Cytological and microsatellite mapping of the genes determining liguleless phenotype in durum wheat

Liguleless phenotypes of wheat lack ligule and auricle structures on all leaves of the plant. Two recessive genes principally control the liguleless character in tetraploid wheat. The F₂ progenies of k17769 (liguleless mutant)/Triticum dicoccoides and k17769/T. dicoccum segregated in a 15:1 ratio, whereas the F₂ progenies of k17769/T. durum and k17769/T. turgidum segregated in a 3:1 ratio. A new gene, lg₃, was found on chromosome 2A. Segregation of F₂ progenies between k17769 and chromosome substitution lines for homoeologous group 2 chromosomes suggested that the liguleless genotype had occurred by mutation at the lg₃ locus on chromosome 2A, and then by mutation at the lg₁ locus on chromosome 2B, in the process of domestication of tetraploid wheat. The gene (lg₁) was linked to Tc₂ (11.9 cM), which determines phenol colour reaction of kernels, on the long arm of chromosome 2B. The distance of lg₁ to the centromere was found to be 60.4 cM, and microsatellite mapping established the gene order, centromere – Xgwm382 – Xgwm619 – Tc₂ – lg₁ on the long arm of chromosome 2B.     

Genetic mapping of the genes for non-glaucous phenotypes in tetraploid wheat

Glaucousness is a visual trait related to the colour of the photosynthetic surface and hence it can be easily selected. It is associated both with the deposition and orientation of wax platelets on the cuticle of the photosynthetic surface. It is known that the glaucous leaf character is determined by the W1 gene, and Iw1 and Iw1 ^DIC genes from Triticum dicoccoides, which act as an epistatic inhibitor to a glaucousness. The aim of the present study was to map W1, Iw1 and Iw1 ^DIC from T. dicoccoides in the short arm of chromosome 2B of tetraploid wheat. Segregation of F₂ populations of three hybrids indicated that the marker Xgwm455 is linked with and distal to Iw1 (16.7 cM), two markers are tightly linked with W1, and Iw1 ^DIC is linked with Xgwm614 and Xwmc661 distally located on chromosome 2BS. From three derived maps, it is suggested that Iw1 ^DIC locus is different from W1. We discuss the difference between Vir and Iw1 ^DIC from T. dicoccoides.     

Telosomic mapping of the homoeologous genes for the long glume phenotype in tetraploid wheat

Triticum turgidum ssp. polonicum and T. ispahanicum were characterized by the long glume phenotype. P ₁ gene determines the long glume phenotype of T. polonicum, and locates on chromosome 7A. T. ispahanicum has shorter glume than T. polonicum and the long glumephenotype is determined by P ₂ gene located on chromosome 7B. In the present study, aneuploid stocks of `Langdon' durum wheat were used to map the genes, P ₁ and P ₂. P ₁ located on the long arms of chromosome 7A and its map distances from the centromere was 14.5 cM. On chromosome 7B, four loci located as cc (chocolate black chaff) – Pc (purple culm) – centromere – P ₂ – cn-BI (chlorina). P ₂ located on the long arms of chromosome 7B and its map distances from the centromere was 11.7 cM. It was suggested that a paralogous gene set conditions long glume phenotype in the homoeologous group 7 chromosomes. The P ₁ and P ₂ genes may be useful as genetic markers in tetraploid wheat.     

Variation of characters in near-isogenic lines of wheat with added genes for leaf rust resistance

Summary Using the cultivar Arina as the recurrent parent, six backcrosses were made with two donor lines carrying the leaf rust resistance genes Lr1 and Lr9, respectively. Selection for leaf rust resistance occurred at the seedling stage in the greenhouse; the first plants transferred to the field were BC₆F₄s. Frequency distribution of the 332 Lr1/7 × Arina and the 335 Lr9/7 × Arina lines showed continuous variation for yellow rust resistance and heading date in these leaf rust near-isogenic lines (NILs). Similar results were also obtained for plant height, for resistance to powdery mildew and glume blotch, as well as for baking quality characters in another set of more advanced NILs. The available information on the behaviour of one of the parents of cultivar Arina led to the conclusion that the expressed yellow rust resistance is quantitative and might possibly be durable.     

Agronomic and quality characters of near-isogenic lines of wheat carrying genes for stem rust resistance

Summary Two sets of near-isogenic lines of wheat carrying single genes for stem rust resistance were grown in yield tests to determine whether the resistance genes were deleterious. One set was based on the cultivar Marquis and the second set on a susceptible, day-length insensitive line, LMPG. The results indicated that the effects of resistance genes vary with different genes and different environments. However, there appeared to be a tendency for resistance genes to reduce yield. In most cases the reductions were too small to be of much concern to wheat breeders.     

