Inheritance of agronomic traits from the Chinese barley dwarfing gene donors 'Xiaoshan Lixiahuang' and 'Cangzhou Luodama'

The inheritance of agronomic traits from the barley dwarfing gene donors ‘Xiaoshan Lixiahuang’ and ‘Cangzhou Luodamai’ was studied. The results indicated that dwarf plants, six‐row and short spikes, dense spikelets and naked kernels, respectively, were controlled by one pair of recessive genes, but a toothed awn was determined by one pair of dominant genes in both barley cultivars. The genes for the six characters in ‘Xiaoshan Lixiahuang’ were allelic to those in ‘Cangzhou Luodamai’. Genetic linkage was found among the genes for plant height, spike length and spikelet density. They were located on the long arm of chromosome 3 (3HL) in the order: plant height, spikelet density, spike length. The genes for naked kernels, six‐row spikes and tooth awns were independent of each other, and carried on the long arms of chromosomes 1(7H), 2(H) and 7(5H), respectively. The dwarfing genes were the same as the gene uz in Japanese and Korean barley cultivars.     

Inheritance of resistance to Pyrenophora teres f. teres in spring barley

The inheritance of seedling resistance to a Swedish isolate of Pyrenophora teres f. teres was investigated in four resistance sources of spring barley. Accessions CI 2330, CI 5791, CI 5822 and CI 9779 were used as resistance sources, and the cultivar ‘Alexis’ was used as a susceptible parent in different crosses. From the disease reaction in the F₁, F₂ and F₃ generations it was concluded that the resistance was governed by the same two complementary genes in CI 5791, CI 822 and CI 9776. One of these genes was present in CI 2330. The first three cultivars were highly resistant to the isolate used in this investigation. These results, when combined with earlier studies, suggest that CI 5791, CI 5822 and CI 9776 may be of great value as sources of resistance to barley net blotch. Spearman's rank correlation between the disease reaction of F₂ plants and their F₃ progeny was highly significant (r = 0.75; P ≥ 0.001) It is suggested that selection in the F₂ generation is effective. In a backcross breeding scheme, single plant reactions in F₁ or F₂ need to be confirmed in later generations.     

Inheritance and allelism of Russian wheat aphid resistance in several wheat lines

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko) has caused serious reduction in wheat production in 17 Western states of the United States since 1986. Inheritance of resistance to RWA in seven wheat lines and the allelism of the resistance genes in these lines with three known resistance genes Dn4, Dn5, and Dn6 were studied. The seven resistant lines were crossed to a susceptible wheat cultivar ‘Carson’ and three resistant wheats: CORWA1 (Dn4), PI 294994 (Dn5), and PI 243781 (Dn6). Seedlings of the parents, F₁, and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21–28 days after the infestation using a 1–9 scale. The resistance level of all the F₁ hybrids was similar to that of the resistant parent, indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ population, suggesting qualitative, nonadditive gene action, in which the presence of any one of the dominant alleles confers complete resistance to RWA. Resistance in CI 2401 is controlled by two dominant genes. Resistance in CI 6501 and PI 94365 is governed by one dominant gene. Resistance in PI 94355 and PI 151918 may be conditioned by either one dominant gene or one dominant and one recessive gene. No conclusion can be made on how many resistance genes are in AUSVA1-F₃, since the parent population was not a pure line. Allelic analyses showed that one of resistance genes in CI 2401 and PI 151918 was the same allele as Dn4, the resistance gene in CI 6501 was the same allele as Dn6, and AUS-VA1-F₃ had one resistance gene which was the same allele as one of the resistance genes in PI 294994. One non-allelic resistance gene different from the Dn4, Dn5, and Dn6 genes in CI 2401, PI 94355, PI 94365, and PI 222668 was identified and should be very useful in diversifying gene sources in wheat breeding.     

