Meiotic instability of common wheat strains derived from Triticum timopheevi Zhuk. crosses

Summary Meiotic instability was studied in four strains of common wheat derived from crosses involving a Triticum timopheevi derivative, C.I. 13093, and the common wheat varieties Cheyenne and Minturki. A wide range in the percentage of normal cells at six stages of meiosis, MI, AI, dyad, MII, AII, and quartet, was found. The date of sporocyte sampling influenced meiotic irregularities at six stages of meiosis. Florets from the same spikes differed in the percentage of normal cells at all the meiotic stages except MII. Plants within strains differed significantly at the MI and quartet stages only. Except at AI stage, spikes collected on the same day did not differ significantly. Apparently, environment during premeiotic phase determined the extent and the pattern of meiotic instability in spikes of these strains.Chromosome counts were taken on 213 of the highly aberrant PMC at MI. Among these, 44 different chromosome numbers ranging from 4 to 76 per cell were recorded. Cells with deviating numbers from 22 to 28 were the most frequent (30.52 percent). Chromosome numbers 12, 17, 21, 22, 24, 26 and 28 occurred in 43.9 percent of the aberrant PMC, and the number of cells with these chromosome numbers ranged from 9 to 15.The metaphase 1 chromosome pairing in deviating PMC was not related to the number of chromosomes in these PMC. Apparently, deviating PMC had a random assortment of chromosomes. In aberrant PMC and others with apparently normal chromosome numbers (but irregular meiosis) homeologous chromosomes may pair. Functional gametes from such cells may perpetuate meiotic instability that had persisted in these three advanced generation common wheat strains of hybrid origin.     

Chromosomal locations in common wheat of three new leaf rust resistance genes from Triticum monococcum

Monosomic analysis was conducted to determine chromosomal locations of three new leaf rust resistance genes recently transferred to common wheat (Triticum aestivum) from T. monococcum. The resistance gene in wheat germplasm line KS92WGRC23 was transferred from T. monococcum ssp. monococcum. The resistance genes found in KS93U3 and KS96WGRC34 were transferred from T. monococcum ssp. aegilopoides. Allelism tests showed that the three resistance genes were unlinked. The three lines were crossed with each of the seven A-genome Wichita monosomic lines. The leaf rust resistance genes in KS92WGRC23, KS93U3, and KS96WGRC34 were located on chromosomes 6A, 1A, and 5A, respectively, by monosomic analysis. These results demonstrate that the three new genes derived from T. monococcum are each different. They also differ from previously reported Lr genes. This information on chromosome location and the development of mapping populations will facilitate molecular tagging of the new genes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Identification of microsatellite markers for a leaf rust resistance gene introgressed into common wheat from Triticum timopheevii

The tetraploid wheat Triticum timopheevii Zhuk (A^tA^tGG) is known as a source of genes determining resistance to many diseases. An introgressive line 842, with durable resistance to leaf rust was established by crossing T. aestivum cv. ‘Saratovskaya29’ with T. timopheevii ssp. viticulosum and used for mapping leaf rust resistance genes. Molecular analysis of the line 842 with polymorphic microsatellite markers detected introgressions of T. timopheevii into the homoeologous group 2 chromosomes of common wheat. Transloca‐tion breakpoints of introgressed fragments were localized between the markers Xgwm95 and Xgwm817 on chromosome 2A, as well as Xgwm1128 and Xgwm1067 on chromosome 2B. Linkage analysis demonstrated the association of disease resistance at the seedling stage with chromosome 2A. The gene was found to be linked with marker Xgwm817 at a genetic distance of 1.5 cM. The alien leaf rust resistance gene was temporarily designated as lrTt1.     