Genetic control of the long glume phenotype in tetraploid wheat by homoeologous chromosomes

The Ispahan emmer wheat, Triticum ispahanicum Heslot, was discovered in Iran 1957 by the French expedition of Vinnot- Bourgen. T. ispahanicum has a long glume and a more slender spike than T. turgidum var. polonicum. The objectives of this study were (1) to determine the inheritance and chromosomal location of the gene for long glume, P2, from T. ispahanicum using the near- isogenic line P2-LD222, and (2) to compare the effects of the genes for long glume. The gene for long glume, P2, was located approximately 36.5 cM from the cn-B1 locus, which controls the chlorina trait and approximately 40 cM from the centromere on the long arm of 7B. The location of P2 approximately 29.6 cM from the Pc locus produced additional evidence that the order of loci was cn-B1, P2, and Pc. This raises the possibility of a paralogous gene set conditioning long glumes. A significant deviation from a 3:1 ratio in the F2 of LDN 7D(7B)/P2-LD222 confirmed the location of P2 on chromosome 7B. It is proposed that T. ispahanicum originated as a mutation of a gene affecting glume length on chromosome 7B of T. dicoccum, a spelt type of cultivated tetraploid wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Increase in grain protein percentage in high-yielding common wheat breeding lines by genes from wild tetraploid wheat

Summary Forty-one breeding lines of common wheat, derived from crosses between the Israeli cultivars Miriam and Lakhish and high-protein lines of wild tetraploid wheat, Triticum turgidum var. dicoccoides, were tested for various protein and yield parameters in field trials, under typical agronomic conditions. All lines had a higher grain protein percentage (GPP) than the leading Israeli cultivar Deganit, which was grown as a control. Grain yield (GY) ranged in the breeding lines from a low of 2.44 t/ha to as high as that of Deganit (6.95 t/ha). Despite the weak negative correlation between GPP and GY, several lines excelled both in GPP and in GY. The grain protein yield (GPY) of some of these selected breeding lines was higher than that of Deganit; e.g., 1.19 t/ha in the best line vs. 1.02 t/ha in Deganit. The 16.7% increase in GPY in this line reflected a more efficient utilization of nitrogen.     

STMS markers for grain protein content and their validation using near-isogenic lines in bread wheat

The present study, undertaken as a continuation of an earlier study on quantitative trait loci (QTL) analysis of grain protein content (GPC) in bread wheat (Prasad et al. 1999), includes the following: (1) identification of an additional molecular marker associated with GPC; (2) development of near‐isogenic lines (NILs) for high GPC; and (3) the use of three sets of NILs (a total of 10 NILs) to validate the two available markers linked with QTL for GPC. A total of 114 sequence‐tagged microsatellite site (STMS) primer pairs (that were not used in the previous study) were used for detection of polymorphism between the two parents (PH132, with high GPC; WL711, with low GPC) of a mapping population of 100 recombinant inbred lines (RILs). A total of 95 primer pairs gave amplification products, of which only 30 detected reproducible polymorphism between the parental genotypes. Bulked segregant analysis was conducted using these 30 primers on two bulks (each comprising eight RILs) representing the two extremes of the normal distribution. A solitary primer pair (WMC415) showed association with GPC, which was further confirmed through selective genotyping. Subsequently, 100 RILs were genotyped. A single‐marker linear regression analysis showed significant association between the marker WMC415 and GPC, thus identifying a quantitative trait locus (designated as QGpcccsu‐5A1), which explained 6.21% of the variation for GPC among the RILs. The above STMS marker, together with the STMS marker (WMC41) identified earlier, explains approximately 25% of the variation for GPC. In order to conduct validation of the above two available markers, 10 NILs were developed for high GPC using two genotypes (WL711 and HD2329) with low GPC as recipient parents and another two genotypes (PH132 and PH133) with high GPC as donor parents. NIL 2233 (with 11.7% GPC), derived from HD2329, when tried with WMC41 gave a characteristic amplification profile similar to that of its donor parent PH132, and NIL 2215 (with 11.9% GPC) derived from WL711, when tried with WMC415 gave an amplification profile that resembled its donor parent PH133. The remaining eight NILs with high GPC gave patterns similar to those of their corresponding recipient parents with both the markers, suggesting that either the QTL, other than those associated with the above markers, were actually transferred from the donor parents and contributed to high GPC in these NILs or that recombination had occurred between the markers identified and the corresponding QTL. Thus, the marker validation conducted using NILs, while demonstrating the utility of these two microsatellite markers for use in marker‐assisted selection in plant breeding, also suggested that many more QTL exist that would need to be identified using closely linked molecular markers.     