Inheritance of nitrogen and phosphorus utilization efficiency in spring barley at the vegetative growth stages under high and low nutrition

Inheritance of the nitrogen and phosphorus utilization efficiencies (NUE and PUE, respectively) and that of the tolerance (T) to limited NP nutrition was investigated in spring barley crosses at the vegetative growth stages. Plants were grown in sand-vermiculite cultures under high and low NP nutrition. In a diallel set (Ps and F₂s), both general combining ability (GCA) and specific combining ability (SCA) effects were significant for the variation in NUE and PUE, while the variation in T was mainly associated with GCA effects. The contribution of nonadditive genes for the utilization efficiencies was found to increase under nutrient shortages. Overdominance of genes was detected. The characters exhibited low heritabilities (0.10-0.42). Generation means analysis in two cross-combinations revealed significant effects of epistatic gene interactions. It was assumed that the involvement of both the dominance effects and epistatic interactions would not facilitate selection efforts to improve the characters in spring barley at its vegetative growth. Such selection should be performed among families of later generations. The genotype-nutrition interactions observed suggest that selection under diverse nutrition rates would be necessary for the more precise evaluation of barley efficiency under less favourable soil fertility.     

Inheritance and allelism for resistance to Russian Wheat Aphid in an Iranian Spring Wheat

Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), is an important pest of wheat (Triticum aestivum L.) in the United States of America. Developing adapted wheat cultivars with genetic resistance to RWA is an effective control strategy. Genetic studies were conducted to determine the mode of inheritance of gene(s) conferring resistance to RWA in an Iranian landrace wheat line, G 5864. For the inheritance study, G 5864 was crossed with the susceptible wheats ‘Yecora Rojo’ and ND 2375. Seedlings of F1, reciprocal F1, F2, BC1 to the susceptible parent (BCS), and BC1 to the resistant parent (BCR) were screened for RWA reaction. Several phenotypic segregation ratios were tested in the F2 populations for goodness of fit; the 9:3:3:1 ratio (resistant: rolled leaves: stunted plants: susceptible) was an acceptable fit in all cases. Thus, resistance in G 5864 seemed to be controlled by two independent dominant genes with additive gene effects. The allelic relationships of gene(s) in this line with genes in other resistant lines, PI 137739 (Dn1), PI 262660 (Dn2), PI 372129 (Dn4), PI 294994 (Dn5), and PI 243781 (Dn6), were also studied. Segregation patterns observed in G 5864 × resistant (R × R) F2 populations were inconclusive. However, no susceptible plants were observed in these F2 populations. If previous reports concerning the number of resistance genes present in the other resistant lines are correct, then given the high manifestation of resistance observed in G 5864, and given the absence of susceptible plants in the R × R F2 populations, it is indicated that RWA resistance in G 5864 is either controlled by different alleles at the same loci as the other resistance genes, or that G 5864 shares a resistance gene with each of the other resistant lines. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance to powdery mildew in selections from Tunisian landraces of barley

Two-hundred and thirty-two accessions of barley landraces collected from Tunisia were screened for resistance to powdery mildew. A number of race-specific genes were detected using the detached leaf technique. Among the 232 accessions tested, 169 were susceptible to powdery mildew, 20 were resistant and 43 showed differential reactions to the three isolates of powdery mildew used. An attempt was made to determine the number of genes, the types of gene, the types of gene action and the gene loci in 20 resistant accessions. Three types of cross were made: (1) the accessions were crossed to the susceptible variety ‘Pallas’, (2) the accessions were crossed with ‘Pallas’ isolines, and (2) accessions with identical powdery mildew reaction patterns were intercrossed. Three isolates of Erysiphe graminis f. sp. hordei were used: Bzm-1, KM 18-75, R13C. A number of different resistance genes were detected among the 19 resistant accessions. Surprisingly, segregation indicating single genes only were detected with the isolates used. Some of these genes could be associated with loci already known. In 19 cases a dominant and in one a recessive mode of inheritance was detected. The recessive gene was not located at the mlo locus. This investigation represents the first systematic study of race-specific genes for powdery mildew resistance in Tunisian landraces. The newly identified sources of resistance may be used in many strategies of breeding for disease resistance.     

