Reduction of <Emphasis Type="Italic">Aegilops sharonensis</Emphasis> chromatin associated with resistance genes <Emphasis Type="Italic">Lr56</Emphasis> and <Emphasis Type="Italic">Yr38</Emphasis> in wheat

The Lr56/Yr38 translocation consists primarily of alien-derived chromatin with only the 6AL telomeric region being of wheat origin. To improve its utility in wheat breeding, an attempt was made to exchange excess Ae. sharonensis chromatin for wheat chromatin through homoeologous crossover in the absence of Ph1. Translocation heterozygotes that lacked Ph1 were test-crossed with Chinese Spring nullisomic 6A tetrasomic 6B and nullisomic 6A-tetrasomic 6D plants and the resistant (hemizygous 6A) progeny were analyzed with four microsatellite markers. Genetic mapping suggested general homoeology between wheat chromosome 6A and the translocation chromosomes, and showed that Lr56 was located near the long arm telomere. Thirty of the 53 recombinants had breakpoints between Lr56 and the most distal marker Xgwm427. These were characterized with additional markers. The data suggested that recombinants #39, 157 and 175 were wheat chromosomes 6A with small intercalary inserts of foreign chromatin containing Lr56 and Yr38, located distally on the long arms. These three recombinants are being incorporated into adapted germplasm. Attempts to identify the single shortest translocation and to develop appropriate markers are being continued.     

Evaluation and reduction of Lr19-149, a recombined form of the Lr19 translocation of wheat

The Lr19 translocation was introgressed from Thinopyrum ponticum in 1966. It has not been used in wheat breeding in many countries despite it being an excellent source of leaf rust resistance as it carries an undesirable gene(s) coding for yellow endosperm pigmentation. A shortened form, Lr19-149, was since produced and lacks the yellow pigment genes. A yield trial with near isogenic lines of both the original and shortened translocations suggested that Lr19 may cause a small reduction in kernel size and anincrease in loaf volume, effects which are not associated with Lr19-149. In Lr19-149 heterozygotes the translocation generally showed reduced pollen transmission whereas its transmission through egg cells was mostly normal. An attempt to shorten Lr19-149 through allosyndetic recombination in the absence of Ph1b produced four recombinants which were characterized by means of RFLP and AFLP polymorphisms and physically mapped with a set of 27 deletion lines. In three recombinants (252, 299 and 462) Thinopyrum chromatin proximally to Lr19 was exchanged for wheat chromatin. In one recombinant (478) chromatin distally from Lr19 was replaced. Based on physical map distance estimates it appears that the Lr19 translocation in the shortest recombinant (299) may have been reduced to about one third or less of its original size. It may now be possible to obtain a further, albeit relatively small, decrease in the size of the translocation through homologous crossover between recombinants 299 and 478. Similar to Lr19-149, the new recombinants show self elimination in heterozygotes and they have apparently retained the Sd2 locus. This revised version was published online in August 2006 with corrections to the Cover Date.     

Leaf Rust and Stripe Rust Resistance Genes Derived from Aegilops Sharonensis

Summary Linked leaf and stripe rust resistance genes introgressed into hexaploid wheat from Aegilops sharonensis provided protection in the seedling stage to a wide range of pathotypes of the two diseases. Monosomic and telosomic analyses showed that the resistance genes occur on wheat chromosome 6A. This result could be confirmed making use of mapped chromosome 6A microsatellite markers. The introgressed chromatin appeared to involve the proximal part of 6AL and the complete 6AS arm and it was thus not possible to deduce the chromosome arm harbouring the resistance genes. The resistance showed non-Mendelian transmission. The genetic background of a heterozygote interacted with the introgressed region to result in either preferential or impaired female transmission. Male transmission appeared to be affected in a different way from female transmission and was exclusive in the genetic background studied. Symbols Lr56 and Yr38 are proposed to designate the respective genes of which line 0352-4 is the appropriate source material.     

