Water taken up through the bark is detected in the transpiration stream in intact upper-canopy branches

Abstract

Alternative water uptake pathways through leaves and bark complement water supply with interception, fog or dew. Bark water-uptake contributes to embolism-repair, as demonstrated in cut branches. We tested whether bark water-uptake could also contribute to supplement xylem-water for transpiration. We applied bandages injected with ² H-enriched water on intact upper-canopy branches of Pinus sylvestris and Fagus sylvatica in a boreal and in a temperate forest, in summer and winter, and monitored transpiration and online isotopic composition (δ² H and δ¹⁸ O) of water vapour, before sampling for analyses of δ² H and δ¹⁸ O in tissue waters. Xylem, bark and leaf waters from segments downstream from the bandages were ² H-enriched whereas δ¹⁸ O was similar to controls. Transpiration was positively correlated with ² H-enrichment. Isotopic compositions of transpiration and xylem water allowed us to calculate isotopic exchange through the bark via vapour exchange, which was negligible in comparison to estimated bark water-uptake, suggesting that water-uptake occurred via liquid phase. Results were consistent across species, forests and seasons, indicating that bark water-uptake may be more ubiquitous than previously considered. We suggest that water taken up through the bark could be incorporated into the transpiration stream, which could imply that sap-flow measurements underestimate transpiration when bark is wet.

Keywords: European beech; Scots pine; bark; deuterium; drought; frost; hydrogen; oxygen; water stable isotopes; xylem.