Leaf rust and stripe rust resistance genes transferred to common wheat from Triticum dicoccoides

Linked leaf rust and stripe rust resistance genes introduced from Triticum dicoccoides protected common wheat seedlings against a range of pathotypes of the respective pathogens. The genes were chromosomally mapped using monosomic and telosomic analyses, C-banding and RFLPs. The data indicated that an introgressed region is located on wheat chromosome arm 6BS. The introgressed region did not pair with the ‘Chinese Spring’ 6BS arm during meiosis possibly as a result of reduced homology, but appeared to pair with 6BS of W84-17 (57% of pollen mother cells) and ‘Avocet S’. The introgressed region had a very strong preferential pollen transmission (0.96–0.98) whereas its transmission through egg cells (0.41–0.66) varied with the genetic background of the heterozygote. Homozygous resistant plants had a normal phenotype, were fertile and produced plump seeds. Symbols Lr53 and Yr35 are proposed to designate the respective genes.     

Use of a synthetic hexaploid Triticum miguschovae for transfer of leaf rust resistance to common wheat

Summary Triticum miguschovae, a genome addition synthetic, was used as a source for transfer of leaf rust (Puccinia recondita tritici) resistance to common wheat. This synthetic, developed from two wild species Triticum militinae and Aegilops squarrosa, proves a valuable donor of the genes for leaf rust resistance. Leaf rust resistance was transferred from T. miguschovae by both dominant and recessive genes. Stable lines phenotypically similar to their recurrent parents Kavkaz and Bezostaya 1 but differing from them in a high level of leaf rust resistance were obtained. The genes for resistance in 3 selected lines differed from each other and from the known effective genes Lr9, Lr19, and Lr24. The resistance of one of them (line 1229) is controlled by two complementary interacting genes located on chromosome 7B and 1D was revealed by monosomic analysis.     

Chromosomal location in triticale of leaf rust resistance genes introduced from Triticum monococcum

This study was undertaken to understand the inheritance of leaf rust resistance in line TM16 of Triticum monococcum ssp. monococcum var. macedonicum Papag. which is the source of resistance transferred into hexaploid triticale lines (Tcl/Tm). Thirty-two secondary tetraploid genotypes were analysed cytologicaly to identify substitutions of A^m-genome chromosomes by their homoeologous A-genome chromosomes from a leaf rust susceptible hexaploid triticale accession. Plants with one (or more) substituted chromosomes were inoculated with leaf rust at two growth stages. The disease phenotypes of these lines indicated that a major resistance gene was located on the short arm of T. monococcum chromosome 2A^m. An additional gene on chromosome 6A^m had complementary effects in enhancing the effects of the gene on chromosome 2A^m.     

Wheat leaf rust resistance gene Lr59 derived from Aegilops peregrina

An Aegilops peregrina (Hackel in J. Fraser) Maire & Weiller accession that showed resistance to mixed leaf rust ( Puccinia triticina Eriks.) inoculum was crossed with, and backcrossed to, hexaploid wheat ( Triticum aestivum L.). During backcrossing a chromosome segment containing a leaf rust resistance gene (here designated Lr59 ) was spontaneously translocated to wheat chromosome 1A. Meiotic, monosomic and microsatellite analyses suggested that the translocated segment replaced most of, or the complete, 1AL arm, and probably resulted from centromeric breaks and fusion. The translocation, of which hexaploid wheat line 0306 is the appropriate source material, provided seedling leaf rust resistance against a wide range of South African and Canadian pathotypes.     

Evaluation of slow rusting resistance components to leaf rust in CIMMYT durum wheats

Information about slow rusting resistance to leaf rust (Puccinia triticina) in durum wheat (Triticum turgidum var. durum) is limited. Three slow rusting components, latent period, receptivity, and uredinium size, were determined at the adult plant stage for seven durums with slow rusting resistance to leaf rust and two susceptible durums in three greenhouse experiments. Additionally, area under the disease progress curve (AUDPC) and final disease severity (FDS) were determined in three field trials under artificial epidemics with the same P. triticina race BBG/BN. Compared to the most susceptible check, the AUDPC and FDS of slow rusting resistant durums were significantly lower and ranged from 13–47 to 22–59%, respectively. The latent period was significantly longer (8.5–10.3 days) and uredinium size significantly smaller (8.1–14.8 × 10⁻² mm²) on slow rusting durums than on the susceptible checks (8.0 days and 17.3–23.8 × 10⁻² mm², respectively). Uredinium size was the most stable slow rusting component across experiments. Correlations between uredinium size versus AUDPC and uredinium size versus FDS for each environment were high (r = 0.86–0.88). Correlations between latent period and field parameters were significant (r = −0.60 to −0.80). Correlations between receptivity and the field parameters were not significant. A multiple regression analysis showed that the variation in AUDPC and FDS was significantly explained only by uredinium size (P < 0.0001). The best slow rusting resistant lines can be used for developing high-yielding durums with more durable resistance to leaf rust.     