Molecular and agronomic characterization of wheat-Agropyron intermedium recombinant chromosomes

Thirty‐six wheat‐Agropyron intermedium (host) Beauv. [Syn. Trichopyrum intermedium (host) A. Love, Elytrigia intermedia (host) Nevski, Thinopyrum intermedium (host) Barkworth and Dewey] 7A/7Ai‐1 recombinant chromosomes were characterized using DNA markers. Analysis of recombinant chromosomes using 15 restriction fragment length polymorphism probes identified the homoeologous crossover products that had varying length of A. intermedium chromatin introgressed onto chromosome 7A of common wheat. The linear order of the probe loci was established along the lengths of the chromosomes. The short arm recombinants that had A. intermedium chromatin distal to the locus Xpsr108 and proximal to the locus Xpsr119 were resistant to wheat stem rust, indicating that the rust resistance gene (Sr44) was located on the distal part of chromosome arm 4Ai‐1s. The barley yellow dwarf virus (BYDV) resistance gene reported to be present on the long arm of chromosome 7Ai‐1 was found to be ineffective against the BYDV serotype used in the present study.     

Induction of a mutation in the male fertility gene of the preferentially transmitted Aegilops sharonensis chromosome 4S1 and its application for hybrid wheat production

Summary Wheat plants nullisomic for chromosome 4B are male sterile due to the absence of the male fertility gene Ms1. However, plants in which chromosome 4B has been substituted by the preferentially transmitted chromosome 4S¹ of Ae. sharonensis are male fertile due to the compensating effect of Ms4 on the alien chromosome. This substitution line has been mutated and three recessive mutation of Ms4 have been selected. Plants homozygous for these mutations are male sterile. The implication of these mutations for hybrid wheat production is discussed.     

Cytogenetical studies in wheat XVI. Chromosome location of a new gene for resistance to leaf rust in a Japanese wheat-rye translocation line

Summary A leaf rust resistant wheat-rye translocation stock, ST-1, introduced from Japan, comprised distinct morphological types. One type possessed a T1BL·1RS chromosome with genes Lr26, Yr9 and Sr31. A second type carried a new gene, Lr45, located in a large segment of rye chromosome translocated to wheat chromosome 2A. Its structure was identified as T2AS-2RS·2RL. Despite the homoeology of the 2A and 2R chromosomes and the high level of compensation provided by the translocation, Lr45 was not normally inherited and is probably associated with agronomic deficiencies that will prevent its exploitation in agriculture.Contribution No. 94-509-J from the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, USA.     

A microsatellite marker linked to leaf rust resistance transferred from Aegilops triuncialis into hexaploid wheat

Aegilops triuncialis (UUCC) is an excellent source of resistance to various wheat diseases, including leaf rust. Leaf rust‐resistant derivatives from a cross of a highly susceptible Triticum aestivum cv.‘WL711’ as the recurrent parent and Ae. triuncialis Ace.3549 as the donor and with and without a pair of acrocentric chromosomes were used for molecular tagging. The use of a set of sequence tagged microsatellite (STMS) markers already mapped to different wheat chromosomes unequivocally indicated that STMS marker gwm368 of chromosome 4BS was tightly linked to the Ae. triuncialis leaf rust resistance gene transferred to wheat. The presence of the Ae. Triuncialis‐specific STMS gwm368 homoeoallele along with the non‐polymorphic 4BS allele in the rust‐resistant derivatives with and without the acrocentric chromosome indicates that the resistance has been transferred from the acrocentric chromosome to either the A or the D genome of wheat. This alien leaf rust resistance gene has been temporarily named as LrTr.     

Changes in the meiotic pairing behaviour of a winter wheat-winter barley hybrid maintained for a long term in tissue culture, and tracing the barley chromatin in the progeny using GISH and SSR markers

The aim of the present study was to produce backcross progenies in a new winter wheat (‘Asakaze komugi’) × winter barley (‘Manas’) hybrid produced in Martonvasar. As no backcross seeds were obtained from the initial hybrids, young inflorescences of the hybrids were used for in vitro multiplication in three consecutive cycles until a backcross progeny was developed. The chromosome constitution of the regenerated hybrids was analysed using genomic in situ hybridization (GISH) after each in vitro multiplication cycle. The seven barley chromosomes were present even after the third in vitro multiplication cycle but abnormalities were observed. Sixteen BC; plants containing, according to GfSH analysis, one to three complete barley chromosomes, two deletion barley chromosomes and a dicentric wheat‐barley translocation were grown to maturity from the single backcross progeny. The barley chromatin was identified using 20 chromosome‐specific barley SSR markers. All seven barley chromosomes were represented in the BC: plants. A deletion breakpoint at FL ±0,3 on the 5HL chromosome arm facilitated the physical localization of microsatellite markers.     

