Nicolas Lenoir

Evidence for hydraulic vulnerability segmentation and lack of xylem refilling under tension

Direct, noninvasive observations of embolism formation and repair reveal a lack of refilling under negative pressure and a xylem hydraulic vulnerability segmentation in grapevine. The vascular system of grapevine (Vitis spp.) has been reported as being highly vulnerable, even though grapevine regularly experiences seasonal drought. Consequently, stomata would remain open below water potentials that would generate a high loss of stem hydraulic conductivity via xylem embolism. This situation would necessitate daily cycles of embolism repair to restore hydraulic function. However, a more parsimonious explanation is that some hydraulic techniques are prone to artifacts in species with long vessels, leading to the overestimation of vulnerability. The aim of this study was to provide an unbiased assessment of (1) the vulnerability to drought-induced embolism in perennial and annual organs and (2) the ability to refill embolized vessels in two Vitis species X-ray micro-computed tomography observations of intact plants indicated that both Vitis vinifera and Vitis riparia were relatively vulnerable, with the pressure inducing 50% loss of stem hydraulic conductivity = −1.7 and −1.3 MPa, respectively. In V. vinifera, both the stem and petiole had similar sigmoidal vulnerability curves but differed in pressure inducing 50% loss of hydraulic conductivity (−1.7 and −1 MPa for stem and petiole, respectively). Refilling was not observed as long as bulk xylem pressure remained negative (e.g. at the apical part of the plants; −0.11 ± 0.02 MPa) and change in percentage loss of conductivity was 0.02% ± 0.01%. However, positive xylem pressure was observed at the basal part of the plant (0.04 ± 0.01 MPa), leading to a recovery of conductance (change in percentage loss of conductivity = −0.24% ± 0.12%). Our findings provide evidence that grapevine is unable to repair embolized xylem vessels under negative pressure, but its hydraulic vulnerability segmentation provides significant protection of the perennial stem.

Xylem resistance to embolism: Presenting a simple diagnostic test for the open vessel artefact

Xylem vulnerability to embolism represents an essential trait for the evaluation of the impact of hydraulics in plant function and ecology. The standard centrifuge technique is widely used for the construction of vulnerability curves, although its accuracy when applied to species with long vessels remains under debate. We developed a simple diagnostic test to determine whether the open-vessel artefact influences centrifuge estimates of embolism resistance. Xylem samples from three species with differing vessel lengths were exposed to less negative xylem pressures via centrifugation than the minimum pressure the sample had previously experienced. Additional calibration was obtained from non-invasive measurement of embolism on intact olive plants by X-ray microtomography. Results showed artefactual decreases in hydraulic conductance (k) for samples with open vessels when exposed to a less negative xylem pressure than the minimum pressure they had previously experienced. X-Ray microtomography indicated that most of the embolism formation in olive occurs at xylem pressures below -4.0 MPa, reaching 50% loss of hydraulic conductivity at -5.3 MPa. The artefactual reductions in k induced by centrifugation underestimate embolism resistance data of species with long vessels. A simple test is suggested to avoid this open vessel artefact and to ensure the reliability of this technique in future studies.

Tomography and imaging at the PSICHE beam line of the SOLEIL synchrotron

PSICHE (Pressure, Structure and Imaging by Contrast at High Energy) is the high-energy beam line of the SOLEIL synchrotron. The beam line is designed to study samples at extreme pressures, using diffraction, and to perform imaging and tomography for materials science and other diverse applications. This paper presents the tomograph and the use of the beam line for imaging, with emphasis on developments made with respect to existing instruments. Of particular note are the high load capacity rotation stage with free central aperture for installing large or complex samples and sample environments, x-ray mirror and filter optics for pink beam imaging, and multiple options for combining imaging and diffraction measurement. We describe how x-ray imaging techniques have been integrated into high-pressure experiments. The design and the specifications of the beam line are described, and several case studies drawn from the first user experiments are presented.

