Wheat yields globally will depend increasingly on good management to conserve rainfall and new varieties that use water efficiently for grain production. Here we propose an approach for developing new varieties to make better use of deep stored water. We focus on water-limited wheat production in the summer-dominant rainfall regions of India and Australia, but the approach is generally applicable to other environments and root-based constraints. Use of stored deep water is valuable because it is more predictable than variable in-season rainfall and can be measured prior to sowing. Further, this moisture is converted into grain with twice the efficiently of in-season rainfall since it is taken up later in crop growth during the grain-filling period when the roots reach deeper layers. We propose that wheat varieties with a deeper root system, a redistribution of branch root density from the surface to depth, and with greater radial hydraulic conductivity at depth would have higher yields in rainfed systems where crops rely on deep water for grain fill. Developing selection systems for mature root system traits is challenging as there are limited high-throughput phenotyping methods for roots in the field, and there is a risk that traits selected in the lab on young plants will not translate into mature root system traits in the field. We give an example of a breeding programme that combines laboratory and field phenotyping with proof of concept evaluation of the trait at the beginning of the selection programme. This would greatly enhance confidence in a high-throughput laboratory or field screen, and avoid investment in screens without yield value. This approach requires careful selection of field sites and years that allow expression of deep roots and increased yield. It also requires careful selection and crossing of germplasm to allow comparison of root expression among genotypes that are similar for other traits, especially flowering time and disease and toxicity resistances. Such a programme with field and laboratory evaluation at the outset will speed up delivery of varieties with improved root systems for higher yield.

The grain yield of cereals has almost doubled this century as a result of genetic manipulation by plant breeding. Surprisingly, there has been no change in the rate of photosynthesis per unit leaf area to accompany these increases. However, total photosynthesis has increased as a result of an increase in leaf area, daily duration of photosynthesis or leaf area duration. There remain substantial opportunities to continue to improve total photosynthesis and crop yield genetically using conventional breeding practices. Selectable traits are discussed here in the context of increasing total above-ground biomass under favourable conditions. Opportunities exist to alter crop duration and the timing of crop development to match it better to radiation, temperature and vapour pressure during crop growth, and to increase the rate of development of early leaf area to achieve rapid canopy closure. The importance of these traits will depend on the environment in which the crop is grown. Increases in crop photosynthesis through breeding are also likely to come via indirect means. Selection for a high and sustained stomatal conductance during the period of stem elongation is one way. Increasing assimilate allocation to the reproductive primordia so as to establish a large potential sink should also indirectly increase total crop photosynthesis. Evidence in the major grain crops suggests that by anthesis the capacity for photosynthesis is high and that photosynthesis is not limiting during grain filling. To use this surplus capacity it is suggested that carbon and nitrogen partitioning to the reproductive meristem be increased so as to establish a high potential grain number and the potential for a large grain size. It is then expected that additional photosynthesis will follow, either by a longer daily duration of photosynthesis or by an extended leaf area duration.

Reproductive stage water stress leads to spikelet sterility in wheat. Whereas drought stress at anthesis affects mainly grain size, stress at the young microspore stage of pollen development is characterized by abortion of pollen development and reduction in grain number. We identified genetic variability for drought tolerance at the reproductive stage. Drought-tolerant wheat germplasm is able to maintain carbohydrate accumulation in the reproductive organs throughout the stress treatment. Starch depletion in the ovary of drought-sensitive wheat is reversible upon re-watering and cross-pollination experiments indicate that the ovary is more resilient than the anther. The effect on anthers and pollen fertility is irreversible, suggesting that pollen sterility is the main cause of grain loss during drought conditions in wheat. The difference in storage carbohydrate accumulation in drought-sensitive and droughttolerant wheat is correlated with differences in sugar profiles, cell wall invertase gene expression and expression of fructan biosynthesis genes in anther and ovary (sucrose : sucrose 1-fructosyl-transferase, 1-SST; sucrose : fructan 6-fructosyl-transferase, 6-SFT). Our results indicate that the ability to control and maintain sink strength and carbohydrate supply to anthers may be the key to maintaining pollen fertility and grain number in wheat and this mechanism may also provide protection against other abiotic stresses.