Leaf rust and stripe rust resistance genes Lr54 and Yr37 transferred to wheat from Aegilops kotschyi

The tendency of unpaired meiotic chromosomes to undergo centric misdivision was exploited to translocate leaf rust and stripe rust resistance genes from an Aegilops kotschyi addition chromosome to a group 2 chromosome of wheat. Monosomic and telosomic analyses showed that the translocation occurred to wheat chromosome arm 2DL. The introgressed region did not pair with the corresponding wheat 2DL telosome during meiosis suggesting that a whole arm may have been transferred. Female transmission of the resistance was about 55% whereas male transmission was strongly preferential (96%). The symbols Lr54 and Yr37 are proposed to designate the new resistance genes.     

Basic studies on hybrid wheat breeding using the 1BL-1RS translocation chromosome / Aegilops kotschyi cytoplasm system 1. Development of male sterile and maintainer lines with discovery of a new fertility-restorer

The Aegilops kotschyi cytoplasm and a 1BL-1RS translocation chromosome that consists of the long arm of wheat chromosome 1B and the short arm of rye chromosome 1R were transferred to six spring common wheat cultivars by repeated backcrossing. Resistance to leaf rust race 21B conditioned by the Lr26 gene and a secalin subunit encoded by the Sec-1 gene, both on the 1RS arm, were used as the selection markers of the translocation chromosome. Five of the six cultivars used were converted to complete male steriles, whereas the remaining one, cv. Kitamiharu 48, retained normal fertility, after transfer of both the 1BL-1RS chromosome and Ae. Kotschyi cytoplasm. Conventional gene analysis suggested that Kitamiharu 48 carries an incompletely dominant fertility-restoring gene. The F₁ hybrids between the male steriles and ordinary common wheat cultivars recovered fertility only at a low level, indicating that a single dose of the Rfv1 gene on the 1BS arm of wheat is insufficient for full fertility restoration under spring-sowing condition. Our results are in clear contrast to complete fertility restoration under fall-sowing condition reported by Nonaka et al. (1993). Combination of the 1BL-1RS chromosome / Ae. Kotschyi cytoplasm system with a new fertility-restoring gene discovered in Kitamiharu 48 may provide a breakthrough for spring-type hybrid wheat. This revised version was published online in August 2006 with corrections to the Cover Date.     

A Robertsonian translocation from Thinopyrum bessarabicum into bread wheat confers high iron and zinc contents

Development of wheat–alien translocation lines has facilitated practical utilization of alien species in wheat improvement. The production of a compensating Triticum aestivum–Thinopyrum bessarabicum whole‐arm Robertsonian translocation (RobT) involving chromosomes 6D of wheat and 6E^b of Th. bessarabicum (2n = 2x = 14, E^bE^b) through the mechanism of centric breakage–fusion is reported here. An F₂ population was derived from plants double‐monosomic for chromosome 6D and 6E^b from crosses between a DS6E^b(6D) substitution line and bread wheat cultivar ‘Roushan’ (2n = 6x = 42, AABBDD) as female parent. Eighty F₂ genotypes (L1–L80) were screened for chromosome composition. Three PCR‐based Landmark Unique Gene (PLUG) markers specific to chromosomes 6D and 6E^b were used for screening the F₂ plants. One plant with a T6E^bS.6DL centric fusion (RobT) was identified. A homozygous translocation line with full fertility was recovered among F₃ families and verified with genomic in situ hybridization (GISH). Grain micronutrient analysis showed that the DS6E^b(6D) substitution line and T6E^bS.6DL stock have higher Fe and Zn contents than the recipient wheat cultivar ‘Roushan’.     

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.     