Adult plant leaf rust resistance from 111 wheat (Triticum aestivum L.) cultivars

Adult plant resistance against Indian leaf rust race 77 and five of its highly virulent variants have been identified from 111 bread wheat cultivars originating from 12 countries. The adult plant resistance of only 16 of these cultivars is due to hypersensitive seedling or adult plant resistance genes. All others expressed nonhypersensitive type of resistance characteristic of the genes Lr34 and Lr46.Forty five of the 111 cultivars showed tip necrosis on flag leaves, a trait linked to the gene Lr34. Therefore, the nonhypersensilive type of resistance of these 45 cultivars is attributed to Lr34. The nonhypersensitive resistance of the remaining cultivars is likely to be due to the gene(s) different than Lr34. The reaction pattern of these 111 cultivars to six races suggests the presence of at least six to seven new hypersensitive adult plant resistance genes and at least three new hypersensitive seedling resistance genes. The known genes Lr10, Lr23 and Lr26 were detected frequently but these genes did not contribute towards the adult plant resistance of any of the 111 cultivars. Based on the presence of new genes for hypersensitive and nonhypersensitive type of resistance, the 111 cultivars have been classified into 31 diverse resistance groups. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chromosome locations of leaf rust resistance genes in selected tetraploid wheats through substitution lines

Langdon durum D-genome disomic substitution lines were used to study the chromosome locations of adult-plant leaf rust resistance genes identified from tetraploid wheat accessions. The accessions are 104 (Triticum turgidum subsp. dicoccum var. arras) and 127 (T. turgidum subsp. durum var. aestivum). The complete sets of the substitution lines were crossed as female parents with the accessions and F₁ double monosomic individuals selected at metaphase I. Segregating F₂ individuals were inoculated during the flag leaf stage with pathotype UVPrt2 of Puccinia triticina. The substitution analysis involving accession 104 showed that the gene for leaf rust resistance is located on chromosome 6B. The analysis with accession 127 indicated that chromosome 4A carries a gene for leaf rust resistance. The two novel genes are temporarily designated as Lrac104 and Lrac127, respectively from accessions 104 and 127.     

Inheritance of temperature-sensitive leaf rust resistance and adult plant stripe rust resistance in common wheat cultivar PBW343

Genetic analysis of common wheat cultivar PBW343 confirmed temperature-sensitive leaf rust resistance and adult plant stripe rust resistance. At low temperatures, PBW343 was resistant to P. triticina (Ptr) pathotype (pt.) 121R63-1, and at high temperature it was resistant to Ptr pt. 121R127. The low temperature resistance to pt. 121R63-1 was attributed to interaction between dominant and recessive genes. The dominant gene involved in low-temperature resistance to pt. 121R63-1 also conferred resistance to pt. 45R35. The high-temperature resistance to Ptr pt. 121R127 was governed by a different single partially dominant gene. Agra Local (a commonly used susceptible check) and IWP94 (a leaf rust differential used in India) are also resistant to pt. 121R127 at high temperatures. An allelism test indicated that PBW343 and IWP94 possessed a common gene for high temperature resistance to this pathotype. The adult plant stripe rust resistance against P. striiformis (Pst) was possibly conferred by one gene in addition to Yr27.     

