Abstract
The geometrical structure and spatial arrangement of lumens, bordered pits, and scalariform perforation plates in xylem vessels modulate water flow from roots to leaves. Understanding their respective hydraulic functions is essential to unveil how plants regulate their hydraulic networks to facilitate the ascent of sap under biotic and abiotic stresses but is challenging because of the opaque nature of the vessel networks and water flow within them. We made the first-ever effort to discern the difference between lumens and scalariform perforation plates in cotton in impeding water flow in single vessels and vessel networks using X-ray tomography and pore-scale numerical simulation. Three-dimensional structures of xylem vessels in the stem of two cotton cultivars were acquired non-invasively using X-ray computed tomography (CT) at high spatial resolution, and a lattice Boltzmann model was developed to simulate water flow through the xylem networks at micrometer scale. The detailed water velocity and pressure simulated using the model were used to calculate the hydraulic resistance caused by the lumens and the scalariform perforation plates in individual vessels and the vessel networks of the two cotton cultivars. The results showed that the hydraulic resistance spiked whenever water flowed across a perforation plate and that the overall hydraulic resistance caused by the perforation plates in an individual vessel accounted for approximately 54% of the total resistance of the vessel. We also calculated the hydraulic conductance of individual vessels and vessel networks using the simulated water velocity and pressure at micrometer scale and compared it with those estimated from the Hagen Poiseuille (HP) equation as commonly used in the literature by approximating the xylem vessels in the cotton as isolated tubes. While it was found that the HP equation overestimated the hydraulic conductance by more than 200%, the overestimate was largely due to the incapability of the HP equation to represent the perforation plates rather than its approximation of the irregular vessels by circular tubes.
Highlights
Vascular plants use their xylem networks to ascend water and mineral nutrients from soil to leaves to sustain photosynthesis (Sack and Holbrook, 2006)
The purpose of this paper is to combine these pore-scale models with X-ray computed tomography (CT) to directly calculate, for the first time, the hydraulic resistances caused by each lumen and perforation plate in single vessels as well as their consequence for hydraulic conductance of the xylem vessel network in cotton
Water ascent in the network was simulated using the lattice Boltzmann model under a pressure gradient, and the simulated water velocity and pressure at micrometer scale were used to analyse the resistance induced by perforation plates and lumens in single vessels and vessel networks, respectively
Summary
Vascular plants use their xylem networks to ascend water and mineral nutrients from soil to leaves to sustain photosynthesis (Sack and Holbrook, 2006). Simplifying the irregular vessels as straight tubes and omitting the perforation plates and bordered pits are known to give rise to errors, but the magnitude of the errors caused by each varied widely from less than 10% (Schulte and Castle, 1993) to more than 200% (Christman and Sperry, 2010). They appear to depend on plant species (Trueba et al, 2019) and the experimental methods used to measure the hydraulic conductance (Chiu and Ewers, 1993)
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