatives also possessed one or more of the three alien translocations involving 1BL, 2AL and 5BS wheat chromosomes which could not be detected through C-banding. A translocation involving 5DSof wheat and the substituted chromosome 5M of Ae. ovata was also observed in one of the derivatives. Susceptibility of this derivative to leaf rust showed that the leaf rust resistance gene(s) is/are located on short arm of 5M chromosome of Ae. ovata. Though the Ae. ovatasegment translocated to 1BL and 2AL did not seem to possess any rust resistance gene, the alien segment translocated to 5BS may also possess gene(s) for rust resistance. The study demonstrated the usefulness of microsatellite markers in characterisation of interspecific derivatives. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Antibiosis and antixenosis to Rhopalosiphum padi among triticale accessions

Tests for antibiosis and antixenosis resistance to Rhopalosiphum padi L., the bird cherry-oat aphid, were conducted among four wheat (Triticum aestivum L.) and eight triticale (XTriticosecale Wittmack) accessions. Tests for antibiosis included measuring R. padi-population growth over 13 days, number of days to reproduction of individual R. padi, and number of aphid progeny produced in the first 7 days of adulthood. Antixenosis was measured in no-choice nymphiposition tests and in choice tests of host selection by winged R. padi. Three of seven triticale accessions limited R. padi populations relative to control accessions. Lower R. padi-population growth on N1185 and Okto Derzhavina could be explained partially by increased developmental times. Lower R. padi-population growth on triticale accessions N1185, N1186 and Okto Derzhavina could be explained at least partially by fewer aphid progeny on these accessions. Developmental time of R. padi on N1185 and Okto Derzhavina was greater than that on Stniism 3 triticale, identified previously as resistant to R. padi. There were less R. padi progeny on N1185 than on Stniism 3, and comparable numbers of R. padi progeny among N1186, Okto Derzhavina, and Stniism 3. None of the accessions limited nymphiposition by R. padi. Choice tests revealed heterogeneity in host selection by R. padi but an overall trend that triticale accessions Okto Derzhavina, N1185, N1186 and Stniism 3 were less preferred hosts than Arapahoe wheat. Relatively strong resistance in these triticale accessions warrant consideration of their future use in breeding programs for cereal-aphid resistance.     

Inheritance of biotype E greenbug resistance in bread wheat CI 17882 and its relationship with wheat streak mosaic virus resistance

Summary Genetic studies were conducted to determine the inheritance of biotype E greenbug resistance in CI 17882 (CI 15092/T. speltoides//Fletcher/3/4^* Centurk), a wheat germplasm line previously released as resistant to wheat streak mosaic virus (WSMV). In addition, the association of greenbug and WSMV resistance in CI 17882 was examined. Results indicated that biotype E greenbug resistance in CI 17882 is conditioned by a single dominant gene that is not linked with the WSMV resistance gene.Cooperative research of the USDA, Agricultural Research Service and the Oklahoma Agricultural Experiment Station. Journal article 4845 of the Oklahoma Agric. Exp. Stn., Oklahoma State Univ., Stillwater, OK 74078.     

Wheat pre-breeding using wild progenitors

To facilitate the use of wheat wild relatives in conventional breedingprograms, a wheat pre-breeding activity started at ICARDA in 1994/1995season. Preliminary results of gene introgression from wild diploidprogenitors, Triticum urartu, T. baeoticum, Aegilops speltoides andAe. tauschii and tetraploid T. dicoccoides are described. Crosseswith wild diploid Triticum spp. yielded high variation in plant andspike morphology. Synthetic hexaploids were produced from crosses of alocal durum wheat landrace `Haurani' with two Ae. tauschiiaccessions. Both Ae. tauschii accessions carry hybrid necrosis allelesthat gave necrotic plant phenotypes in crosses with some bread wheats.Backcross progenies with agronomical desirable traits, i.e. high spikeproductivity, short plant stature, earliness, drought tolerance and highproductive tillering, were identified in crosses of durum wheat with wild Triticum spp. and in a cross of one of the hexaploid synthetics with alocally adapted bread wheat cv. `Cham 6'. Resistance to yellow rust wasfound in durum wheat crosses with the three wild Triticum spp. andAe. speltoides and leaf rust resistance was identified in crosses withT. baeoticum and Ae. speltoides. The results show that wheatimmediate progenitors may be a valuable and readily accessible source ofnew genetic diversity for wheat improvement.     

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.     

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.     

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.     

Screening for resistance against Rhopalosiphum padi (L.). 2. Hordeum species and interspecific hybrids

Summary Twenty-seven accessions of Hordeum species and interspecific hybrids were screened in a growth chamber for resistance against the aphid Rhopalosiphum padi. Resistance measurement was based on the aphids' population growth during a 16-day period. The mean number of nymphs produced per female was only about 14% of that on the control cv. Tellus. H. bogdani was the most resistant accession. As a group, interspecific F₁-hybrids with common barley (H. vulgare × H. sp.) held an intermediate position. Differences in resistance between hybrids as well as between species (H. jubatum and H. lechleri) were observed. The most resistant species were all diploids. No effects of the host plant on formation of alatae or on the within-plant distribution of aphids were found.     

Utilisation of <Emphasis Type="Italic">Aegilops</Emphasis> (goatgrass) species to widen the genetic diversity of cultivated wheat

Wild Aegilops species related to cultivated wheat (Triticum spp.) possess numerous genes of agronomic interest and can be valuable sources of resistance to diseases, pests and extreme environmental factors. These genes can be incorporated into the wheat genome via intergeneric crossing, following, where necessary, the development of chromosome addition and substitution lines from the resulting hybrids. The transfer of a single segment from an alien chromosome can be achieved by translocations. The Aegilops (goatgrass) species, which are the most closely related to wheat, exhibit great genetic diversity, the exploitation of which has been the subject of experimentation for more than a century. The present paper gives a survey of the results achieved to date in the field of wheat–Aegilops hybridisation and gene transfer. The Aegilops genus consists of 11 diploid, 10 tetraploid and 2 hexaploid species. Of these 23 Aegilops species, most of the diploids (Ae. umbellulata Zhuk., Ae. mutica Boiss., Ae. bicornis (Forssk.) Jaub. & Spach, Ae. searsii Feldman & Kislev ex Hammer, Ae. caudata L., Ae. sharonensis Eig, Ae. speltoides Tausch, Ae. longissima Schweinf. & Muschl.) and several polyploids (Ae. ventricosa Tausch, Ae. peregrina (Hack. In J. Fraser) Marie & Weiller, Ae. geniculata Roth, Ae. kotschyi Boiss., Ae. biuncialis L.) have been used to develop wheat–Aegilops addition lines. Wheat–Aegilops substitution lines were developed using several species, including Ae. umbellulata, Ae. caudata, Ae. tauschii, Ae. speltoides, Ae. sharonensis, Ae. longissima and Ae. geniculata. Translocations carrying genes responsible for useful agronomic traits were developed with Ae. umbellulata, Ae. comosa, Ae. ventricosa, Ae. longissima, Ae. speltoides and Ae. geniculata. A large number of genes were transferred from Aegilops species to cultivated wheat, including those for resistance to leaf rust, stem rust, yellow rust and powdery mildew, and various pests (cereal cyst nematode, root knot nematode, Hessian fly, greenbug). Many molecular markers are linked to these resistance genes. The development of new molecular markers is also underway. There are still many untapped genetic resources in Aegilops species that could be used as resistance sources for plant breeding.