Combined effects of defoliation and water stress on pine growth and non-structural carbohydrates.

Climate change is expected to increase both pest insect damage and the occurrence of severe drought. There is therefore a need to better understand the combined effects of biotic and abiotic damage on tree growth in order to predict the multi-factorial effect of climate change on forest ecosystem productivity. Indeed, the effect of stress interactions on tree growth is an increasingly important topic that greatly lacks experiments and data, and it is unlikely that the impact of combined stresses can be extrapolated from the outcomes of studies that focused on a single stress. We developed an original manipulative study under real field conditions where we applied artificial defoliation and induced water stress on 10-year-old (∼10 m high) maritime pine trees (Pinus pinaster Ait.). Tree response to combined stresses was quantitatively assessed following tree secondary growth and carbohydrate pools. Such a design allowed us to address the crucial question of combined stresses on trees under stand conditions, sharing soil supplies with neighboring trees. Our initial hypotheses were that (i) moderate defoliation can limit the impact of water stress on tree growth through reduced transpiration demand by a tree canopy partly defoliated and that (ii) defoliation results in reduced non-structural carbohydrate (NSC) pools, affecting tree tolerance to drought. Our results showed additive effects of defoliation and water stress on tree growth and contradict our initial hypothesis. Indeed, under stand conditions, we found that partial defoliation does not limit the impact of water stress through reduced transpiration. Our study also highlighted that, even if NSC in all organs were affected by defoliation, tree response to water stress was not triggered. We found that stem NSC were maintained or increased during the entire growing season, supporting literature-based hypotheses such as an active maintenance of the hydraulic system or another limiting resource for tree growth under defoliation. We also observed a significant decrease in root carbohydrates, which suggests a shift in the root carbon balance under defoliation. The decrease in carbohydrate supply under defoliation may not counterbalance the carbon use for mineral and water uptakes or a translocation to other tissues.