Dormancy breaking procedures and the breeding of white-grained wheat with resistance to pre-harvest sprouting

Summary Gibberellic acid, cold (4°C) and a combination of these two treatments were tested for use in breaking dormancy in 27 lines of white-grained wheat with varying levels of resistance to pre-harvest sprouting. Germination increased in all lines treated with gibberellic acid. Dormancy could be broken with 1 M gibberellic acid. Response to cold varied. A combination of gibberellic acid and cold treatment was the most effective. This technique has been found useful in treating seed in a breeding program aimed at producing wheats with resistance to pre-harvest sprouting.     

Novel resistance to pre-harvest sprouting in Australian wheat from the wild relative Triticum tauschii

The diploid D-genome progenitor of hexaploid wheat, Triticum tauschii (Coss.) Schmahl., was screened to identify mechanisms for resistance to pre-harvest sprouting. A number of promising mechanisms were identified, and transferred to hexaploid wheat via wide-hybridisation. One identified mechanism, an inhibitory phenolic compound present in the bracts surrounding the grain, has been shown to function effectively in synthetic hexaploid wheats. A number of seed-borne dormancy mechanisms were also identified. Expression of embryo dormancy in synthetic hexaploid wheats was demonstrated when compared with non-dormant hexaploid wheat. Effects of the seed coat on dormancy were also studied, with the seed coat of synthetic hexaploids accelerating rather than inhibiting germination. Embryo dormancy was also demonstrated in two `direct-cross' hybrids. The results suggest that a combination of the described mechanisms may produce white wheats with resistance to pre-harvest sprouting adequate for most Australian climatic conditions. This revised version was published online in August 2006 with corrections to the Cover Date.     

QTL analysis and mapping of pre-harvest sprouting resistance in Sorghum

One of the most important agronomic problems in the production of sorghum [Sorghum bicolor (L.) Moench] in humid climates is pre-harvest sprouting (PHS). A molecular linkage map was developed using 112molecular markers in an F₂ mapping population derived from a cross between IS 9530 (high resistance to PHS) and Redland B2 (susceptible to PHS). Two year phenotypic data was obtained. By means of interval mapping analysis, two significant QTL were detected in two different linkage groups with LOD scores of 8.77and 4.39. Each of these two QTL individually explained approximately 53% of the phenotypic variance, but together, in a two-QTL model, they explained 83% of the phenotypic variance with a LOD score of 12.37.These results were corroborated by a one-way ANOVA in which the four flanking markers of the most likely QTL positions displayed highly significant values in theF-test, and significant variation in trait expression was associated with marker genotypic classes. The four markers with highest effect in the one-way ANOVA were also detected in the second year replication of the F₂ population, and significant genotype × environment interactions was observed. The putative relationship between PHS resistance in sorghum and the maize Vp1 gene is also discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Detection of QTLs for traits associated with pre-harvest sprouting resistance in bread wheat (Triticum aestivum L.)

Pre-harvest sprouting (PHS) is one of the serious problems for wheat production, especially in rainy regions. Although seed dormancy is the most critical trait for PHS resistance, the control of heading time should also be considered to prevent seed maturation during unfavorable conditions. In addition, awning is known to enhance water absorption by the spike, causing PHS. In this study, we conducted QTL analysis for three PHS resistant related traits, seed dormancy, heading time and awn length, by using recombinant inbred lines from ‘Zenkouji-komugi’ (high PHS resistance) × ‘Chinese Spring’ (weak PHS resistance). QTLs for seed dormancy were detected on chromosomes 1B (QDor-1B) and 4A (QDor-4A), in addition to a QTL on chromosome 3A, which was recently cloned as TaMFT-3A. In addition, the accumulation of the QTLs and their epistatic interactions contributed significantly to a higher level of dormancy. QDor-4A is co-located with the Hooded locus for awn development. Furthermore, an effective QTL, which confers early heading by the Zenkouji-komugi allele, was detected on the short arm of chromosome 7B, where the Vrn-B3 locus is located. Understanding the genetic architecture of traits associated with PHS resistance will facilitate the marker assisted selection to breed new varieties with higher PHS resistance.     