Genetic control of long glume phenotype in tetraploid wheat derived from Triticum petropavlovskyi Udacz. et Migusch.

The Chinese wheat landrace, Xinjiang rice wheat (T. petropavlovskyi Udacz. et Migusch., 2n = 42), known as ‘Daosuimai’ or rice-head wheat is characterized by long glumes, and was found in the agricultural areas in the west part of Talimu basin, Xinjiang, China in 1948. The gene for long glume from T. petropavlovskyi was introduced into a line of spring durum wheat, LD222. The gene for long glume is located approximately46.8 cm from the cn-A1 locus, which controls the chlorinatrait. Significant deviation from a 3:1 in the F₂ of LDN7D(7A)/ANW5C confirmed that the long glume of T. petropavlovskyi can be controlled by a gene located on chromosome 7A. The gene locates approximately 12.4 ± 0.5 cM from the centromere on the long arm of 7A. It is considered that the gene for long glume from T. petropavlovskyi is an allele on the P ₁ locus, and it should be designated as P _1a. It is suggested that T. petropavlovskyi originated from either the natural hybrid between T. aestivum that has an awn-like appendage on the glume and T. polonicum or a natural point mutation of T. aestivum. This revised version was published online in August 2006 with corrections to the Cover Date.     

Phosphorus use efficiency in tall, semi-dwarf and dwarf near-isogenic lines of spring wheat

The impact of the Rht dwarfing genes on P utilization efficiency (PUTE = grain dry matter per kg P in above-ground biomass), total P uptake (Pt) and related traits was studied in the varietal backgrounds of two tall wheat cultivars, Maringa and Nainari 60. Four sets of near-isogenic lines carrying different combinations of the alleles Rht-B1b, Rht-D1b and Rht-B1c for gibberellin-insensitive dwarfism in the hexaploid wheat (Triticum aestivum L.) were compared with tall controls in two field trials under conditions of adequate nutrient supply and irrigation in Northwest Mexico. The yield-increasing effect of the dwarfing genes Rht-D1b and Rht-B1b led to improved PUTE in Maringa and total P uptake in both cultivars. Also, the double dwarf line of Maringa had larger grain yields and P uptake compared to the tall control. The Rht-B1c genotypes showed low PUTE, thick roots and high P concentration in vegetative biomass indicating a surplus of assimilates and P, which could not be translocated into the grains. A similar problem could be observed in Nainari 60 with Rht-B1b and Rht-D1b, which produced the largest grain dry matter with the lowest P concentrations in grains although they showed high P accumulation in straw. Most of the net P uptake occurred before anthesis. P absorption after anthesis was more critical for the dwarf genotypes. For double dwarfs and Rht-B1c, respectively, only 3% and 21% of the total accumulated P at maturity was absorbed at post-anthesis. The grain P of the dwarf lines came more from P accumulated at pre-anthesis and translocated from the vegetative biomass into the grain. The pre-anthesis P accumulation was positively correlated with spikes per m² (r = 0.91), whereas post-anthesis P accumulation correlated better with grains per spike(r = 0.72), and thousand kernel weight (r = 0.51). P uptake efficiency played a secondary role under these non-P-limiting conditions, and differences in root length density were only slightly affected by Rht-genes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Radiation interception and radiation use efficiency of near-isogenic wheat lines with different height

Resistance to Pyrenophora teres Drechs. f. teres Smedeg., the net blotch pathogen, was studied in six 6-row Nordic spring barleys (Hordeum vulgare L.) in the field and in the greenhouse. The barley genotypes were: Arve, Agneta, Artturi, H6221, Pohto and WW7977. Disease progress was monitored in the field (1994 and 1995) in small artificially infected plots, sown at commercial seeding rates, and in infected hill plots (1994). Areas under the disease progress curves (AUDPC) and apparent infection rates (r) were calculated for the uppermost 3 or 4 leaves. Terminal severities (TS) were also recorded. Infection response of seedlings to a range of P. teres isolates was assessed in the greenhouse using a standard scale. In small plots in the field, Arve and Agneta were very susceptible to P. teres infection, as indicated by large values for AUDPC and TS. H6221 and WW7977 were highly resistant, while Artturi and Pohto were moderately resistant. In hill plots the situation was similar, except that Artturi and Pohto appeared less resistant than in the small plots. The relatively greater resistance of H6221 and WW7977 was reflected in seedling infection responses. According to the results of these experiments, H6221 and WW7977 possess adequate levels of quantitative resistance to P. teres to make them useful parents in future crossing programs aimed at improving net blotch resistance in Finnish spring barleys. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic mapping of sedimentation volume across environments using recombinant inbred lines of durum wheat

