Effects of drought hardening on the carbohydrate dynamics of

Introduction: As precipitation patterns are predicted to become increasingly erratic, the functional maintenance of warm-temperate forests constitutes a key challenge for forest managers. In this study, 2-year-old Methods: Seedlings were trained under different soil water conditions for 2 months: drought (D), well-watered (W), 1-month drought and then 1-month well-watered (D-W), and 1-month well-watered and then 1-month drought (W-D). The functional traits involved in water- and carbon-use strategies were explored at the end of the hardening period. Compared with seedlings in group W, seedlings in groups D, D-W, and W-D had increased potential for carbon uptake (i.e., light saturated point, maximum ribulose-1,5-bisphosphate (RuBP) saturated rate, and electron transport rate) and water uptake (i.e., fine root-to-coarse root ratio) and downregulated growth and mitochondrial respiration to decrease carbon consumption. After water fluctuation hardening, we performed a successional dry-down experiment for 1 month to detect carbohydrate dynamics and explore the acclimation caused by prior hardening. Results and discussion: Our results revealed that there were more soluble sugars allocated in the leaves and more starch allocated in the stems and roots of seedlings hardened in the D, W-D, and D-W treatments than that of seedlings hardened in the W treatment. No significant changes in total non-structural carbohydrates were found. In addition, we found near-zero (seedlings trained by D and D-W treatments) or negative (seedlings trained by W-D treatment) growth of structural biomass at the end of the dry-down experiment, which was significantly lower than that of W-hardened seedlings. This suggests that there was a shift in allocation patterns between carbon storage and growth under recurrent soil drought, which can be strengthened by drought memory. We conclude that