On the problem of pre-harvest sprouting of wheat

Summary The problem of pre-harvest sprouting in wheat is very intricate. New concepts in breeding for resistance to pre-harvest sprouting regard many components such as -amylase production potential, response to gibberellic acid and inhibition in the bracts as important as dormancy. In the research work carried out at The University of Sydney's Plant Breeding Institute, we found varietal differences for each of these components. We also developed criteria for screening breeding material for these components. The implications of our findings in planning efficient breeding programmes to evolve varieties with multiple resistance to pre-harvest sprouting have been discussed briefly.     

Relationship between sprouting in wheat and embryo response to endogenous inhibition

Summary Seed dormancy in wheat (Triticum aestivum L. em Thell) is important for minimizing pre-harvest sprouting. To facilitate breeding cultivars that tolerate pre-harvest sprouting conditions, we assessed mode and magnitude of variation of seed dormancy among genotypes and investigated involvement of endogenous water-soluble inhibitor(s) in seed germination. Embryo bio-assays established that water-soluble inhibitor was ubiquitous among the wheat cultivars studied and did not diminish in quantity during after-ripening. Germination response of embryos was decreased by endogenous inhibitor, but the effect markedly declined as embryos aged at room temperature. Variation in dormancy among cultivars was primarily caused by differential response of their embryos to endogenous inhibitor. Gibberellic acid counteracted the initial inhibitory effect of endogenous inhibitor on germination but not the subsequent inhibitory effect on seedling growth. We concluded that pre-harvest sprouting resistance involves multiple factors, particularly embryo receptivity to endogenous inhibitor, and that variation in inhibitor quantity is not solely responsible for genotypic differences in susceptibility to pre-harvest sprouting. The possibility of additional approaches to breeding for pre-harvest sprouting resistance is indicated.Contribution no. 81-389-j, Department of Agronomy, Kansas State University, Manhattan, KS 66506, U.S.A.     

Utilization of tom thumb source of pre-harvest sprouting tolerance in a wheat breeding programme

Summary Segegating populations of the cross Tordo (a spring white-grained Tom Thumb derivative) and Tr 454-16 (a spring white-grained Sonora-Gabo derivative) were studied for plant height, apparent -amylase synthesis (AAS) and response to gibberellic acid. Means, variances and heritabilities for these characters were computed and their association was studied by computing correlation coefficients. Plant height showed high heritability while AAS and response to GA₃ showed moderate heritability. Plant height was positively correlated with AAS and response to GA₃. The results pointed out the importance of using Tom Thumb as a source of pre-harvest sprouting tolerance in a wheat breeding programme, particularly in areas where wheat breeding is restricted to evolving white-grained cultivars.     

Germplasm improvement for preharvest sprouting resistance in Chinese white-grained wheat: An overview of the current strategy

Most modern white-grained wheat (Triticum aestivum L.) cultivars in China lack sprouting resistance. Some white grain varieties showed lower preharvest sprouting percentage in different years. Tolerance to sprouting in most cultivars comes from the varieties, Wanxian White, Fulingxuxu White, Suiningtuotuo, and Yongchuan White. The α-amylase inhibitor gene under a wheat genetic background inhibited α-amylase activity and delayed sprouting. The falling number of six suspected sources, out of 835Chinese leading varieties since 1950', was independent of late maturityα-amylase. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of resistance to Russian wheat aphid in barley

Summary The inheritance of resistance to Russian wheat aphid Diuraphis noxia (Mordvilko), in two resistant barleys, Hordeum vulgare L., ASE/2CM//B76BB and Gloria/Come, was studied in the field and in the greenhouse. The resistant genotypes were crossed with susceptible genotypes Esperanza and Shyri. Resistance reactions of F1, BC1, and F2 plants, and individual F2 plant derived F3 families indicated that resistance in each genotype was controlled by the same single dominant gene.     

Spike-based and seed-based selection for preharvest sprouting resistance in wheat

Summary The utility of spike- and seed-based mass selection techniques for improving preharvest sprouting resistance in heterogeneous wheat (Triticum spp.) populations was evaluated. Sorting seed by size improved selection efficiency in some cases, putatively by physiological synchronization. Progeny testing, as well as changes in frequency of red-kernelled types, indicate effectiveness of both spike- and seed-based mass selection for reduced preharvest sprouting. Differential effectiveness of mass selection, in populations segregating for dormancy from different sources, is consistent with previous work on mechanisms of dormancy from these sources. These results are of value to improvement of preharvest sprouting resistance in large, heterogeneous wheat populations.     

