Tolerance of Combined Drought and Heat Stress Is Associated With Transpiration Maintenance and Water Soluble Carbohydrates in Wheat Grains

Wheat ( Triticum aestivum L.) production is increasingly challenged by simultaneous drought and heatwaves. We assessed the effect of both stresses combined on whole plant water use and carbohydrate partitioning in eight bread wheat genotypes that showed contrasting tolerance. Plant water use was monitored throughout growth, and water-soluble carbohydrates (WSC) and starch were measured following a 3-day heat treatment during drought. Final grain yield was increasingly associated with aboveground biomass and total water use with increasing stress intensity. Combined drought and heat stress immediately reduced daily water use in some genotypes and altered transpiration response to vapor pressure deficit during grain filling, compared to drought only. In grains, glucose and fructose concentrations measured 12 days after anthesis explained 43 and 40% of variation in final grain weight in the main spike, respectively. Starch concentrations in grains offset the reduction in WSC following drought or combined drought and heat stress in some genotypes, while in other genotypes both stresses altered the balance between WSC and starch concentrations. WSC were predominantly allocated to the spike in modern Australian varieties (28–50% of total WSC in the main stem), whereas the stem contained most WSC in older genotypes (67–87%). Drought and combined drought and heat stress increased WSC partitioning to the spike in older genotypes but not in the modern varieties. Ability to maintain transpiration, especially following combined drought and heat stress, appears essential for maintaining wheat productivity.