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Title: Dynamic surface tension of xylem sap lipids
Abstract The surface tension of xylem sap has been traditionally assumed to be close to that of the pure water because decreasing surface tension is thought to increase vulnerability to air seeding and embolism. However, xylem sap contains insoluble lipid-based surfactants, which also coat vessel and pit membrane surfaces, where gas bubbles can enter xylem under negative pressure in the process known as air seeding. Because of the insolubility of amphiphilic lipids, the surface tension influencing air seeding in pit pores is not the equilibrium surface tension of extracted bulk sap but the local surface tension at gas–liquid interfaces, which depends dynamically on the local concentration of lipids per surface area. To estimate the dynamic surface tension in lipid layers that line surfaces in the xylem apoplast, we studied the time-dependent and surface area-regulated surface tensions of apoplastic lipids extracted from xylem sap of four woody angiosperm plants using constrained drop surfactometry. Xylem lipids were found to demonstrate potent surface activity, with surface tensions reaching an equilibrium at ~25 mN m-1 and varying between a minimum of 19 mN m-1 and a maximum of 68 mN m-1 when changing the surface area between 50 and 160% around the equilibrium surface more » area. It is concluded that xylem lipid films in natural conditions most likely range from nonequilibrium metastable conditions of a supersaturated compression state to an undersaturated expansion state, depending on the local surface areas of gas–liquid interfaces. Together with findings that maximum pore constrictions in angiosperm pit membranes are much smaller than previously assumed, low dynamic surface tension in xylem turns out to be entirely compatible with the cohesion–tension and air-seeding theories, as well as with the existence of lipid-coated nanobubbles in xylem sap, and with the range of vulnerabilities to embolism observed in plants. « less
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Tree Physiology
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National Science Foundation
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  1. ABSTRACT Pit membranes in bordered pits of tracheary elements of angiosperm xylem represent primary cell walls that undergo structural and chemical modifications, not only during cell death but also during and after their role as safety valves for water transport between conduits. Cellulose microfibrils, which are typically grouped in aggregates with a diameter between 20 to 30 nm, make up their main component. While it is clear that pectins and hemicellulose are removed from immature pit membranes during hydrolysis, recent observations of amphiphilic lipids and proteins associated with pit membranes raise important questions about drought-induced embolism formation and spread via air-seeding from gas-filled conduits. Indeed, mechanisms behind air-seeding remain poorly understood, which is due in part to little attention paid to the three-dimensional structure of pit membranes in earlier studies. Based on perfusion experiments and modelling, pore constrictions in fibrous pit membranes are estimated to be well below 50 nm, and typically smaller than 20 nm. Together with the low dynamic surface tensions of amphiphilic lipids at air-water interfaces in pit membranes, 5 to 20 nm pore constrictions are in line with the observed xylem water potentials values that generally induce spread of embolism. Moreover, pit membranes appear to showmore »ideal porous medium properties for sap flow to promote hydraulic efficiency and safety due to their very high porosity (pore volume fraction), with highly interconnected, non-tortuous pore pathways, and the occurrence of multiple pore constrictions within a single pore. This three-dimensional view of pit membranes as mesoporous media may explain the relationship between pit membrane thickness and embolism resistance, but is largely incompatible with earlier, two-dimensional views on air-seeding. It is hypothesised that pit membranes enable water transport under negative pressure by producing stable, surfactant coated nanobubbles while preventing the entry of large bubbles that would cause embolism.« less
  2. Abstract

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