Evidence of a seasonal trade-off between growth and starch storage in declining beeches: assessment through stem radial increment, non-structural carbohydrates and intra-ring δ13C

Forest decline is reported in recent decades all over the world. However, developing a clear vision of the associated tree dysfunctioning is still a challenge for plant physiologists. In this study, our aim was to examine the seasonal carbon adjustments of beech trees in the case of a long-term drought-induced decline. We compared healthy and declining trees in terms of stem radial growth, phloem sugar content and δ13C, together with xylem carbohydrates and intra-ring δ13C patterns. The radial growth of declining trees was clearly reduced by lower growth rates and shorter growing season length (44 days compared with healthy trees). The soluble sugar content was higher in the xylem of declining trees compared with the healthy ones, but similar in the phloem except at the end of their growth. Declining trees increased their levels of xylem starch content from budburst until the date of maximal growth rate. These reserve dynamics revealed an early trade-off between radial growth and starch storage that might be the result of an active or passive process. For declining trees, the slight decrease of intra-ring cellulose δ13C pattern during the early growing season was attributed to the synthesis of 13C enriched starch. For healthy trees, δ13C patterns were characterized by a progressive 13C increase along the ring, attributed to increased water-use efficiency (WUE) in response to decreased water availability. Individual variations of the crown area were negatively correlated to the intra-ring δ13C amplitude, which was ascribed to variations in canopy WUE and resource competition for healthy trees and partly to variations in the amount of reserves accumulated during spring for declining ones. Our study highlights the carbon physiological adjustment of declining trees towards reducing spring growth while storing starch, which can be reflected in the individual intra-ring cellulose δ13C patterns.