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Intervessel pits are considered to function as valves that avoid embolism spreading and optimize efficient transport of xylem sap across neighbouring vessels. Hydraulic transport between vessels would therefore follow a safety-efficiency trade-off, which is directly related to the total intervessel pit area (A p ), inversely related to the pit membrane thickness (T PM ) and driven by a pressure difference. To test this hypothesis, we modelled the relative transport rate of gas (k a ) and water (Q) at the intervessel pit level for 23 angiosperm species and correlated these parameters with the water potential at which 50% of embolism occurs (Ψ 50 ). We also measured k a for 10 species using pneumatic measurements. The pressure difference across adjacent vessels and estimated values of k a and Q were related to Ψ 50 , following a convex safety-efficiency trade-off based on modelled and experimental data. Minor changes in T PM and A p exponentially affected the pressure difference and flow, respectively. Our results provide clear evidence that a xylem safety-efficiency trade-off is not linear, but convex due to flow across intervessel pit membranes, which represent mesoporous media within microporous conduits. Moreover, the convex nature of long-distance xylem transport may contribute to an adjustable fluid balance of plants, depending on environmental conditions., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Embolism spreading in angiosperm xylem occurs via mesoporous pit membranes between vessels. Here, we investigate how the size of pore constrictions in pit membranes is related to pit membrane thickness and embolism resistance. Pit membranes were modelled as multiple layers to investigate how pit membrane thickness and the number of intervessel pits per vessel determine pore constriction sizes, the probability of encountering large pores, and embolism resistance. These estimations were complemented by measurements of pit membrane thickness, embolism resistance, and number of intervessel pits per vessel in stem xylem (n = 31, 31 and 20 species, respectively). The modelled constriction sizes in pit membranes decreased with increasing membrane thickness, explaining the measured relationship between pit membrane thickness and embolism resistance. The number of pits per vessel affected constriction size and embolism resistance much less than pit membrane thickness. Moreover, a strong relationship between modelled and measured embolism resistance was observed. Pore constrictions provide a mechanistic explanation for why pit membrane thickness determines embolism resistance, which suggests that hydraulic safety can be uncoupled from hydraulic efficiency. Although embolism spreading remains puzzling and encompasses more than pore constriction sizes, angiosperms are unlikely to have leaky pit membranes, which enables tensile transport of water., (© 2021 The Authors New Phytologist © 2021 New Phytologist Trust.)

Maintaining xylem water transport under drought is vital for plants, but xylem failure does occur when drought-induced embolisms form and progressively spread through the xylem. The hydraulic method is widely considered the gold standard to quantify drought-induced xylem embolism. The method determines hydraulic conductivity (Kh) in cut branch samples, dehydrated to specific drought levels, by pushing water through them. The technique is widely considered for its reliable Kh measurements, but there is some uncertainty in the literature over how to define stable Kh and how that relates to the degree of xylem embolism formation. Therefore, the most common setup for this method was extended to measure four parameters: (i) inlet Kh, (ii) outlet Kh, (iii) radial flow from xylem to surrounding living tissue and (iv) the pressure difference across the sample. From a strictly theoretical viewpoint, hydraulic steady state, where inflow equals outflow and radial flow is zero, will result in stable Kh. Application of the setup to Malus domestica Borkh. branches showed that achieving hydraulic steady state takes considerable time (up to 300 min) and that time to reach steady state increased with declining xylem water potentials. During each experimental run, Kh and xylem water potentials dynamically increased, which was supported by X-ray computed microtomography visualizations of embolism refilling under both high- (8 kPa) and low-pressure (2 kPa) heads. Supplying pressurized water can hence cause artificial refilling of vessels, which makes it difficult to achieve a truly stable Kh in partially embolized xylem., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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 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. [ABSTRACT FROM AUTHOR]

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 show 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. [ABSTRACT FROM AUTHOR]

The article focuses on the process of ascent of sap in xylem tissue of plant in presence of bubbles. Topics discussed include response of plant toward climate change, cohesion-tension theory of water transport in plant, entering of air bubbles in xylem, increase in vulnerability of xylem for embolism formation due to addition of artificial surfactants, and intriguing property of bubbles which are surfactant coated.