RFLP mapping of resistance to chlorosis induction by Pyrenophora tritici-repentis in wheat

Tan spot, caused by Pyrenophora tritici-repentis, is an economically important disease in major wheat production areas. The fungus can produce two genetically distinct symptoms on leaves of susceptible wheat genotypes: tan necrosis (nec) and extensive chlorosis (chl). Our objectives were to determine the number of genes conditioning resistance to tan spot in a population of wheat recombinant inbred lines, and map the chromosomal location of the resistance genes using RFLPs. Conidia produced by the P. tritici-repentis isolate Pti2 (nec+chl+) were used to inoculate seedlings of 135 recombinant inbred lines derived from the cross of the synthetic hexaploid wheat W-7984 with Opata 85. A subset of the population was inoculated with conidia produced by the isolates D308 (nec−chl+) and 86-124 (nec+chl−). Inoculated seedlings were rated on a scale of 1 to 5 based on lesion type. Necrosis-inducing culture filtrate produced by the isolate 86-124 was also used to screen the entire population. A map consisting of 532 markers was employed to identify significant associations between marker loci and tan spot resistance. The entire population was insensitive to culture filtrate produced by the isolate 86-124, and the entire subset was resistant to conidial inoculation of the same isolate. The population segregated for reaction to isolates D308 and Pti2, indicating that this population segregates for resistance to extensive chlorosis only, and not to tan necrosis. RFLP analysis indicated the presence of a gene with a major effect in 1AS, a gene with a minor effect in 4AL, and an interaction between the 1AS gene and a gene in 2DL. Together, these loci explained 49.0% of the variation in this population for resistance to tan spot produced by the isolate Pti2. Two regions one in 1BL and one in 3BL, were significantly associated with resistance to extensive chlorosis, but were not significant in the multiple regression model. It should be feasible to introgress these resistance loci into adapted genetic backgrounds by using a marker-assisted selection scheme. Received: 30 March 1996 / Accepted: 31 May 1996     

Genetic mapping of the powdery mildew resistance gene Pm6 in wheat by RFLP analysis

Pm6 in bread wheat (Triticum aestivum L.), which was transferred from Triticum. timopheevii L., is a gene conferring resistance to the powdery mildew disease caused by Erysiphe graminis f. sp. tritici. Six near-isogenic lines ( NILs ) of Pm6 in a cultivar ’Prins’ background were analyzed to map this gene using restriction fragment length polymorphism (RFLP). Each of the six NILs possessed a T. timopheevii-derived segment, varying in length, and associated with powdery mildew resistance. Lines IGV1–465 (FAO163b/ 7*Prins) and IGV1–467 (Idaed 59B/7*Prins) had the shortest introgressed segments, which were detected only by DNA probes BCD135 and PSR934, respectively. The polymorphic loci detected by both probes were mapped to the long arm of chromosome 2B. Lines IGV1–458 (CI13250/7*Prins) and IGV1–456 (CI12559/8*Prins) contained the longest T. timopheevii segments involving both arms of donor chromosome 2G across the centromere. All these introgressed segments had an overlapping region flanked by the loci xpsr934 and xbcd135 on 2BL. Thus, Pm6 was located in this region since the powdery mildew resistance in all the NILs resulted from the introgressed fragments. Using the F₂ mapping population from a cross of IGV1–463 (PI170914/7*Prins)×Prins, Pm6 was shown to be closely linked to the loci xbcd135 and xbcd266 at a genetic distance of 1.6 cM and 4.8 cM, respectively. BCD135 was successfully used in detecting the presence of Pm6 in different genetic backgrounds. Received: 29 June 1999 / Accepted: 6 July 1999     

Identification of molecular markers linked to the Yr15 stripe rust resistance gene of wheat originated in wild emmer wheat, Triticum dicoccoides