The inheritance of leaf rust resistance in four varieties of common wheat

Summary The genetic nature of resistance to four Australian strains of Puccinia recondita was studied in the cultivar Timgalen and in three entries of the 1963 ISWRN, viz, 25 (W3300) 318 (W3301) and 409 (W3303). Seedling resistance of Timgalen was controlled by the dominant gene Lr3 and by a recessive gene, tentatively designated lrT. A third gene, identical with one in C.I.12632, was operating in adult plants only.In each of the three varieties W3300, W3301, W3303 the same dominant and the same recessive gene were operating in seedlings. The former may be identical with a major gene previously reported in Klein Titan, Frontana and Rio Negro. The latter gene was different from lrT of Timgalen. In addition, W3301 and W3303 each carried a dominant factor being only operative in the mature plant stage.     

Environmental stability of partial resistance in spring wheat to wheat leaf rust

Summary Five spring wheat cultivars differing in partial resistance (PR) to wheat leaf rust were tested at Wageningen (the Netherlands) on a sandy and a clay site, El Batan (CIMMYT, Mexico) and Ponta Grossa (Brazil) over two years. The cultivars were Skalavatis 56, Little Club (both very susceptible), Westphal 12A, Akabozu and BH 1146 (all three with high levels of PR). The results showed that PR was expressed at all four locations in both years. The level of expression was influenced by the environment but the cultivar ranking was hardly affected. Selection for PR in the field can therefore be carried out over a wide range of environments.     

Molecular mapping of Aegilops speltoides derived leaf rust resistance gene Lr28 in wheat

In a segregating homozygous F₂ population of bread wheat involving a leaf rust resistance gene Lr28 derived from Aegilops speltoides, six randomly amplified polymorphic DNA (RAPD) markers, three each in coupling and repulsion phase were identified as linked to Lr28, mapped to a region spanning 32 cM including the locus. The F₂ and F₃ populations were studied in the phytotron challenged with the most virulent pathotype 77-5 of leaf rust. A coupling phase linked RAPD marker S464₇₂₁ and a repulsion phase linked RAPD marker S326₅₅₀ flanked the gene Lr28 by a distance of 2.4± 0.016 cM on either side. The flanking markers genetically worked as co-dominant markers when analyzed together after separate amplification in the F₂ population by distinguishing the homozygotes from the heterozygotes and increased the efficiency of marker assisted selection by reducing the false positives and negatives. One of the three RAPD markers, S421₆₄₀ was converted to locus specific SCAR marker SCS421₆₄₀ which was further truncated by designing primers internal from both ends of the original RAPD amplicon to eliminate a non-specific amplification of nearly same size. The truncated polymorphic sequence characterized amplified region marker (TPSCAR) SCS421₅₇₀ was 70 bp smaller, but resulted in a single band polymorphism specific to Lr28 resistance. The TPSCAR marker was validated for its specificity to the gene Lr28 in nine different genetic backgrounds and on 43 of the 50 Lr genes of both native and alien origin, suggesting the utility of the SCAR markers in pyramiding leaf rust resistance genes in wheat.     

Growth of wheat leaf rust colonies in susceptible and partially resistant spring wheats

Summary The average size of wheat leaf rust colonies, measured using epifluorescence microscopy was significantly larger in the highly susceptible genotype Morocco than in the susceptible genotype Kaspar and the partially resistant genotypes Westphal 12A, Akabozu and BH 1146. This was already so three days after inoculation. Colony growth in partially resistant genotypes was continuously retarded compared to colonies in the highly susceptible genotype Morocco. No evidence was found for an initial inhibition of the growth of colonies in partially resistant genotypes. In partially resistant genotypes formation of uredial beds and sporulating areas started at a smaller colony size than in susceptible genotypes. Wheat leaf rust colonies in primary leaves of all genotypes studied were much larger than colonies in flag leaves measured at the same number of days after inoculation. Growth and sporulation of not intertwined colonies was not influenced by either a high or a low number of neighbouring colonies.     

