Die-hard seedlings. A global meta-analysis on the factors determining the effectiveness of drought hardening on growth and survival of forest plantations

Drought hardening is a nursery technique aimed to enhance early forest plantation establishment under dry conditions, which is a main limiting factors for plantation success. However, the quantitative effectiveness of drought hardening remains unclear. We conducted a meta-analysis to evaluate the influence of different factors in the effectiveness of drought hardening on seedling post-planting survival and growth. Overall, drought hardening did not significantly affect survival or growth, as several factors induced great heterogeneity, but analyses of those factors explained its effectiveness, especially on survival. A longer time between hardening and transplanting strongly reduced survival. Indoor-grown seedlings did not benefit more from hardening than outdoor-grown seedlings. Evaluations of drought hardening effectiveness in pots showed positive effects on survival but negative effects on growth, while no effects were found in large bed experiments. In field experiments, hardening significantly increased survival and growth with site aridity. Survival benefits were independent of species drought tolerance, measured by osmotic potential at the turgor loss point (π_tlp), in moderate to high aridity sites. However, in low aridity sites, hardening increased survival in drought-tolerant species but decreased it in drought-intolerant species. Field results showed that hardening benefited shrubs more than trees in angiosperms. In conclusion, drought hardening at the end of nursery cultivation tend to increase post-planting seedling performance particularly in scenarios limiting post-planting root growth such as in arid climates and pot experiments. Our findings highlight the importance of future research on modelling the interaction between these technical features and species water use strategies..