Inspection of gas bubbles in frozen Betula pendula xylem with micro-CT: Conduit size, water status and bark permeability affect bubble characteristics

Abstract Bubbles of gas trapped in the xylem during freezing are a major cause of damage for trees growing at high altitudes or latitudes, as the bubbles may cause embolism during thawing. Yet the factors controlling bubble formation upon freeze-thaw cycles remain poorly understood. Especially the size of the bubbles formed in the ice is crucial for winter embolism formation. We used high-resolution X-ray microtomography combined with freezing experiments to investigate the size and shape of 68 343 gas bubbles in frozen conduits in branches of Betula pendula. We also studied how conduit size, tree water status (-0.2 MPa vs -0.6 MPa) and bark permeability to gases (decreased by Vaseline-coating) affect the gas bubbles characteristics. High-resolution X-ray images allowed us to detect gas bubbles down to 1.0 ?m in diameter and revealed that not only small spherical gas bubbles but gaseous volumes of various shapes and sizes were found from the frozen xylem indicating that gas bubbles may have started to grow already during the freezing propagation. Most of the gas bubbles were found in fibers, but the rare gas bubbles found in the vessels were larger than those in the fibers. Bubble volume increased with conduit volume in both fibers and vessels, but conduit size alone could not explain gas bubble volume. Low water potential and restriction of gas escape from the branch seem to cause more, larger, and less spherical bubbles and thus increase the risk of embolism formation. These findings open new research avenues for further studies of winter embolism formation. This article is protected by copyright. All rights reserved.