SDS-sedimentation volume (SV) is a biochemical index widely used to evaluate flour quality in durum and bread wheats. Significant association between SV and endosperm proteins (gliadin, high-molecular-weight- and low-molecular-weight-glutenin subunits) have been reported. Protein loci, however, account for only a portion of the total genetic variability. The objective of this study was to identify and locate quantitative trait loci (QTLs) associated with SV in a set of recombinant inbred (RI) lines, derived from a cross between the cv.‘Messapia’ of durum wheat and the accession MG4343 of the var. dicoccoides, and characterized for 259 genetic and molecular (RFLP) markers. Significant differences were detected for the quality index in the six environments examined, while the pattern of variability was that of a quantitative trait. Regression analysis of marker loci and sedimentation volume indicated, as expected, that chromosome 1B, on which are located the Gli-B 1/Glu-B 3 loci for some gliadin and glutenin subunits, is important for wheat quality. Two additional regions located on chromosomes 6AL and 7BS, and four regions on 1AL, 3AS, 3BL and 5AL, were shown to have single-factor effects on sedimentation volume at P < 0.001 and P < 0.01, respectively. Positive effects were contributed by both parents. A multiple linear regression model consisting of seven significant loci on different chromosomes explained 62–91% of the genotypic variation of the trait. The availability of linked markers to QTLs may facilitate the genetic dissection of quantitative traits and the early selection in wheat breeding programmes.     

Development of near-isogenic lines for a major QTL on 3BL conferring Fusarium crown rot resistance in hexaploid wheat

By essentially fixing the genetic background, near-isogenic lines (NILs) are ideal for studies of the function of specific loci. We report in this paper the development of NILs for a major QTL located on the long arm of chromosome 3B conferring Fusarium crown rot (FCR) resistance in hexaploid wheat. These NILs were generated based on the method of the heterogeneous inbred family analysis. 13 heterozygous lines were initially selected from three segregating populations using a single SSR marker linked with the major FCR QTL. The two isolines for each of the putative NILs obtained showed no obvious morphological differences, but differences among the NIL pairs were large. Significant differences in FCR resistance between the isolines were detected for nine of the 13 putative NIL pairs. The presence of the FCR allele from the resistant parent reduced FCR severity by 29.3–63.9% with an average of 45.2% across these NILs. These NILs will be invaluable in further characterising this major FCR locus, in studying the mechanism of FCR resistance and in investigating possible interactions between FCR resistance and other traits of agronomic importance.     

Comparison of near-isogenic lines for anthracnoseAre are genes in common bean,Phaseolus vulgaris L.

Summary This study was undertaken to determine whether the Are gene controlling anthracnose resistance had an adverse effect on maturity and yield because backcross-derived cultivars/lines had numerically lower yield and later maturity than the respective recurrent parent. Three pairs of common bean lines near-isogenic for theAre gene were developed fromAre are F₂ plants of the sixth backcross of Seafarer, Fleetwood, and Ex Rico 23. The near-isogenic lines were tested along with the recurrent cultivars and backcross-derived cultivars/lines. The results showed no evidence of any adverse effect of theAre gene on the agronomic characteristics in 11 trails at 5 locations in 3 years.     

Fine mapping of open-bud duplicate genes in homoelogous chromosomes of tetraploid cotton

The open-bud mutant plant has a flower bud that is open at its tip due to the shortening of the corollas, resulting in exposure of the stigma and upper anthers. This mutant plant is potentially useful as a parent for producing hybrid cotton seeds. We have identified the open-bud trait to be inherited as recessive duplicate genes and finely mapped them in the homoelogous chromosome (chr) pair of chr. 18 (D13) and 13 (A13). Gossypium hirsutum substitution line 18, which has chr. 18 replaced by its corresponding homozygous chromosome in G. barbadense acc. 3-79 and the genotyped open-bud duplicate genes ob ₁ ob ₁ ob ₂ ob ₂, was crossed with the monomeric genotypic G. hirsutum acc. TM-1 and G. barbadense acc. 3-79 to produce two mapping populations. We then fine-mapped the duplicate open-bud gene ob ₁ in the chr. 18 (D13) and ob ₂ in the chr. 13 (A13) homoelogous chromosome pair in tetraploid cotton. Molecular markers closely linked with ob ₁ and ob ₂ will be useful tools in the development of open-bud lines by marker-assisted selection.     

