Drought and heat stress cause losses in wheat productivity in major growing regions worldwide, and both the occurrence and the severity of these events are likely to increase with global climate change. Water deficits and high temperatures frequently occur simultaneously at sensitive growth stages, reducing wheat yields by reducing grain number or weight. Although genetic variation and underlying quantitative trait loci for either individual stress are known, the combination of the two stresses has rarely been studied. Complex and often antagonistic physiology means that genetic loci underlying tolerance to the combined stress are likely to differ from those for drought or heat stress tolerance alone. Here, we review what is known of the physiological traits and genetic control of drought and heat tolerance in wheat and discuss potential physiological traits to study for combined tolerance. We further place this knowledge in the context of breeding for new, more tolerant varieties and discuss opportunities and constraints. We conclude that a fine control of water relations across the growing cycle will be beneficial for combined tolerance and might be achieved through fine management of spatial and temporal gas exchange.

Background: Improving abiotic stress tolerance in wheat requires large scale screening of yield components such as seed weight, seed number and single seed weight, all of which is very laborious, and a detailed analysis of seed morphology is time-consuming and visually often impossible. Computed tomography offers the opportunity for much faster and more accurate assessment of yield components. Results: An X-ray computed tomographic analysis was carried out on 203 very diverse wheat accessions which have been exposed to either drought or combined drought and heat stress. Results demonstrated that our computed tomography pipeline was capable of evaluating grain set with an accuracy of 95-99%. Most accessions exposed to combined drought and heat stress developed smaller, shrivelled seeds with an increased seed surface. As expected, seed weight and seed number per ear as well as single seed size were significantly reduced under combined drought and heat compared to drought alone. Seed weight along the ear was significantly reduced at the top and bottom of the wheat spike. Conclusions: We were able to establish a pipeline with a higher throughput with scanning times of 7 min per ear and accuracy than previous pipelines predicting a set of agronomical important seed traits and to visualize even more complex traits such as seed deformations. The pipeline presented here could be scaled up to use for high throughput, high resolution phenotyping of tens of thousands of heads, greatly accelerating breeding efforts to improve abiotic stress tolerance.

Drought and heat waves commonly co-occur in many wheat-growing regions causing significant crop losses. The identification of stress associated quantitative trait loci, particularly those for yield, is problematic due to their association with plant phenology and the high genetic × environment interaction. Here we studied a panel of 315 diverse, spring type accessions of bread wheat (Triticum aestivum) in pots in a semi-controlled environment under combined drought and heat stress over 2 years. Importantly, we treated individual plants according to their flowering time. We found 134 out of the 145 identified loci for grain weight that were not associated with either plant phenology or plant height. The majority of loci uncovered here were novel, with favorable alleles widespread in Asian and African landraces providing opportunities for their incorporation into modern varieties through breeding. Using residual heterozygosity in lines from a nested association mapping population, we were able to rapidly develop near-isogenic lines for important target loci. One target locus on chromosome 6A contributed to higher grain weight, harvest index, thousand kernel weight, and grain number under drought and heat stress in field conditions consistent with allelic effects demonstrated in the genome-wide association study.

Drought and heat stress constrain wheat (Triticum aestivum L.) yields globally. To identify putative mechanisms and candidate genes associated with combined drought and heat stress tolerance, we developed bread wheat near-isogenic lines (NILs) targeting a quantitative trait locus (QTL) on chromosome 6B which was previously associated with combined drought and heat stress tolerance in a diverse panel of wheats. Genotyping-by-sequencing was used to identify additional regions that segregated in allelic pairs between the recurrent and the introduced exotic parent, genome-wide. NILs were phenotyped in a gravimetric platform with precision irrigation and exposed to either drought or to combined drought and heat stress from three days after anthesis. An increase in grain weight in NILs carrying the exotic allele at 6B locus was associated with thicker, greener leaves, higher photosynthetic capacity and increased water use index after re-watering. RNA sequencing of developing grains at early and later stages of treatment revealed 75 genes that were differentially expressed between NILs across both treatments and timepoints. Differentially expressed genes coincided with the targeted QTL on chromosome 6B and regions of genetic segregation on chromosomes 1B and 7A. Pathway enrichment analysis showed the involvement of these genes in cell and gene regulation, metabolism of amino acids and transport of carbohydrates. The majority of these genes have not been characterized previously under drought or heat stress and they might serve as candidate genes for improved abiotic stress tolerance.

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