 The Yr15 gene of wheat confers resistance to the stripe rust pathogen Puccinia striiformis West., which is one of the most devastating diseases of wheat throughout the world. In the present study, molecular markers flanking the Yr15 gene of wheat have been identified using the near-isogenic-lines approach. RFLP screening of 76 probe-enzyme combinations revealed one polymorphic marker (Nor/TaqI) between the susceptible and the resistant lines. In addition, out of 340 RAPD primers tested, six produced polymorphic RAPD bands between the susceptible and the resistant lines. The genetic linkage of the polymorphic markers was tested on segregating F₂ population (123 plants) derived from crosses between stripe rust-susceptible Triticum durum wheat, cv D447, and a BC₃F₉ resistant line carrying Yr15 in a D447 background. A 2.8-kb fragment produced by the Nor RFLP probe and a 1420-bp PCR product generated by the RAPD primer OPB13 showed linkage, in coupling, with the Yr15 gene. Employing the standard maximum-likelihood technique it was found that the order OPB13 ₁₄₂₀ –Yr15–Nor1 on chromosome 1B appeared to be no less than 1000-times more probable than the closest alternative. The map distances between OPB13 ₁₄₂₀ –Yr15–Nor1 are 27.1 cM and 11.0 cM for the first and second intervals, respectively. The application of marker-assisted selection for the breeding of new wheat cultivars with the stripe rust resistance gene is discussed. Received: 27 February 1997/Accepted: 7 March 1997     

Comparative analyses of RFLP diversity in landraces of Triticum aestivum and collections of T. tauschii from China and Southwest Asia

 Chinese accessions of Triticum tauschii and T. aestivum L. from the Sichuan white (SW), Yunnan hulled (YH), Tibetan weedrace (TW), and Xinjiang rice (XR) wheat groups were subjected to RFLP analysis. T. tauschii and landraces of T. aestivum from countries in Southwest Asia were also evaluated. For T. tauschii, a west to east gradient was apparent where the Chinese accessions exhibited less diversity than those from Southwest Asia. Compared to the Southwest Asian gene pool, the Chinese T. tauschii was highly homogeneous giving a low frequency of polymorphic bands (16%) and banding patterns (1.33 per probe) with 75 RFLP probe-HindIII combinations. Accessions of T. tauschii from Afghanistan and Pakistan were genetically more similar to the Chinese T. tauschii than those from Iran. Of 368 bands found for 39 Chinese hexaploid wheat accessions with 63 RFLP probe-HindIII combinations, 28.3% were polymorphic with an average of 2.6 banding patterns per probe and 5.0 bands per genotype. The individual Chinese landrace wheat groups revealed less variation than those from Afghanistan, Iran, and Turkey. When classified into country based groups, however, the diversity level over all Chinese landraces was greater than that of some Southwest Asian landraces, especially those from Afghanistan and Iran . The XR wheat group was genetically distinct from the other three Chinese landrace groups and was more related to the Southwest Asian landraces. The TW group was genetically similar to, but more diverse than, the SW and YH groups. The Chinese landraces had a higher degree of genetic relatedness to the Southwest Asian T. tauschii, particularly to accessions from Iran, rather than to the Chinese T. tauschii. ‘Chinese Spring’ was most related to ‘Chengdu-guang-tou’, a cultivar from the SW wheat group. Received: 13 May 1997 / Accepted: 19 September 1997     

Isolating individual wheat (Triticum aestivum) chromosome arms by flow cytometric analysis of ditelosomic lines

 We are reporting the successful isolation of wheat chromosome arm 1DS by flow cytometry. A chromosome suspension was prepared for the 1DS ditelosomic line and the normal ‘Chinese Spring’ (CS) by chopping 2-day-old root tip meristems, synchronized by hydroxyurea, in HEPES-magnesium sulfate buffer containing propidium iodide. Chromosomes were analyzed and sorted with a FACS Vantage flow cytometer and cell sorter. An extra peak was observed in the flow karyotype of the ditelosomic line that was absent in ‘CS’. The estimated size of chromosomes from the extra peak matched with the expected size of chromosome 1DS. Chromosomes from the putative 1DS peak were analyzed by both fluorescent microscopy and N-banding analysis. A total of 571 chromosomes from two separate experiments were analyzed, and all were observed to be telosomics except for 2 which were broken. About 82% of these telosomics showed the diagnostic N-band of 1DS, the remaining were unbanded and are probably also 1DS. This strategy can also be used to sort other wheat arms. Received: 30 September 1998 / Accepted: 2 November 1998     

Microclimatic stress and adaptive DNA differentiation in wild emmer wheat, Triticum dicoccoides