Methods for characterization of preharvest sprouting resistance in a wheat breeding program

Preharvest sprouting (PHS) can be a serious problem in western Canada resulting in economically important yield and grade losses for wheat producers. Improved PHS resistance is an important breeding objective and wheat breeders require effective and reliable methods to select for PHS resistance. The objective of this study was to evaluate whether differences exist between field and artificial weathering in the evaluation of PHS resistance in breeding lines and to determine whether sprouting scores are a good estimator of PHS resistance when compared to Hagberg falling numbers. Results suggest that both field and artificial weathering can be effectively used to screen for PHS resistance in wheat breeding lines. Although in this study, field weathering resulted in greater sprouting damage than artificial weathering, the ability to control conditions may make artificial weathering the more reliable test for PHS resistance in a breeding program. Mean sprouting scores greater than 7.5 consistently identified lines with low PHS resistance; hence, sprouting scores can be used to discard the most sprouting susceptible lines. However, falling numbers appear to be more reliable to evaluate the PHS resistance of advanced lines. This revised version was published online in August 2006 with corrections to the Cover Date.     

Intergenotypic competition and border effect in bread wheat and barley

Summary A coordinated series of experiments aimed to estimate border effect and intergenotypic competition was carried out on bread wheat and barley in Italy and Germany. The performance of the varieties grown in alternate rows 14–17 cm apart proved to be strictly dependent on the bordering variety. The varieties grown in adjacent plots consisting of eight rows showed significant border effects: the two external rows yielded up to 40% more than the two innermost, and the two 30 cm end hills facing the alleys yielded 29% more than the central part of the plot. Is has been suggested that in variety trials the first 30–40 cm at both ends of each plot should be removed mechanically. In spite of the border effect detected in the outer rows of the plots, the varieties similarly exploited the extra space available at the borders, and the space of 30 cm left between adjacent plots seems to be sufficient to dilute the intergenotypic competition at plot level to an extent only marginally affecting variety performance.In conclusion, grain yield based on whole plot (8 rows, 6–10 m²) appears to provide a good estimate of true yield, providing that it is based on the actual area of the plot, including the uncropped strip between adjacent plots. By contrast, the reduction of plot size to 1–3 rows is not feasible without introducing considerable bias into the estimation of the yield potential of the varieties.Abbreviations WW winter wheat - WB winter barley - SB spring barley     

QTL mapping of the domestication traits pre-harvest sprouting and dormancy in wheat (<Emphasis Type="BoldItalic">Triticum aestivum L.</Emphasis>)

A set of 75 recombinant inbred lines (RILs) of the ITMI mapping population was grown under field conditions in Gatersleben. The lines were evaluated for the domestication traits pre-harvest sprouting and dormancy (germinability). Main QTLs could be localized for pre-harvest sprouting on chromosome 4AL and dormancy on chromosome 3AL. In addition, 85 Triticum aestivum cv. “Chinese Spring”-Aegilops tauschii introgression lines grown under greenhouse conditions were researched. No QTL could be found for pre-harvest sprouting but a major QTL could be detected for dormancy on chromosome 6DL.     

Exploiting the diversity of <Emphasis Type="Italic">Viviparous-1</Emphasis> gene associated with pre-harvest sprouting tolerance in European wheat varieties

Pre-harvest sprouting (PHS) reduces the quality of wheat (Triticum aestivum L.) and the economic value of the grain. The objective of this study was to evaluate the diversity of the Viviparous-1B (Vp-1B) gene associated with PHS tolerance in a collection of 490 widely grown winter wheat varieties from central and northern Europe. Four alleles of Vp-1B were found in the wheat varieties tested, three of which (Vp-1Ba, Vp-1Bb and Vp-1Bc) had previously been identified in Chinese wheat varieties. The fourth was a new allele which had a 25-bp of deletion in the third intron region compared with the nucleotide sequence of Vp-1Ba, and was designated as Vp-1Bd. The frequencies of different alleles in this set of European wheat germplasm were: Vp-1Ba (54%) > Vp-1Bc (21%) > Vp-1Bd (20%) > Vp-1Ba + c (4%) > Vp-1Bb (1%), with Vp-1Bb being present only in two French varieties, ‘Altria’ and ‘Recital’. In addition, the frequencies of the alleles differed in varieties from different European countries. For example, Vp-1Ba had the highest frequency (76%) in varieties included in the UK National List (NL), but was least frequent in the Recommended List (RL) of Sweden (19%). Similarly, Vp-1Bc was present with the highest frequency (58%) in wheat varieties from Sweden, and the lowest in UK NL varieties (8%) while Vp-1Bd had the highest frequency of 32% in German varieties, and the lowest in Sweden varieties with only 8%. The Vp-1Ba allele was present in over half of the UK wheat varieties tested but the frequency was lower in RL varieties than in NL ones. Furthermore, heterogeneities were found between Vp-1Ba and Vp-1Bc in the varieties from Sweden, Netherlands, Germany and UK.     