Time-resolved 2D concentration maps in flowing suspensions using X-ray

In this paper, we introduce a new technique based on X-ray radiography with high temporal (O(0.1u2009s)) and spatial (O(10u2009μm)) resolutions to study fast suspension flows regardless of optical access. We benefit from the axial symmetry of our flow configuration, a wide gap Couette setup, to extract a 3D solid volume fraction field from a single X-ray projection image. We propose a mathematical algorithm based on the inversion of Abel transform in conjunction with H1 regularization and data denoising to measure the solid volume fraction field in suspensions in a fraction of a second. We show that the results are in excellent agreement with those obtained from micro Computed Tomography (CT scan) in one hour. This significant reduction in the data acquisition time opens a new avenue in the field of suspensions. As a proof of concept, we study the kinetics of shear-induced migration for suspensions of particles in both Newtonian and yield stress suspending fluids. The latter experiments include two different conditions: With and without a plug region. In both cases, we are able to capture in detail the kinetics of migration. In the presence of a plug region, we manage to accurately describe the particle accumulation at the interface between the sheared and the static regions. Remarkably, even in the absence of sedimentation, the concentration profiles show a complex 2D structure, with no z-invariant region, which illustrates the strong impact of top and bottom boundary effects on migration. We also show the importance of boundary effects on the shear induced migration of particles in a Newtonian suspending fluid. This further shows the necessity of developing techniques that give access to the full spatial concentration field, as the one we present here.In this paper, we introduce a new technique based on X-ray radiography with high temporal (O(0.1u2009s)) and spatial (O(10u2009μm)) resolutions to study fast suspension flows regardless of optical access. We benefit from the axial symmetry of our flow configuration, a wide gap Couette setup, to extract a 3D solid volume fraction field from a single X-ray projection image. We propose a mathematical algorithm based on the inversion of Abel transform in conjunction with H1 regularization and data denoising to measure the solid volume fraction field in suspensions in a fraction of a second. We show that the results are in excellent agreement with those obtained from micro Computed Tomography (CT scan) in one hour. This significant reduction in the data acquisition time opens a new avenue in the field of suspensions. As a proof of concept, we study the kinetics of shear-induced migration for suspensions of particles in both Newtonian and yield stress suspending fluids. The latter experiments include two different co...

An inconvenient truth about xylem resistance to embolism in the model species for refilling Laurus nobilis L.

Key messageDirect, non-invasive X-ray microtomography and optical technique observations applied in stems and leaves of intact seedlings revealed that laurel is highly resistant to drought-induced xylem embolism. Contrary to what has been brought forward, daily cycles of embolism formation and refilling are unlikely to occur in this species and to explain how it copes with drought.ContextThere has been considerable controversy regarding xylem embolism resistance for long-vesselled angiosperm species and particularly for the model species for refilling (Laurus nobilis L.).AimsThe purpose of this study was to resolve the hydraulic properties of this species by documenting vulnerability curves of different organs in intact plants.MethodsHere, we applied a direct, non-invasive method to visualize xylem embolism in stems and leaves of intact laurel seedlings up to 2-m tall using X-ray microtomography (microCT) observations and the optical vulnerability technique. These approaches were coupled with complementary centrifugation measurements performed on 1-m long branches sampled from adult trees and compared with additional microCT analyses carried out on 80-cm cut branches.ResultsDirect observations of embolism spread during desiccation of intact laurels revealed that 50% loss of xylem conductivity (Ψ50) was reached at −u20097.9u2009±u20090.5 and −u20098.4u2009±u20090.3xa0MPa in stems and leaves, respectively, while the minimum xylem water potentials measured in the field were −u20094.2xa0MPa during a moderate drought season. Those findings reveal that embolism formation is not routine in Laurus nobilis contrary to what has been previously reported. These Ψ50 values were close to those based on the flow-centrifuge technique (−u20099.2u2009±u20090.2xa0MPa), but at odds with microCT observations of cut branches (−u20094.0u2009±u20090.5xa0MPa).ConclusionIn summary, independent methods converge toward the same conclusion that laurel is highly resistant to xylem embolism regardless its development stage. Under typical growth conditions without extreme drought events, this species maintains positive hydraulic safety margin, while daily cycles of embolism formation and refilling are unlikely to occur in this species.