 Genetic diversity was examined by RAPD-PCR analysis in 118 registered individuals of wild emmer wheat, Triticum dicoccoides, from a microsite at Yehudiyya, northeast of the Sea of Galilee, Israel. The test involved two climatic microniches in the open oak-park forest of Quercus ithaburensis: (1) sunny between trees and (2) shady under the trees’ canopies. Comparisons were based on 97 loci amplified by 20 oligonucleotide primers. Significant genetic differentiations were found at single-, two- and multilocus structures between the neighbouring shady and sunny niches. These DNA polymorphisms appear to be associated with microclimatic stresses. The pronounced niche-effect on the significance of linkage disequilibrium and niche-specific linkage disequilibrium may suggest that natural selection directed the two-locus associations. The structure of the multilocus associations also mainly results from natural selection, and not by chance from population subdivision, or founder effects. These findings are largely parallel to the previous allozymic results at single-locus and multilocus levels. Both the DNA and the allozymic results suggest that microclimatic selection appears to play an important role in DNA differentiation as well as in protein polymophism. Received: 30 October 1998 / Accepted: 2 November 1998     

Microsatellite polymorphism in natural populations of wild emmer wheat,Triticum dicoccoides,in Israel

Diversity in 20 microsatellite loci of wild emmer wheat, Triticum dicoccoides, was examined in 15 populations (135 genotypes) representing a wide range of ecological conditions of soil, temperature, and water availability, in Israel and Turkey. An extensive amount of diversity at microsatellite loci was observed despite the predominantly selfing nature of this plant species. The 20 Gatersleben wheat microsatellites (GWM), representing 13 chromosomes of genomes A and B of wheat, revealed a total of 364 alleles, with an average of 18 alleles per GWM marker (range: 5–26). The proportion of polymorphic loci per population averaged 0.90 (range: 0.45– 1.00); genic diversity, He, averaged 0.50 (range 0.094– 0.736); and Shannon’s information index averaged 0.84 (range 0.166–1.307). The coefficients of genetic distance between populations were high and averaged D=1.862 (range 0.876–3.320), an indication of sharp genetic divergence over short distances. Interpopulation genetic distances showed no association with geographic distance between the population sites of origin, which ruled out a simple isolation by distance model. Genetic dissimilarity values between genotypes were used to produce a dendrogram of the relationships among wild wheat populations by the unweighted pair-group method with arithmetic averages (UPGMA). The results showed that all the wild emmer wheat populations could be distinguished. Microsatellite analysis was found to be highly effective in distinguishing genotypes of T. dicoccoides, originating from diverse ecogeographical sites in Israel and Turkey, with 88% of the 135 genotypes correctly classified into sites of origin by discriminant analysis. Our present microsatellite results are non-random and in agreement with the previously obtained allozyme and RAPD patterns, although the genetic-diversity values obtained with microsatellites are much higher. Significant correlates of microsatellite markers with various climatic and soil factors suggest that, as in allozymes and RAPDs, natural selection causes adaptive microsatellite ecogeographical differentiation, not only in coding, but most importantly in non-coding genomic regions. Hence, the concept of ”junk DNA” needs to be replaced by at least partly regulatory DNA. The obtained results suggest that microsatellite markers are useful for the estimation of genetic diversity in natural populations of T. dicoccoides and for the tagging of agronomically important traits derived from wild emmer wheat. Received: 27 February 2001 / Accepted: 22 March 2001     

Microsatellite mapping of the induced sphaerococcoid mutation genes in Triticum aestivum

The S1, S2 and S3 genes of the induced sphaerococcoid mutation in common wheat (Triticum aestivum) were mapped using three different F₂ populations consisting of 71–96 individual plants. Twenty-four microsatellite markers from homeologous group 3 of T. aestivum were used to map the S1, S2 and S3 genes on chromosomes 3D, 3B and 3A, respectively. The S1 locus was found to be closely linked to the centromeric marker Xgwm456 of the long arm (2.9 cM) and mapped not far (8.0 cM) from the Xgdm72 marker of the short arm of chromosome 3D. The S2 gene was tightly linked to 2 centromeric markers (Xgwm566, Xgwm845) of chromosome 3B. S3 was located between Xgwm2 (5.1 cM), the marker of the short arm, and Xgwm720 (6.6 cM), the marker of the long arm, both of chromosome 3A. Mapping the S1, S2 and S3 loci of the induced sphaerococcoid mutation near the centromeric regions supports the hypothesis that the sphaerococcum type may be due to gene duplication resulting from DNA recombination in the centromeric region. Received: 20 June 1999 / Accepted: 29 July 1999     