Inheritance and allelism of resistances to the Russian wheat aphid in seven wheat lines

Summary Studies were conducted to determine the inheritance and allelic relationships of genes controlling resistance to the Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), in seven wheat germplasm lines previously identified as resistant to RWA. The seven resistant lines were crossed to a susceptible wheat cultivar Carson, and three resistant wheats, CORWA1, PI294994 and PI243781, lines carrying the resistance genes Dn4, Dn5 and Dn6, respectively. Seedlings of the parents, F₁ and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21 to 28 days after the infestation using a 1 to 9 scale. All the F₁ hybrids had equal or near equal levels of resistance to the resistant parent indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ segregation suggesting major gene actions. The resistance in PI225262 was controlled by two dominant genes. Resistance in all other lines was controlled by a single dominant gene. KS92WGRC24 appeared to have the same resistance gene as PI243781 and STARS-9302W-sib had a common allele with PI294994. The other lines had genes different from the three known genes.     

Evidence of allelism between genes Pm8 and Pm17 and chromosomal location of powdery mildew and leaf rust resistance genes in the common wheat cultivar'Amigo'

TIBL-1RS wheat-rye translocation cultivars utilized in wheat programmes worldwide carry powdery mildew resistance gene Pm8. Cultivar‘Amigo’possesses resistance gene Pm17 on its TIAL-1RS translocated chromosome. To be able to use Pm17efficiently in breeding programmes, this gene was transferred to a TIBL-1RS translocation in line Helami-105, and allelism between Pm8 and Pm17was studied. The progenies of the hybrids in the F₂ generation and F₃ families provided evidence that the two genes are allelic. Genetic studies using monosomic analyses confirmed that in cultivar‘Amigo', Pm17 and leaf rust resistance gene Lr24 are located on a translocated chromosome involving 1 A and 1B, respectively.     

Inheritance and chromosome location of Alp, a gene controlling aluminum tolerance in 'Dayton' barley

Aluminum (Al) toxicity is a major limiting factor in acid soils and more adequate genetic tolerance is needed to improve barley adaptation and production in affected regions. To study the inheritance and chromosome location of the Alp gene controlling Al tolerance in ‘Dayton’ barley the primary trisomics of sensitive ‘Shin Ebisu 16’ were crossed to ‘Dayton’. Parental, F₁ and F₂ seedlings were grown in nutrient solution containing 0.03, 0.06 and 0.09mM Al. and classified for tolerance by haematoxylin staining of the roots. In diploid F₂ progeny, Alp was inherited as a single gene, dominant at 0.06mM and recessive at 0.09 mM concentrations, as indicated by the 3:1 and 1:3 (tolerant: sensitive) segregation ratios, respectively. Segregation of the trisomic Frderived F₂ seedlings at 0.06mM Al deviated significantly from the 3:1 only for the triplo 4/‘Dayton’ cross. Data for this cross fit the expected trisomic ratios, indicating that the Alp gene is distally located from the centromere on chromosome 4. These results confirm that tolerance is simply inherited, but expression of tolerance is dependent on Al concentration and allele dose.     

Parial resistance in the barley (Hordeum vulgare L.) cultivar 'Chikurin Ibaraki 1' to two PAV-like isolates of barley yellow-dwarf virus: allelic variability at the Yd2 gene locus

