In Situ Visualization of the Dynamics in Xylem Embolism Formation and Removal in the Absence of Root Pressure: A Study on Excised Grapevine Stems
Knipfer T; Cuneo I.F; Brodersen C.R; McElrone A.J;
Gas embolisms formed during drought can disrupt long-distance water transport through plant xylem vessels but some species have the ability to remove these blockages. Despite evidence suggesting that embolism removal is linked to the presence of vessel-associated parenchyma the underlying mechanism remains controversial and is thought to involve positive pressure generated by roots. Here we used in situ x-ray microtomography on excised grapevine stems to determine if embolism removal is possible without root pressure and if the embolism formation/removal affects vessel functional status after sample excision. Our data show that embolism removal in excised stems was driven by water droplet growth and was qualitatively identical to refilling in intact plants. When stem segments were rehydrated with H2O after excision vessel refilling occurred rapidly (<1 h). The refilling process was substantially slower when polyethylene glycol was added to the H2O source thereby providing new support for an osmotically driven refilling mechanism. In contrast segments not supplied with H2O showed no refilling and increased embolism formation. Dynamic changes in liquid/wall contact angles indicated that the processes of embolism removal (i.e. vessel refilling) by water influx and embolism formation by water efflux were directly linked to the activity of vessel-associated living tissue. Overall our results emphasize that root pressure is not required as a driving force for vessel refilling and care should be taken when performing hydraulics measurements on excised plant organs containing living vessel-associated tissue because the vessel behavior may not be static.