Sugar infusion into trees: A novel method to study tree carbon relations and its regulations.

Many carbon-related physiological questions in plants such as carbon (C) limitation or starvation have not yet been resolved thoroughly due to the lack of suitable experimental methodology. As a first step towards resolving these problems, we conducted infusion experiments with bonsai trees (Ficus microcarpa) and young maple trees (Acer pseudoplatanus) in greenhouse, and with adult Scots pine trees (Pinus sylvestris) in the field, that were "fed" with 13C-labelled glucose either through the phloem or the xylem. We then traced the 13C-signal in plant organic matter and respiration to test whether trees can take up and metabolize exogenous sugars infused. Ten weeks after infusion started, xylem but not phloem infusion significantly increased the δ13C values in both aboveground and belowground tissues of the bonsai trees in the greenhouse, whereas xylem infusion significantly increased xylem δ13C values and phloem infusion significantly increased phloem δ13C values of the adult pines in the field experiment, compared to the corresponding controls. The respiration measurement experiment with young maple trees showed significantly increased δ13C-values in shoot respired CO2 at the time of four weeks after xylem infusion started. Our results clearly indicate that trees do translocate and metabolize exogenous sugars infused, and because the phloem layer is too thin, and thus xylem infusion can be better operated than phloem infusion. This tree infusion method developed here opens up new avenues and has great potential to be used for research on the whole plant C balance and its regulation in response to environmental factors and extreme stress conditions.

From the comfort zone to crown dieback: Sequence of physiological stress thresholds in mature European beech trees across progressive drought.

Drought responses of mature trees are still poorly understood making it difficult to predict species distributions under a warmer climate. Using mature European beech (Fagus sylvatica L.), a widespread and economically important tree species in Europe, we aimed at developing an empirical stress-level scheme to describe its physiological response to drought. We analysed effects of decreasing soil and leaf water potential on soil water uptake, stem radius, native embolism, early defoliation and crown dieback with comprehensive measurements from overall nine hydrologically distinct beech stands across Switzerland, including records from the exceptional 2018 drought and the 2019/2020 post-drought period. Based on the observed responses to decreasing water potential we derived the following five stress levels: I (predawn leaf water potential >-0.4 MPa): no detectable hydraulic limitations; II (-0.4 to -1.3): persistent stem shrinkage begins and growth ceases; III (-1.3 to -2.1): onset of native embolism and defoliation; IV (-2.1 to -2.8): onset of crown dieback; V (<-2.8): transpiration ceases and crown dieback is >20%. Our scheme provides, for the first time, quantitative thresholds regarding the physiological downregulation of mature European beech trees under drought and therefore synthesises relevant and fundamental information for process-based species distribution models. Moreover, our study revealed that European beech is drought vulnerable, because it still transpires considerably at high levels of embolism and because defoliation occurs rather as a result of embolism than preventing embolism. During the 2018 drought, an exposure to the stress levels III-V of only one month was long enough to trigger substantial crown dieback in beech trees on shallow soils. On deep soils with a high water holding capacity, in contrast, water reserves in deep soil layers prevented drought stress in beech trees. This emphasises the importance to include local data on soil water availability when predicting the future distribution of European beech.