The Japanese barley cultivar, ‘Chikurin Ibaraki 1’, is partially resistant to the PAV serotype of barley yellow-dwarf virus (BYDV), but its induced mutant line, Ea52, is susceptible. The inheritance of resistance in cv. ‘Chikurin Ibaraki 1’ to BYDV-PAV was investigated. The F, and F2 plants of crosses of cvs ‘Chikurin Ibaraki 1’, Ea52, ‘Vixen’, carrying the Yd2 gene of resistance, and ‘Plaisant’, a susceptible French cultivar, were tested in growth chamber and field conditions. Isolate RG, against which ‘Chikurin Ibaraki 1’ is partially resistant in growth chamber and field conditions, and isolate 2t, which overcomes the partial resistance of ‘Chikurin Ibaraki 1’ in field conditions (Chalhoub et al. 1994) were used. The segregation of F2 plants of crosses between ‘Chikurin Ibaraki 1’ and the susceptible cultivars to isolate RG (one resistant to three susceptible) suggests that the resistance of ‘Chikurin Ibaraki 1’ is controlled by a single recessive gene. All 537 F2 plants of ‘Chikurin Ibaraki 1’בVixen’ tested with isolate RG in growth chamber and field conditions were resistant. The F2 plants of this cross were all resistant to isolate 2t in growth chamber conditions but segregated with a ratio of one resistant to three susceptible in field conditions owing to the susceptibility of ‘Chikurin Ibaraki 1’ to this isolate. Results suggest that the resistance gene in ‘Chikurin Ibaraki 1’ is tightly linked or allelic with the Yd2 gene in ‘Vixen’. However, it differs from this gene in ‘Vixen’ in that it can be overcome by isolate 2t in field conditions.     

Inheritance of heading time in spring barley evaluated in multiple environments

The inheritance of heading time of spring barley was studied in three extremely early genotypes IB, RL and ‘Mona’ (M), which is homozygous recessive for the early maturity ea8 (=ea_k) gene conferring extreme earliness under short daylengths and is relatively photoperiod insensitive, and five (GP, MA, PS, NU and BA) spring genotypes that are early to intermediate for heading time. Frequency distributions of F₂ generations grown at Ouled Gnaou, Morocco (32°15′ N), an environment which maximizes differences between photoperiod‐insensitive and photoperiod‐sensitive genotypes, indicated that across populations many loci were segregating in a complex Mendelian manner. IB and RL were both homozygous recessive for the ea8 gene, which conferred an early heading time. RL had partially dominant alleles at second locus (Enea8), which enhanced its earliness. Recovery of only progeny within the parental range of genotypes for heading time from the crosses of RL/M and IB/M suggests that numerous loci remained suppressed, perhaps latent, given their diverse parentage. The ea8 recessive homozygote in RL suppressed another unidentified locus which, when homozygous recessive in the absence of the ea8 recessive homozygote, conferred extreme earliness in one short daylength environment (Ouled Gnaou, Morocco) but was undetected in another environment (Davis, CA, USA). Epistatic gene action and genotype × environment effects strongly influenced heading time. In addition to a genetic system consisting of single‐locus recessive homozygotes conferring photoperiod insensitivity, a second genetic system, based on dominant alleles at one or a few loci, derived from the early heading Finnish landrace ‘Olli’, also confers extremely early heading time under short daylengths and relative photoperiod insensitivity in the genotype GP.     

Genetic analysis of the nitrogen and phosphorus utilization efficiencies in mature spring barley plants

Inheritance of the efficiency of nitrogen and phosphorus utilization in grain production was studied in diallel hybrids of spring barley. Effects of varied NPK fertilization and soil moisture on the genetic variance was evaluated at maturity in pot‐ and field‐grown plants. A preponderance of general combining ability effects was found for the efficiency ratios and utilization efficiencies, suggesting that the generative efficiency indices were mainly controlled by an additive genetic system. However, the relatively high narrow‐sense heritabilities (0.33‐0.81) found under optimal conditions tended to decline to 0.00‐0.36 following water and nutrient shortages. The importance of non‐additive gene effects for utilization efficiencies was demonstrated in the field under reduced nutrition. This leads to the conclusion that the involvement of additive gene action should facilitate selection efforts only under favourable environments. Selection delayed until later hybrid generations should be used when trying to improve the nutrient efficiency of barley under less favourable nutrition conditions.     