Responses of wheat and barley genotypes to toxic concentrations of soil boron

Summary The growth and yield of seven wheat and two barley cultivars or lines, previously found to show different degrees of boron tolerance under field conditions, were compared in a pot experiment at a range of soil boron treatments. Soil treatments ranged up to 150 mg/kg applied B. Extractable B in soils ranged up to 103 mg/kg.At the highest B treatment seedling emergence was delayed, but the percentage emergence was not reduced. The degree of boron toxicity symptom expression varied between the wheat cultivars and lines, with the two most tolerant, Halberd and (Wq^*KP)^*WmH)/6/12, displaying the least symptoms.The concentration of boron applied to the soil which produced a significant depression of growth and yield varied between cultivars. For example, the yield of (Wq^*KP)^*WmH)/6/12 was not affected at the 100 mg/kg applied boron treatment, while the grain yield for (Wl^*MMC)/W1/10 was significantly reduced at the 25 mg/kg treatment.There was a linear increase in boron concentration in tillers at the boot-stage with increasing concentration of boron in the soil. The most boron tolerant genotypes had the lowest tissue boron concentrations in each of the treatments. Halberd and (Wq^*KP)^*WmH)/6/12 had approximately half the boron concentrations of the more sensitive genotypes at the 25 and 50 mg/kg treatments. Differential tolerance of boron within the tissue was also observed. Both Stirling and (Wl^*MMC)/W1/10 had significantly reduced total dry matter and grain yields at the 25 mg/kg treatment, while the concentrations of boron in boot stage tillers at this treatment were 118 and 100 mg/kg, respectively. On the other hand, Halberd and (Wq^*KP)^*WmH)/6/12 had tissue boron concentrations of 144 and 131 mg/kg, respectively, at the 50 mg/kg treatment but yield was unaffected.The relative responses in the pot experiment, for wheat, were in close agreement with field results. Halberd and (Wq^*KP)^*WmH)/6/12 had the highest grain yields, with the lowest concentrations of boron in the grain when grown under high boron conditions in the field. In pots these two genotypes proved to be the most tolerant of boron. For barley the advantage in grain yield in the field, expressed by WI-2584 compared with Stirling, was not repeated in pots. WI-2584 was, however, more tolerant than Stirling on the basis of total dry matter production.The results show that useful variation in boron tolerance exists among wheat, and that breeding should be able to provide cultivars tolerant to high levels of boron.     

Genetic relationships between preharvest sprouting and dormancy in barley

Preharvest sprouting (PHS) and dormancy (DOR) can be problems in barley production and end use quality, especially for barley used for seed and malting. Three crosses previously analyzed for DOR inheritance, were reanalyzed for PHS and DOR inheritance using artificial rain to calculate sprout score (SSc) and measure alpha-amylase activity (AA). Germination percentage of untreated grain for DOR was also measured. The crosses are ‘Steptoe’/’Morex’ (previously published), ‘Harrington’/TR306, and ‘Triumph’/Morex. Among the three crosses, DOR QTLs were located to six and PHS QTLs to five chromosomes, respectively. Chromosome 6H was never implicated. Previously identified DOR QTLs were confirmed in each cross, and most PHS QTLs coincided with DOR QTLs, but not all. Unique PHS QTLs were identified on chromosomes 1H (AA), 2H (SSc, AA), 3H (SSc, AA), and 7H (SSc, AA) and unique DOR QTLs on 1H, 2H, and 7H. Results indicate that PHS susceptibility and DOR are not always represented by opposite alleles at a locus. Some QTL regions for a given trait are conserved across crosses and some are not. Several QTLs are suitable for marker-assisted selection to balance PHS and DOR in breeding new cultivars.     

