Worldwide, dryland salinity is a major limitation to crop production. Breeding for salinity tolerance could be an effective way of improving yield and yield stability on saline-sodic soils of dryland agriculture. However, this requires a good understanding of inheritance of this quantitative trait. In the present study, a doubled-haploid bread wheat population (Berkut/Krichauff) was grown in supported hydroponics to identify quantitative trait loci (QTL) associated with salinity tolerance traits commonly reported in the literature (leaf symptoms, tiller number, seedling biomass, chlorophyll content, and shoot Na(+) and K(+) concentrations), understand the relationships amongst these traits, and determine their genetic value for marker-assisted selection. There was considerable segregation within the population for all traits measured. With a genetic map of 527 SSR-, DArT- and gene-based markers, a total of 40 QTL were detected for all seven traits. For the first time in a cereal species, a QTL interval for Na(+) exclusion (wPt-3114-wmc170) was associated with an increase (10%) in seedling biomass. Of the five QTL identified for Na(+) exclusion, two were co-located with seedling biomass (2A and 6A). The 2A QTL appears to coincide with the previously reported Na(+) exclusion locus in durum wheat that hosts one active HKT1;4 (Nax1) and one inactive HKT1;4 gene. Using these sequences as template for primer design enabled mapping of at least three HKT1;4 genes onto chromosome 2AL in bread wheat, suggesting that bread wheat carries more HKT1;4 gene family members than durum wheat. However, the combined effects of all Na(+) exclusion loci only accounted for 18% of the variation in seedling biomass under salinity stress indicating that there were other mechanisms of salinity tolerance operative at the seedling stage in this population. Na(+) and K(+) accumulation appear under separate genetic control. The molecular markers wmc170 (2A) and cfd080 (6A) are expected to facilitate breeding for salinity tolerance in bread wheat, the latter being associated with seedling vigour.
Soil salinity and sodicity are major constraints to global cereal production, but breeding for tolerance has been slow. Narrow gene pools, over-emphasis on the sodium (Na+) exclusion mechanism, little attention to osmotic stress/tissue tolerance mechanism(s) in which accumulation of inorganic ions such as Na+ is implicated, and lack of a suitable screening method have impaired progress. The aims of this study were to discover novel genes for Na+ accumulation using genome-wide association studies, compare growth responses to salinity and sodicity in low-Na+ bread Westonia with Nax1 and Nax2 genes and high-Na+ bread wheat Baart-46, and evaluate growth responses to salinity and sodicity in bread wheats with varying leaf Na+ concentrations. The novel high-Na+ bread wheat germplasm, MW#293, had higher grain yield under salinity and sodicity, in absolute and relative terms, than the other bread wheat entries tested. Genes associated with high Na+ accumulation in bread wheat were identified, which may be involved in tissue tolerance/osmotic adjustment. As most modern bread wheats are efficient at excluding Na+, further reduction in plant Na+ is unlikely to provide agronomic benefit. The salinity and sodicity tolerant germplasm MW#293 provides an opportunity for the development of future salinity/sodicity tolerant bread wheat.
Three cDNAs encoding the antifungal protein Ag-AFP from the fungus Aspergillus giganteus, a barley class II chitinase and a barley type I RIP, all regulated by the constitutive Ubiquitin1 promoter from maize, were expressed in transgenic wheat. In 17 wheat lines, stable integration and inheritance of one of the three transgenes has been demonstrated over four generations. The formation of powdery mildew (Erysiphe graminis f. sp. tritici) or leaf rust (Puccinia recondita f. sp. tritici) colonies was significantly reduced on leaves from afp or chitinase II- but not from rip I-expressing wheat lines compared with non-transgenic controls. The increased resistance of afp and chitinase II lines was dependent on the dose of fungal spores used for inoculation. Heterologous expression of the fungal afp gene and the barley chitinase II gene in wheat demonstrated that colony formation and, thereby, spreading of two important biotrophic fungal diseases is inhibited approximately 40 to 50% at an inoculum density of 80 to 100 spores per cm2.
SummaryThe separation of toxic effects of sodium (Na + ) and chloride (Cl À ) by the current methods of mixed salts and subsequent determination of their relevance to breeding has been problematic. We report a novel method (Na + humate) to study the ionic effects of Na + toxicity without interference from Cl À , and ionic and osmotic effects when combined with salinity (NaCl).Three cereal species (Hordeum vulgare, Triticum aestivum and Triticum turgidum ssp. durum with and without the Na + exclusion gene Nax2) differing in Na + exclusion were grown in a potting mix under sodicity (Na + humate) and salinity (NaCl), and water use, leaf nutrient profiles and yield were determined. Under sodicity, Na + -excluding bread wheat and durum wheat with the Nax2 gene had higher yield than Na + -accumulating barley and durum wheat without the Nax2 gene. However, under salinity, despite a 100-fold difference in leaf Na + , all species yielded similarly, indicating that osmotic stress negated the benefits of Na + exclusion. In conclusion, Na + exclusion can be an effective mechanism for sodicity tolerance, while osmoregulation and tissue tolerance to Na + and/or Cl À should be the main foci for further improvement of salinity tolerance in cereals. This represents a paradigm shift for breeding cereals with salinity tolerance.
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