Application of quantitative trait locus mapping to the development of winter-habit malting barley*

Malting quality and winter-hardiness in barley are ‘ultimate’ phenotypes composed of constituent quantitatively inherited traits. A synthesis of molecular-marker linkage data and field phenotyping to reveal the location of quantitative trait loci (QTL) may assist in the development of winter-habit malting barley varieties. One-hundred doubled haploid progeny from a winter x spring cross were evaluated under fall and spring-planted conditions. Malting quality phenotypes and a 76-point map were used to identify QTL and to assess the effect of spring-and autumn-sowing on QTL expression. Many QTL effects were common to both environments and corresponded to QTL detected in other barley germplasm. While there were significant differences in the magnitude of effects across environments, there were no changes in the favourable allele phase. QTL effects for grain protein and diastatic power level coincided with the locations of known function genes. Coincident QTL for a number of mailing-quality traits on chromosome 7 suggests either the presence of a multi-locus cluster of genes controlling components of malting quality or a regulatory gene or genes controlling the cascade of enzymatic processes that function during the malting process. Based on these results, simultaneous selection for malting quality and cold tolerance should be possible in this genetic background.     

Inheritance of plant height in the dwarf mutant 'Enana' of safflower

The dwarf safflower mutant ‘Enana’ has been developed from the cultivar ‘Rancho’ by chemical mutagenesis. The objective of the present research was to study the inheritance of plant height in crosses between ‘Enana’ and ‘Rancho’. Plants of both lines were reciprocally crossed and the F₁, F₂ and F₃ generations were obtained. The evaluation of plant height in the F₂ generation suggested the presence of a single locus controlling this trait. This was confirmed after the evaluation of 164 F_2:3 lines, which revealed an F₂ segregation fitting a 1:2:1 (dwarf: intermediate: standard) ratio. The locus was designated D^w. As the mutated allele was partly dominant over the wild‐type allele, the proposed genotype for ‘Rancho’ was d^w d^w, whereas that for ‘Enana’ was D^w D^w. Furthermore, a partial cytoplasmic effect on plant height was detected, with the heterozygote D^w d^w being about 6 cm taller when ‘Rancho’ was used as female.     

Generation Mean Analysis of Inheritance of Resistance to Pyrenophora teres in Barley

F1, F2, Fl × parentl (BC1), and Fl × parent2 (BC2) generations resulting from four crosses among seven cultivars of barley used in national and international breeding programmes were tested at the seedling stage for their resistance to a mixture of five isolates of Pyrenophora teres. Four methods were used to assess disease resistance: infection type (IT), average lesion size (ALS), number of lesions per unit leaf area (NL) and percent leaf area infected (PLAI). Gene actions were estimated by generation mean analysis on each of the four crosses and on each of the evaluation methods. Significant additive and additive × additive epistatic effects were found. Infection type and percent leaf area infected were found to be highly correlated in all four crosses. These results suggest that barley breeders could improve the level of resistance to P. teres by making appropriate crosses between highly susceptible barley cultivars.     

Allelic variation at the dwarfing gene Rht8 locus and its significance in international breeding programmes

A microsatellite marker WMS261 developed at I.P.K., Gatersleben,Germany, co-segregates with the semi-dwarfing gene Rht8. Screens ofover 800 wheat varieties from 20 countries show 90% carry WMS-261alleles with 165, 174 or 192 base pairs (bp). The 192bp allele diagnosticof Rht8 occurs in most screened Southern European varieties. Anallele with 165 bp occurs in the majority of CIMMYT Mexican varieties andin most varieties bred in countries that utilize CIMMYT germplasm.Agronomic studies of single chromosome recombinant line mappingpopulations segregating for WMS 261 alleles with 165, 174 or 192 bp,show compared to the 174bp allele the 192bp allele reduces height byaround 8 cm and the 165 bp allele increases height by around 3cm. Theresults indicate the importance and widespread utilization of Rht8 asa dwarfing gene in southern Europe and suggest an adaptative significanceto the height promoting 165 bp allele in CIMMYT material.     

