Description: Plants exhibit a variety of drought responses involving multiple interacting traits and processes, which makes predictions of drought survival challenging. Careful evaluation of responses within species, where individuals share broadly similar drought resistance strategies, can provide insight into the relative importance of different traits and processes. We subjected Pinus sylvestris L. saplings to extreme drought (no watering) leading to death in a greenhouse to (i) determine the relative effect of predisposing factors and responses to drought on survival time, (ii) identify and rank the importance of key predictors of time to death and (iii) compare individual characteristics of dead and surviving trees sampled concurrently. Time until death varied over 3 months among individual trees (from 29 to 147 days). Survival time was best predicted (higher explained variance and impact on the median survival time) by variables related to carbon uptake and carbon/water economy before and during drought. Trees with higher concentrations of monosaccharides before the beginning of the drought treatment and with higher assimilation rates prior to and during the treatment survived longer (median survival time increased 25–70 days), even at the expense of higher water loss. Dead trees exhibited less than half the amount of nonstructural carbohydrates (NSCs) in branches, stem and relative to surviving trees sampled concurrently. Overall, our results indicate that the maintenance of carbon assimilation to prevent acute depletion of NSC content above some critical level appears to be the main factor explaining survival time of P. sylvestris trees under extreme drought.

Description: Drought-related tree die-off episodes have been observed in all vegetated continents. Despite much research effort, however, the multiple interactions between carbon starvation, hydraulic failure and biotic agents in driving tree mortality under field conditions are still not well understood. We analysed the seasonal variability of non-structural carbohydrates (NSCs) in four organs (leaves, branches, trunk and roots), the vulnerability to embolism in roots and branches, native embolism (percentage loss of hydraulic conductivity (PLC)) in branches and the presence of root rot pathogens in defoliated and non-defoliated individuals in a declining Scots pine ( Pinus sylvestris L.) population in the NE Iberian Peninsula in 2012, which included a particularly dry and warm summer. No differences were observed between defoliated and non-defoliated pines in hydraulic parameters, except for a higher vulnerability to embolism at pressures below –2 MPa in roots of defoliated pines. No differences were found between defoliation classes in branch PLC. Total NSC (TNSC, soluble sugars plus starch) values decreased during drought, particularly in leaves. Defoliation reduced TNSC levels across tree organs, especially just before (June) and during (August) drought. Root rot infection by the fungal pathogen Onnia P. Karst spp. was detected but it did not appear to be associated to tree defoliation. However, Onnia infection was associated with reduced leaf-specific hydraulic conductivity and sapwood depth, and thus contributed to hydraulic impairment, especially in defoliated pines. Infection was also associated with virtually depleted root starch reserves during and after drought in defoliated pines. Moreover, defoliated and infected trees tended to show lower basal area increment. Overall, our results show the intertwined nature of physiological mechanisms leading to drought-induced mortality and the inherent difficulty of isolating their contribution under field conditions.

Description: Non-structural carbohydrates (NSC) in plant tissue are frequently quantified to make inferences about plant responses to environmental conditions. Laboratories publishing estimates of NSC of woody plants use many different methods to evaluate NSC. We asked whether NSC estimates in the recent literature could be quantitatively compared among studies. We also asked whether any differences among laboratories were related to the extraction and quantification methods used to determine starch and sugar concentrations. These questions were addressed by sending sub-samples collected from five woody plant tissues, which varied in NSC content and chemical composition, to 29 laboratories. Each laboratory analyzed the samples with their laboratory-specific protocols, based on recent publications, to determine concentrations of soluble sugars, starch and their sum, total NSC. Laboratory estimates differed substantially for all samples. For example, estimates for Eucalyptus globulus leaves (EGL) varied from 23 to 116 (mean = 56) mg g –1 for soluble sugars, 6–533 (mean = 94) mg g –1 for starch and 53–649 (mean = 153) mg g –1 for total NSC. Mixed model analysis of variance showed that much of the variability among laboratories was unrelated to the categories we used for extraction and quantification methods (method category R 2 = 0.05–0.12 for soluble sugars, 0.10–0.33 for starch and 0.01–0.09 for total NSC). For EGL, the difference between the highest and lowest least squares means for categories in the mixed model analysis was 33 mg g –1 for total NSC, compared with the range of laboratory estimates of 596 mg g –1 . Laboratories were reasonably consistent in their ranks of estimates among tissues for starch ( r = 0.41–0.91), but less so for total NSC ( r = 0.45–0.84) and soluble sugars ( r = 0.11–0.83). Our results show that NSC estimates for woody plant tissues cannot be compared among laboratories. The relative changes in NSC between treatments measured within a laboratory may be comparable within and between laboratories, especially for starch. To obtain comparable NSC estimates, we suggest that users can either adopt the reference method given in this publication, or report estimates for a portion of samples using the reference method, and report estimates for a standard reference material. Researchers interested in NSC estimates should work to identify and adopt standard methods.