The synergistic effect of hydraulic and thermal impairments accounts for the severe crown damage in Fraxinus mandshurica seedlings following the combined drought-heatwave stress

Drought combined with extreme heatwaves has been increasingly identified as the important trigger of worldwide tree mortality in the context of climate change; nonetheless, our understanding of the potential hydraulic and thermal im-pairments of hot droughts to trees and the subsequent post-recovery process remains limited. To investigate the re-sponse of tree water and carbon relations to drought, heatwave, and combined drought-heatwave stresses, three-year-old potted seedlings of Fraxinus mandshurica Rupr., a dominant tree species in temperate forests of northeast China, were grown under well-watered and drought-stressed conditions and exposed to a rapid, acute heatwave treat-ment. During the heatwave treatment with a maximum temperature exceeding 40 degrees C for two days, the leaf temperature of drought-stressed seedlings was, on average, 5 degrees C higher than that of well-watered counterparts due to less effective evaporative cooling, indicating that soil water availability influenced leaf thermoregulatory capacity during hot ex-tremes. Consistently, more pronounced crown damage, as shown by 13 % irreversible leaf scorch, was found in seed-lings under the drought-heatwave treatment relative to sole heatwave treatment, alongside the more severe stem xylem embolism and leaf electrolyte leakage. While the heatwave treatment accelerated the depletion of non-structural carbohydrates in drought-stressed seedlings, the increase of branch soluble sugar concentration in response to heatwave might be related to the requirement for maintaining hydraulic functioning via osmoregulation under high dehydration risk. The coordination between leaf stomatal conductance and total non-structural carbohydrate content during the post-heatwave recovery phase implied that plant-water relations and carbon physiology were closely coupled in coping with hot droughts. This study highlights that, under scenarios of aggravating drought co-occurring with heatwaves, tree seedlings could face a high risk of crown decline in relation to the synergistically increased hydraulic and thermal impairments.