New sources of resistance for stripe rust in barley

Eight spring barley accessions from the gene bank in Gatersleben, Germany, and 10 cultivars were tested for stripe rust resistance. Tests were performed at the seedling stage in the growth chamber and as adult plants in the field. All accessions and six cultivars were scored as resistant against race 24 under all test conditions, with very few plants as exceptions, while the susceptible control cultivars ‘Karat’ and ‘Certina’, and four other cultivars were attacked in all cases. Differences between accessions and between cultivars were detected after infection with isolates from ‘Trumpf’ and ‘Bigo’ (seedling tests only). Infection structures within seedling leaves without pustules and for the first time within leaves of adult plants from the field were analysed by fluorescence microscopy. With this method additional genetic Differences in the resistance reaction could be detected which could not to be seen in the resistance test. Crosses between the accessions and the susceptible cultivar ‘Karat’ led to segregating F₂ progenies. The percentage of resistant plants varied between the accessions. This also indicates a different genetic basis of resistance in the accessions. The infection structures observed by fluorescence microscopy stopped earlier in leaves of the two accessions HOR 8979 and HOR 8991 than in leaves of other accessions in all the tests. These accessions were the only ones with more than 50% resistant plants in all F₂ tests. In general, the accessions from the gene bank can be used as new resistance sources against stripe rust.     

In vitro quantification of the reaction of barley to common root rot

An in vitro technique was used to quantify the infection level of common root rot. This disease produces a brown to black discoloration of the subcrown internodes of barley. Quantification was based on the percentage of germinated infected pieces (1.5 mm) of subcrown internodes cultured on potato dextrose agar media. The disease severity was apparent among four different visually classified categories and numerical values for each category were applied. The results were highly correlated (r = 0.97, P < 0.01) among the different in vitro experiments, indicating that this testing procedure is repeatable. Highly significant differences (P < 0.001) were found for the length of first leaf and fresh weight between plants inoculated and uninoculated with common root rot. However, the effect of inoculation on fresh weight only differed significantly (P < 0.02) among the genotypes.     

Effects of selection and opportunities for recombination in doubled-haploid populations of barley (Hordeum vulgare L.)

Doubled-haploid breeding systems are typically based on sampling gametes from F₁ plants. However, in the case of repulsion linkages, additional recombination could be advantageous. Pre-selection of gamete donors might also shift progeny performance in a desired direction. The objectives of this study were to quantify the effects of an additional round of recombination and assess the effectiveness of pre-anthesis selection in the production of barley doubled haploids. Assessments were conducted on: 1. 100 F₁-derived lines representing a subset of lines used in previous genome-mapping studies; 2. 100 random F₂-derived lines; and 3. 50 F₂-derived lines from gamete donors selected for early heading. An additional round of recombination had only a modest effect on generating more favourable genotypes. Pre-anthesis selection was ineffective in generating an earlier heading population. According to published quantitative-trait locus (QTL) analyses based on the F₁ -derived population, there are few repulsion linkages between QTL determining the traits measured in this experiment. Any advantages to be gained from postponing the generation of derivation of doubled haploids must be weighed against the delay and additional cost.     

The effect of major dwarfing genes on yield potential in spring wheats

Summary A composite convergent cross of 16 spring wheat parents produced a set of unselected progeny lines among which the major dwarfing genes, Rht1, Rht2 and Rht3, were distributed against a common random genetic background. Random subsets of these lines were grown under irrigation and optimal conditions in 4 experiments with replicated bordered plots in southern New South Wales in order to measure the dwarfing gene effect on yield potential. The dwarfing gene composition of each line was determined by test crossing and seedling responsiveness to gibberellic acid.Lodging was negligible in the two experiments in 1982. While present in the two in 1983, it was not strongly associated with yield. Grain yield levels were appropriately high (mean 5.9 t/ha). In all but 1 experiment the Rht1+Rht2 dwarf genotypes gave highest yields while the Rht3 group yielded on average 3% lower, Rht2 9% lower, Rht1 11% lower, and the non-dwarf or tall group yielded 24% lower. These yield differences were positively associated with harvest index, kernels per m² and kernels per spike, but negatively associated with mature plant height. Even within major dwarfing gene classes, grain yield was significantly and negatively associated with height.