Characterization of spelt (Triticum spelta L.) forms by gel-electrophoretic analyses of seed storage proteins. III. Comparative analyses of spelt and Central European winter wheat (Triticum aestivum L.) cultivars by SDS-PAGE and acid-PAGE

 Seed storage proteins of a few selected spelt forms and crosses have already been electrophoretically analysed by SDS-PAGE and acid-PAGE and compared with a few winter wheat cultivars. In the analyses presented here further important Central European spelt varieties were included, as well as modern winter wheat cultivars which were chosen as standards. In this study gliadin and glutenin band patterns of modern Central European winter wheat cultivars were analysed, in particular for a comparison with spelt varieties. An improved differentiation within and between the two species was obtained. Received: 27 April 1998 / Accepted: 26 May 1998     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.). 5. Alleles at the Pm1 locus

 The chromosomal location and genetic characterization of powdery mildew resistance genes were determined in the common wheat lines MocZlatka, Weihenstephan Stamm M1N and in a resistant line of Triticum aestivum ssp. spelta var. duhamelianum. Monosomic analyses revealed that one major dominant gene is located on chromosome 7A in each of the lines tested. Allelism tests with Pm1 in the backcross-derived line Axminster/8^*Cc on 7A indicated that the resistance genes are alleles at the Pm1 locus. These alleles are now designated Pm1a in line Axminster/8^*Cc, Pm1b in MocZlatka, Pm1c in Weihenstephan Stamm M1N, and Pm1d in T. spelta var. duhamelianum, respectively. Received: 10 November 1997 / Accepted: 29 January 1998     

Cytogenetic and molecular mapping of the leaf rust resistance gene Lr39 in wheat

Leaf rust, caused by the fungus Puccinia triticina Eriks,is one of the most serious diseases of wheat (Triticum aestivum AABBDD, 2n=6x=42) worldwide. Growing resistant cultivars is an efficient and economical method of reducing losses to leaf rust. Here we report a new leaf rust resistance gene, Lr39, transferred from Aegilops tauschii into common wheat. Lr39 conditions both seedling and adult plant resistance to the leaf rust pathogen. The inter- and intra-chromosomal mapping of the Lr39 gene showed that it is different from all previously described Lr genes. We used monosomic analysis for the inter-chromosomal mapping and wheat microsatellite markers for the intra-chromosomal mapping. The monosomic and ditelosomic analysis indicated that Lr39 is independent of the centromere on the short arm of chromosome 2D. Eight microsatellite markers for 2DS were used for linkage analysis on a population of 57 F₂ plants derived from a cross of an Ae. tauschii-derived wheat, cv. Wichita line TA4186 (possessing Lr39), with Wichita monosomics for the D-genome chromosomes. The microsatellite marker analysis confirmed the location of the gene on 2DS. Three markers were polymorphic and linked to the gene. The closest marker Xgwm210 mapped 10.7 cM from Lr39. The location of Lr39 near the telomere of 2DS distinguishes it from the Lr2 and Lr22 loci, which are located on 2DS proximal to Xgwm210. Received: 19 April 2000 / Accepted: 15 May 2000     

Allelopathy in wheat (Triticum aestivum)

Wheat (Triticum aestivum) allelopathy has potential for the management of weeds, pests and diseases. Both wheat residue allelopathy and wheat seedling allelopathy can be exploited for managing weeds, including resistant biotypes. Wheat varieties differ in allelopathic potential against weeds, indicating that selection of allelopathic varieties might be a useful strategy in integrated weed management. Several categories of allelochemicals for wheat allelopathy have been identified, namely, phenolic acids, hydroxamic acids and short‐chain fatty acids. Wheat allelopathic activity is genetically controlled and a multigenic model has been proposed. Research is underway to identify genetic markers associated with wheat allelopathy. Once allelopathic genes have been located, a breeding programme could be initiated to transfer the genes into modern varieties for weed suppression. The negative impacts of wheat autotoxicity on agricultural production systems have also been identified when wheat straws are retained on the soil surface for conservation farming purposes. A management package to avoid such deleterious effects is discussed. Wheat allelopathy requires further study in order to maximise its allelopathic potential for the control of weeds, pests and diseases, and to minimise its detrimental effects on the growth of wheat and other crops.     