Manipulation of discriminatory T-DNA delivery by Agrobacterium into cells of immature embryos of barley and wheat

Immature embryos (IEs) of barley (Hordeum vulgare L.) and wheat (Triticum aestivum L.) were co-cultivated with Agrobacterium tumefaciens and the transfer of T-DNA to the cereals was assessed by analysis of the expression of auidA reporter gene. This approach revealed that T-DNA transfer is preferentially targeted to distinct anatomical regions of the cereal embryos that have a low capacity for regenerative growth. This preference may limit the efficiency of generating genetically transformed plants. With the aims of maximizing the frequency of T-DNA delivery and altering the preference of delivery in favour of regeneration-competent areas of IEs, various parameters of the inoculation and co-cultivation procedure were evaluated. It was found that controlling the density of the Agrobacteriuminoculum and/ or the concentration of the Murashige and Skoog (MS) basal salts in the medium produced changes both in the total number and relative distribution of T-DNA transfer events, and these changes may have positive implications for the production of stable transformants. The use of pre-cultured embryos were also important for T-DNA targeting. This revised version was published online in August 2006 with corrections to the Cover Date.     

Identification of bread wheat,durum wheat and barley cultivars adapted to dry areas of Southern Italy

Summary Among the cultivars of bread wheat, durum wheat and barley grown in the South of Italy, genetic variation for adaptation to the high temperature and drought stress conditions typical of the Mediterranean environment has been found.The basic data have been extrapolated from 5 years of Italian national network cultivar trials, where 20–30 cultivars were grown in replicated plot trials in 30–50 locations per year, including some where stress strongly affected grain yield.After careful identification of the most representative years and testing sites it was possible to characterise the cultivars on the basis of the grain yield in stress conditions and the Fischer & Maurer (1978) susceptibility index and to find genotypic differences sufficiently repeatable in years.The cultivars giving the best yield under stress associated with low susceptibility indices were in bread wheat: Etruria, Spada, Pandas, Centauro, Oderzo, Costantino and Gladio, in durum wheat: Aldura, Arcangelo, Adamello, Vespro and Capeiti, in barley: Fleuret, Barberousse, Jaidor, Express, Trebbia, Georgie, Dahlia, Criter and Magie.     

The effect of temperature on seed set and embryo development in reciprocal crosses of wheat and barley

Summary The effect of temperature on seed set and embryo development in reciprocal crosses of barley and wheat was assessed in crosses involving two spring barley varieties (Betzes, Martonvásári 50) and one wheat variety (Chinese Spring). Detached tillers placed in nutrient solution were pollinated in controlled environments at constant day-night temperature regimes (12, 15, 18 and 21° C) with a light intensity of 30,000 lux and a relative humidity of 80%. When barley was used as the female, lower temperatures (12 and 15° C) produced the maximum seed set, whereas for the reciprocal cross, the highest temperature (21° C) produced the best seed set in the Chinese Spring × Betzes combination.Low temperature retarded the embryo development. The highest numbers of hybrid plants were produced at 18° C and 21° C in the barley × wheat cross and in the wheat × barley cross, respectively. Embryos of about 1.5 mm length in the barley × wheat cross, and of about 1.0 mm length in the wheat × barley cross germinated successfully. The smallest embryo giving rise to hybrid plants was 0.57 mm in the barley × wheat cross and 0.51 mm in the wheat × barley cross.     

Worldwide distribution of β-amylase thermostability in barley

The thermostability of β-amylase in 6752 lines of worldwide barley genetic resources were investigated. Most of the lines were classified into high (type A), medium (type B) , and low (type C) thermostability. Subsequently the geographical distribution of these types was clarified. About 90% of the East Asian (Japan, the Korean Peninsula, China) lines were type A. More than 95% of Ethiopian barley was type C. The thermostability types of varieties in the western areas (north Africa, southwest Asia, Turkey, Europe) consisted of types A, B and C. These results suggest that there is a clear geographical differentiation in β-amylase thermostability, especially in East Asia and Ethiopia. The phenotype characteristics of each thermostability type line reflected the geographical differentiation. Besides types A, B and C, we found new thermostability types, including such useful mutants as a β-amylase-less mutant and highly-thermostable mutants than type A in both China and Nepal. This revised version was published online in August 2006 with corrections to the Cover Date.