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

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

Genetic analysis of adult plant leaf rust resistance in three bread wheat (Triticum aestivum L.) cultivars

Genetic basis of adult plant leaf rust resistance in three released Indian wheat cultivars viz. DWR195, RAJ3765 and HP1731 was investigated through detailed inheritance study under controlled polythene house condition at Flowerdale, India. The F₂, F_3, F₄ and F₅ generations were analyzed with the most frequent and virulent Indian leaf rust pathotype 121R63-1. Two complementary recessive genes imparted resistance in DWR195, two complementary dominant genes governed the resistance of RAJ3765 whereas two independent dominant genes were involved in the resistance of HP1731. The genes responsible for adult plant resistance in the three cultivars were not allelic. The two complementary genes of DWR195 and two independent dominant genes of HP1731 have been isolated as single gene lines. Utilization of resistance from HP1731, which carries two independent dominant genes, will be easy as compared to DWR195 and RAJ3765.     

Heterosis of grain weight in wheat hybrids with Triticum timopheevi cytoplasm

Summary Two kinds of wheat hybrids with the same nuclear genotype but different cytoplasms (one with T.timopheevi cytoplasm, i.e., A/R, and the other with T.aestivum cytoplasm, i.e., B/R) were produced by two 3 × 5 incomplete diallel crosses of 3 A-lines, 3 corresponding B-lines and 5 R-lines, respectively. Experimental results did not show significant differences between the hybrids of A/R and B/R in grain filling characteristics and grain weight. The beterosis of grain weight seems mainly determined by the nuclear genotype. Although the seeds set on most A-lines were shrivelled, such a phenomenon was not found in grains set on F₁. The duration of the lag period (D₁) and the average grain filling rates during the linear period and the mature period (i.e., FR₂ and FR₃) were significantly and positively related to 1000-grain weight. It was in these factors that most hybrids displayed clear mid-parent (MP) heterosis. The amount of grain weight heterosis was not significantly related to MP value. This indicates that the grain weight heterosis of wheat hybrids will not decrease with an increase of the MP value.     

Gametocidal effects and resistance to wheat leaf rust and stem rust in derivatives of a Triticum turgidum ssp. durum/Aegilops speltoides hybrid

Summary Genes for leaf rust and stem rust resistance and segregation distortion (Gc), that seemed to derive from an Aegilops spetroides ssp. ligustica accession, were transferred to common wheat. While the advanced backcrosses had normal meioses and 42 chromosomes, high levels of male and female sterility, abnormal endosperm development and chromosome aberrations were evident. These effects were more pronounced in Gc-heterozygotes than in homozygotes. Gametes without Gc genes did not survive, and the Gc-associated defects were always inherited with the resistance. Since the resistance genes were effective against local pathotypes of the leaf rust and stem rust pathogens, an attempt was made to disrupt the Gc-system through irradiation, treatment with the mutagen N-nitroso-N-methyl-urea or growing the material at elevated temperatures. A very low frequency of the treated material showed slightly better fertility and seed development. However, these effects did not persist in subsequent generations and were apparently not strong enough to allow the recovery of segregates which had lost the Gc gene(s).     

Detection and inheritance of leaf rust resistance in common wheat lines Agra Local and IWP94

Genetic studies were undertaken to determine the number and identities of leaf rust resistance genes in common wheat lines Agra Local and IWP94. The infection type arrays of the two lines with eight pathotypes (pt.) of P. triticina were different from those of lines possessing known leaf rust resistance (Lr) genes. Agra Local possessed two recessive resistance genes, one conditioning resistance to pathotype 4R9-7, and the other, a temperature-sensitive factor, gave resistance to pt. 121R127 at high temperature (27°C). IWP94 was previously demonstrated to carry Lr23. From the present study IWP94 was determined to have at least four leaf rust resistance genes. The first of these was the same recessive gene conferring resistance to pathotype 4R9-7 which was found in Agra Local. A second partially dominant gene conferred resistance to pathotype 121R127 at high temperature and two additional recessive genes governed resistance to pathotype 93R15. When present together, these two recessive genes complemented each other and provided resistance to pathotype 69R13 as well. One of the two recessive genes conferring resistance to pathotypes 93R15 and 69R13 was Lr23.