Origin and variation of gliadin proteins associated with pasta quality in durum wheat

Summary Studies were made of the presence and frequency of occurrence of gliadin bands 42 and 45 in three samples of Aegilops sharonensis Eig and 59 samples of wild tetraploid wheat (Triticum dicoccoides Korn.) from natural distributions of these species in Israel.Two samples of Ae. sharonensis possessed a band in position 45 and one possessed no bands corresponding to either band 45 or band 42. In T. dicoccoides, band 45 was either present or not and 42 was always absent. In its grassy and intermediate growth habit forms, (believed to be more primitieve than the cercal forms) band 45 appeared to be more frequent than in the cereal form.The presence of band 45 in the Ae. sharonensis, and its relatively high frequency in T. dicoccoides, populations from Mt. Hermon (likely to be relatively free from introgression from cultivated tetraploid wheat) indicate the likelihood of a primary origin of the allele coding for band 45. The absence of band 42 from all Ae. sharonensis and T. dicoccoides populations in this study, indicates a more recent evolutionary origin of the allele coding for this band, possibly arising as a mutation during the domestication of tetraploid wheat.The results have implications for breeding programmes in tetraploid wheat.     

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.     

Ph I-induced transfer of leaf and stripe rust-resistance genes from Aegilops triuncialis and Ae. geniculata to bread wheat

Leaf and stripe rusts are severe foliar diseases of bread wheat. Recently, chromosomes 5M^g from the related species Aegilops geniculata that confers resistance to both leaf and stripe rust and 5U^t from Ae. triuncialis conferring resistance to leaf rust have been transferred to bread wheat in the form of disomic DS5M^g(5D) and DS5U^t(5A) chromosome substitution lines. The objective of this study was to shorten the alien segments in these lines using Ph ^I-mediated, induced homoeologous recombination. Putativerecombinants were evaluated for their rust resistance, and by genomic in situ hybridization and microsatellite analyses. One agronomically useful wheat-Ae. geniculata recombinant resistant to leaf and stripe rust was identified that had only a small terminal segment of the 5M^gL arm transferred to the long arm of an unidentified wheat chromosome. This germplasm can be used directly in breeding programs. Only one leaf rust-resistant wheat-Ae. triuncialis recombinant, which consists of most of the complete 5U^t chromosome with a small terminal segment derived from 5AS, was identified. This germplasm will need further chromosome engineering before it can be used in wheat improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic analysis of the Aegilops longissima 3S chromosome carrying the Pm13 resistance gene

The distal region of the short arm of chromosome 3S from Aegilopslongissima, which carries the powdery mildew resistance gene Pm13, was introgressed into common wheat. Due to suppression of recombination between this region and corresponding wheat homoeologous segments, a possible strategy to construct a genetic map around the Pm13 gene was based on crosses between a wheat addition line carrying the Ae.longissima 3S chromosome and the corresponding 3S addition lines of Ae.searsii and Ae. variabilis. The efficiency of this strategy was evaluated by scoring recombination frequencies inprogenies derived from these crosses. Recombination between 3S chromosomes fromAe. searsii and Ae. longissimawas very low, whereas 26.5% recombinant alien chromosomes were obtained from the cross involving the Ae. variabilisand Ae. longissima 3S addition lines. These data were used to construct a3S chromosome map that resulted largely colinear to the consensus map of the homoeologous group 3 of wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Quantitative trait loci mapping of adult‐plant resistance to leaf rust in a Fundulea 900 × ‘Thatcher’ wheat cross

Wheat leaf rust (LR), caused by the obligate biotrophic fungus Puccinia triticina (Pt), is a destructive foliar disease of common wheat (Triticum aestivum L.) worldwide. The most effective, economic means to control the disease is resistant cultivars. The Romanian wheat line Fundulea 900 showed high resistance to LR in the field. To identify the basis of resistance to LR in Fundulea 900, a population of 188 F_2:3 lines from the cross Fundulea 900/‘Thatcher’ was phenotyped for LR severity during the 2010–2011, 2011–2012 and 2012–2013 cropping seasons in the field at Baoding, Hebei Province. Bulked segregant analysis and simple sequence repeat markers were used to identify the quantitative trait loci (QTLs) for LR adult‐plant resistance in the population. Three QTLs were detected and designated as QLr.hebau‐1BL, QLr.hebau‐2DS and QLr.hebau‐7DS. Based on the chromosome positions and molecular marker tests, QLr.hebau‐1BL is Lr46, and QLr.hebau‐7DS is Lr34. QLr.hebau‐2DS was derived from ‘Thatcher’ and was close to Lr22. This result suggests that Lr22b may confer residual resistance on field nurseries when challenged with isolates virulent on Lr22b, or another gene linked to Lr22b confers this resistance from ‘Thatcher’. This study confirms the value of Lr34 and Lr46 in breeding for LR resistance in China; the contribution of the QTL to chromosome 2D needs further validation.     