Development of near-isogenic lines of wheat carrying different null Wx alleles and their starch properties

The granule-bound starch synthase (GBSS I) encoded by the Wxgenes, is involved in amylose synthesis. For analyses of mechanisms of amylose synthesis and associated starch properties in hexaploid wheat, eight possible genotypes having different combinations of the three null alleles at the Wx loci with a common genetic background are a prerequisite. A near-isogenic population of doubled haploid (DH) lines was produced from Chinese Spring × waxy Chinese Spring F₁ plants using the wheat × maize method. The Wx protein phenotypes of the DH progeny were examined by sodium dodecyl sulphate-polyacrylamide gel electrophoresis and found that the null alleles at each of the three Wx loci segregated in a Mendelian fashion. A field trial demonstrated no differences between the eight types for ear emergence time, plant height and grain yield traits. Amylose content in the endosperm starch was highest in the wild type while lowest in the waxy type having no Wx proteins. Comparison between single null types and double null types indicated that the amylose synthesis capacity of Wx-A1a allele is the lowest. Pasting properties of starch are the highest in the waxy type, followed by the double null types. Consequently, both peak viscosity and breakdown were negatively correlated with amylose content. The chain-length distribution analysis of amylopectin structure revealed no clear difference among the eight types,suggesting that the reduced GBSS I activity due to introgression of the null Wx alleles does not affect either the chain length or the degree of branching of amylopectin. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic mapping of the genes for glaucous leaf and tough rachis in Aegilops tauschii, the D-genome progenitor of wheat

Glaucous leaf and tough rachis phenotypes are rare in Aegilops tauschii, the D genome donor to common wheat (Triticum aestivum). The genes for glaucous leaf and tough rachis were mapped using microsatellite probes in A. tauschii. The glaucous phenotype was suppressed by the inhibitor W2^I located on chromosome 2DS. The gene W2^I was mapped to the distal part of 2DS, and was unlinked to the centromere. This suggests that the distance of the W2^I locus from the centromere was maintained during the evolution of hexaploid wheat from its diploid progenitors as the inhibitor gene is at the same position in A. tauschii and bread wheat. The Br^t (Brittle rachis of A. tauschii) locus was located on the short arm of chromosome 3D, and was 19.7 cM from the centromeric marker, Xgdm72.3D. Br^t causes breakage of the spike at the nodes, thus creating barrel-shaped spikelets, while Br₁ in hexaploid wheat causes breakage above the junction of the rachilla with the rachis such that a fragment of rachis is attached below each spikelet.     

Molecular and physical mapping of genes affecting awning in wheat

Quantitative trait loci (QTL) for three traits related to awning (awn length at the base, the middle and the top of the ear) in wheat were mapped in a doubled‐haploid line (DH) population derived from the cross between the cultivars ‘Courtot’ (awned) and ‘Chinese Spring’ (awnless) and grown in Clermont‐Ferrand, France, under natural field conditions. A molecular marker linkage map of this cross that was previously constructed based on 187 DH lines and 550 markers was used for the QTL mapping. The genome was well covered (more than 95%) and a set of anchor loci regularly spaced (one marker every 20.8 cM) was chosen for marker regression analysis. For each trait, only two consistent QTL were identified with individual effects ranging from 8.5 to 45.9% of the total phenotypic variation. These two QTL cosegregated with the genes Hd on chromosome 4A and B2 on chromosome 6B, which are known to inhibit awning. The results were confirmed using ‘Chinese Spring’ deletion lines of these two chromosomes, which have awned spikes, while ‘Chinese Spring’ is usually awnless. No quantitative trait locus was detected on chromosome 5A where the B1 awn‐inhibitor gene is located, suggesting that both ‘Courtot’ and ‘Chinese Spring’ have the same allelic constitution at this locus. The occurrence of awned speltoid spikes on the deletion lines of this chromosome suggests that ‘Chinese Spring’ and ‘Courtot’ have the dominant B1 allele, indicating that B1 alone has insufficient effect to induce complete awn inhibition.