Abstract

Stored non-structural carbohydrates (NSC) have been proposed as a key determinant of drought resistance in plants. However, the evidence for this role is controversial, as it comes mostly from observational, short-term studies. Here, we take advantage of a long-term experimental throughfall reduction to elucidate the response of NSC to increased drought 14 years after the beginning of the treatment in three Mediterranean resprouter trees (Quercus ilex L., Arbutus unedo L. and Phillyrea latifolia L.). In addition, we selected 20 Q. ilex individuals outside the experimental plots to directly assess the relationship between defoliation and NSC at the individual level. We measured the seasonal course of NSC concentrations in leaves, branches and lignotuber in late winter, late spring, summer, and autumn 2012. Total concentrations of NSC were highest in the lignotuber for all species. In the long-term drought experiment we found significant depletion in concentrations of total NSC in treatment plots only in the lignotuber of A. unedo. At the same time, A. unedo was the only species showing a significant reduction in BAI under the drought treatment during the 14 years of the experiment. By contrast, Q. ilex just reduced stem growth only during the first 4 years of treatment and P. latifolia remained unaffected over the whole study period. However, we found a clear association between the concentrations of NSC and defoliation in Q. ilex individuals sampled outside the experimental plots, with lower total concentrations of NSC and lower proportion of starch in defoliated individuals. Taken together, our results suggest that stabilizing processes, probably at the stand level, may have been operating in the long-term to mitigate any impact of drought on NSC levels, and highlight the necessity to incorporate long-term experimental studies of plant responses to drought.

Highlights

  • Climate-related tree mortality has been observed worldwide in all major forest biomes (Allen et al, 2010)

  • DISTRIBUTION AND TEMPORAL VARIATION OF non-structural carbohydrates (NSC) CONCENTRATIONS ACROSS SPECIES The NSC concentrations measured in the present study were comparable to the values previously reported for the same species [e.g., in Q. ilex lignotuber by López et al (2009) and Galiano et al (2012) and in A. unedo leaves by Meletiou-Christou et al (1994)]

  • The concentrations of NSC and soluble sugars tended to be highest in A. unedo and lowest in P. latifolia

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Summary

Introduction

Climate-related tree mortality has been observed worldwide in all major forest biomes (Allen et al, 2010). Because more frequent and intense droughts are predicted, in the Mediterranean basin (IPCC, 2007), different sensitivity of species to drought may cause widespread changes in species distribution and community composition (Engelbrecht et al, 2007; Choat et al, 2012; but see Lloret et al, 2012) Such vegetation shifts may have important implications for ecosystem function, land–atmosphere interactions and ecosystem services to humans in general (Dale et al, 2000; Bonan, 2008; Anderegg et al, 2012a). The carbon starvation hypothesis predicts that stomatal closure to prevent desiccation causes photosynthetic carbon uptake to diminish to near zero, and that continued demand for carbohydrates to maintain metabolism will deplete carbohydrate reserves, leading eventually to starvation and death (McDowell et al, 2008; Adams et al, 2009) Biotic agents, such as insects and pathogens, can amplify or be amplified by both carbon starvation and hydraulic failure (Shaw et al, 2005; Fettig et al, 2007). Recent research has emphasized the links between the www.frontiersin.org

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