Characterization of wheat-alien translocations conferring resistance to diseases and pests: current status

Summary Wild relatives of common wheat, Triticum aestivum, and related species are an important source of disease and pest resistance and several useful traits have been transferred from these species to wheat. C-banding and in situ hybridization analyses are powerful cytological techniques allowing the detection of alien chromatin in wheat. C-banding permits identification of the wheat and alien chromosomes involved in wheat-alien translocations, whereas genomic in situ hybridization analysis allows determination of their size and breakpoint positions. The present review summarizes the available data on wheat-alien transfers conferring resistance to diseases and pests. Ten of the 57 spontaneous and induced wheat-alien translocations were identified as whole arm translocations with the breakpoints within the centromeric regions. The majority of transfers (45) were identified as terminal translocations with distal alien segments translocated to wheat chromosome arms. Only two intercalary wheat-alien transloctions were identified, one induced by radiation treatment with a small segment of rye chromosome 6RL (H25) inserted into the long arm of wheat chromosome 4A, and the other probably induced by homoeologous recombination with a segment derived from the long arm of a group 7 Agropyron elongatum chromosome with Lr19 inserted into the long arm of 7D. The presented information should be useful for further directed chromosome engineering aimed at producing superior germplasm.Contribution No. 96-55-J from the Kansas Experimental Station, Kansas State University, Manhattan, KS 66506-5502, USA.     

An assessment of the potential of 4DS.4DL-4s l L translocation lines as a means of eliminating tall off types in semi-dwarf wheat varieties

Summary Wheat varieties tend to be chromosomally unstable producing on average 2–3% of plants with abnormal chromosome numbers. A number of semi dwarf wheat varieties, carrying the gibberellic acid insensitive dwarfing genes Rht1 or Rht2, have been seen to produce distinct tall off types due to reduction in dosage of the chromosome carrying the dwarfing gene. The UK variety Brigand, carrying Rht2 on chromosome 4D, produced very distinct tall off types when this chromosome was reduced in dosage. The frequency of tall off types was sufficiently high to cause the variety to fail United Kingdom statutory uniformity tests. An attempt to prevent the loss of chromosome 4D was made by constructing translocation chromosomes involving the short arm of chromosome 4D, which carries Rht2, and the long arm of chromosome 4S ^l from Aegilops sharonensis, which carries a gene(s) conferring preferential transmission. The work in this paper describes the field evaluation of two lines carrying 4DS.4DL-4S ^l L translocations, and demonstrates their success in preventing spontaneously occurring monosomy of chromosome 4D in semi-dwarf wheats.     

Genetic polymorphisms at Gli-Dt gliadin loci in Aegilops tauschii as revealed by acid polyacrylamide gel and capillary electrophoresis

Gliadin variation at Gli‐D^t1 and Gli‐D^t2 loci in 198 Aegilops tauschii accessions was studied by acid polyacrylamide gel electrophoresis (A‐PAGE) and capillary electrophoresis (CE). High genetic polymorphisms were found at both gliadin coding loci, revealing a total of 184 and 169 gliadin variants at the Gli‐D^t1 and Gli‐D^t2 loci, respectively. In particular, 12 gliadin blocks encoded by different alleles were apparently expressed and readily identified in six synthetic hexaploids produced by hybridization between Triticum durum and Ae. tauschii accessions. Compared with Ae. tauschii ssp. eusquarrosa, the gliadin profile of the D genome in Ae. tauschii ssp. strangulata more resembles that of T. aestivum, supporting the view that the subspecies strangulata is the most likely progenitor of bread wheat. Capillary electrophoresis analysis showed that the method is capable of separating and characterizing gliadins with speed, in high resolution using small sample amounts, and is well‐suited to detect protein alleles and to identify desirable genotypes in wheat quality improvement.