Quantitative trait loci mapping of dark-induced senescence in winter wheat (Triticum aestivum)

In order to explore the genetics of dark-induced senescence in winter wheat(Triticum aestivum L.),a quantitative trait loci(QTL)analysis was carried out in a doubled haploid population developed from a cross between the varieties Hanxuan 10(HX)and Lumai 14(LM).The senescence parameters chlorophyll content(Chl a+b,Chl a,and Chl b),original fluorescence(Fo),maximum fluorescence level(Fm),maximum photochemical efficiency(Fv/Fm),and ratio of variable fluorescence to original fluorescence(Fv/Fo)were evaluated in the second leaf of whole three-leaf seedlings subjected to 7 d of darkness.A total of 43 QTLs were identified that were associated with dark-induced senescence using composite interval mapping.These QTLs were mapped to 20 loci distributed on 11 chromosomes:1B,1D,2A,2B,3B,3D,5D,6A,6B,7A,and 7B.The phenotypic variation explained by each QTL ranged from 7.5% to 19.4%.Eleven loci coincided with two or more of the analyzed parameters.In addition,14 loci co-located or were linked with previously reported QTLs regulating flag leaf senescence,tolerance to high light stress,and grain protein content(Gpc),separately.     

Comparative RFLP mapping of Triticum monococcum genes controlling vernalization requirement

 The adaptability of Triticum aestivum to a large range of environments is partially due to genetic differences in sensitivity to vernalization. The most potent gene reducing the vernalization requirement in hexaploid wheat is Vrn-A1. An orthologous vernalization gene, designated Vrn-A ^m 1, was mapped in the diploid wheat Triticum monococcum between RFLP markers Xwg908 and Xabg702 on the long arm of chromosome 5A^mL. The orthology of VrnA ^m 1 with Vrn-A1 (5A wheat, originally Vrn1), Vrn-D1 (5D wheat, originally Vrn3), Vrn-R1 (5R rye, originally Sp1) and Vrn-H1 (5H barley, originally Sh2) was shown by mapping RFLP markers linked to these vernalization genes on the T. monococcum linkage map. A second vernalization gene, designated Vrn-A ^m 2, was found in the distal region of chromosome 5A^mL within a segment translocated from homoeologous group 4. This gene is completely linked to RFLP marker Xbcd402 and located between the same RFLP markers (Xβ-Amy-1 and Xmwg616) as the Vrn-H2 (originally Sh) locus in Hordeum vulgare. Received: 6 January 1998 / Accepted: 31 March 1998     

Genetic polymorphism in low-molecular-weight glutenin genes from Triticum aestivum, variety Chinese Spring

Low-molecular-weight (LMW) glutenin subunits consist mainly of two domains, one at the N- terminus which contains repeats of short amino-acid motifs, and a non-repetitive one rich in cysteine, at the C- terminal region. In previous reports, polyacrylamide-gel electrophoresis has been used to show that large size variation exists among LMW and HMW glutenin subunits, and it has been suggested that deletions and insertions within the repetitive region are responsible for these variations in length. In this study, PCR-amplification of genomic DNA (Triticum aestivum variety Chinese Spring) was used to isolate three full-length LMW glutenin genes: LMWG-MB1, LMWG-MB2 and LMWG-MB3. The deduced amino-acid sequences show a high similarity between these ORFs, and with those of other LMW glutenin genes. Comparisons indicate that LMWG-MB1 has probably lost a 12-bp fragment through deletion and that LMWG-MB1 and LMWG-MB2 have an insertion of 81 bp within the repetitive domain. The current study has shown direct evidence that insertions and/or deletions provide a mechanistic explanation for the allelic variation, and the resultant evolution, of prolamin genes. Single-base substitutions at identical sites generate stop codons in both LMWG-MB2 and LMWG-MB3 indicating that these clones are pseudogenes. Received: 7 May 1999 / Accepted: 17 June 1999     

Gliadin polymorphism in wild and cultivated einkorn wheats

To study the relationships between different species of the Einkorn group, 408 accessions of Triticum monococcum, T. boeoticum, T. boeoticum ssp. thauodar and T. urartu were analyzed electrophoretically for their protein composition at the Gli-1 and Gli-2 loci. In all the species the range of allelic variation at the loci examined is remarkable. The gliadin patterns of T. monococcum and T. boeoticum were very similar to one another but differed substantially from those of T. urartu. Several accessions of T. boeoticum and T. monococcum were shown to share the same alleles at the Gli-1 and Gli-2 loci, confirming the recent nomenclature that considers these wheats as different subspecies of the same species, T. monococcum. The gliadin composition of T. urartu resembled that of the A genome of polyploid wheats more than did T. boeoticum or T. monococcum, supporting the hypothesis that T. urartu, rather than T. boeoticum, is the donor of the A genome in cultivated wheats. Because of their high degree of polymorphism the gliadin markers may help in selecting breeding parents from diploid wheat germ plasm collections and can be used both to search for valuable genes linked to the gliadin-coding loci and to monitor the transfer of alien genes into cultivated polyploid wheats. Received: 8 July 1996 / Accepted: 12 July 1996     

Extended physical maps and a consensus physical map of the homoeologous group-6 chromosomes of wheat (Triticum aestivum L. em Thell.)

Extended physical maps of chromosomes 6A, 6B and 6D of common wheat (Triticum aestivum L. em Thell., 2n=6x=42, AABBDD) were constructed with 107 DNA clones and 45 homoeologous group-6 deletion lines. Two-hundred and ten RFLP loci were mapped, including three orthologous loci with each of 34 clones, two orthologous loci with each of 31 clones, one locus with 40 clones, two paralogous loci with one clone, and four loci, including three orthologs and one paralog, with one clone. Fifty five, 74 and 81 loci were mapped in 6A, 6B and 6D, respectively. The linear orders of the mapped orthologous loci in 6A, 6B and 6D appear to be identical and 65 loci were placed on a group-6 consensus physical map. Comparison of the consensus physical map with eight linkage maps of homoeologous group-6 chromosomes from six Triticeaespecies disclosed that the linear orders of the loci on the maps are largely, if not entirely, conserved. The relative distributions of loci on the physical and linkage maps differ markedly, however. On most of the linkage maps, the loci are either distributed relatively evenly or clustered around the centromere. In contrast, approximately 90% of the loci on the three physical maps are located either in the distal one-half or the distal two-thirds of the six chromosome arms and most of the loci are clustered in two or three segments in each chromosome. Received: 19 April 1999 / Accepted: 28 July 1999     

Salicylic acid changes the properties of extracellular peroxidase activity secreted from wounded wheat (Triticum aestivum L.) roots

Summary.  Wheat (Triticum aestivum L.) roots released proteins showing peroxidase activity in the apoplastic solution in response to wound stress. Preincubation of excised roots with 1 mM salicylic acid at pH 7.0 enhanced the guaiacol peroxidase activity of the extracellular solution (so-called extracellular peroxidase). The soluble enzymes were partially purified by precipitation with ammonium sulfate followed by size exclusion and ion exchange chromatography. Despite an increase in the total activity of secreted peroxidase induced by pretreatment of excised roots with salicylic acid, the specific activity of the partially purified protein was significantly lower compared to that of the control. Purification of the corresponding proteins by ion exchange chromatography indicates that several isoforms of peroxidase occurred in both control and salicylic acid-treated samples. The activities of the extracellular peroxidases secreted by the salicylic acid-treated roots responded differently to calcium and lectins compared with those from untreated roots. Taken together, our data suggest that salicylic acid changes the isoforms of peroxidase secreted by wounded wheat roots. Received June 10, 2002; accepted September 24, 2002; published online May 21, 2003 RID="*" ID="*" Correspondence and reprints: Institute of Biochemistry and Biophysics, Russian Academy of Sciences, P.O. Box 30, Kazan 420111, Russia.     

Antioxidative enzymes and the Russian wheat aphid (Diuraphis noxia) resistance response in wheat (Triticum aestivum)

A crucial function of antioxidative enzymes is to remove excess reactive oxygen species (ROS), which can be toxic to plant cells. The effect of Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), infestation on the activities of antioxidative enzymes was investigated in the resistant (cv. Tugela DN) and the near-isogenic susceptible (cv. Tugela) wheat (Triticum aestivum L.). RWA infestation significantly induced the activity of superoxide dismutase, glutathione reductase and ascorbate peroxidase to higher levels in the resistant than in susceptible plants. These findings suggest the involvement of antioxidative enzymes in the RWA-wheat resistance response, which was accompanied by an early oxidative burst. The results are consistent with the role of ROS in the resistance response and the control of their levels to minimise toxic effects.