Xylem Conduits Research Articles

A bioinspired bubble removal method in microchannels based on angiosperm xylem embolism repair

It is difficult to remove and eliminate bubbles in microchannels in many devices used in various biomedical fields, such as those needed for microfluidic immunoassays, point-of-care testing, and cell biology evaluations. Accumulated bubbles are associated with a number of negative outcomes, including a decrease in device sensitivity, inaccuracy of analysis results, and even functional failure. Xylem conduits of angiosperm have the ability to remove bubbles in obstructed conduits. Inspired by such an embolism repair mechanism, this paper proposes a bioinspired bubble removal method, which exhibits a prominent ability to dissolve bubbles continuously within a large range of flow rates (2 µL/min–850 µL/min) while retaining the stability and continuity of the flow without auxiliary equipment. Such a method also shows significant bubble removal stability in dealing with Newtonian liquids and non-Newtonian fluids, especially with high viscosity (6.76 Pa s) and low velocity (152 nL/min). Such advantages associated with the proposed bioinspired method reveal promising application prospects in macro/microfluidic fields ranging from 3D printing, implantable devices, virus detection, and biomedical fluid processing to microscale reactor operation and beyond.

Xylem embolism spread is largely prevented by interconduit pit membranes until the majority of conduits are gas-filled.

Xylem embolism resistance varies across species influencing drought tolerance, yet little is known about the determinants of the embolism resistance of an individual conduit. Here we conducted an experiment using the optical vulnerability method to test whether individual conduits have a specific water potential threshold for embolism formation and whether pre-existing embolism in neighbouring conduits alters this threshold. Observations were made on a diverse sample of angiosperm and conifer species through a cycle of dehydration, rehydration and subsequent dehydration to death. Upon rehydration after the formation of embolism, no refilling was observed. When little pre-existing embolism was present, xylem conduits had a conserved, individual embolism-resistance threshold that varied across the population of conduits. The consequence of a variable conduit-specific embolism threshold is that a small degree of pre-existing embolism in the xylem results in apparently more resistant xylem in subsequent dehydrations, particularly in angiosperms with vessels. While our results suggest that pit membranes separating xylem conduits are critical for maintaining a conserved individual conduit threshold for embolism when little pre-existing embolism is present, as the percentage of embolized conduits increases, gas movement, local pressure differencesand connectivity between conduits increasingly contribute to embolism spread.

Seeing is believing: what visualising bubbles in the xylem has revealed about plant hydraulic function.

Maintaining water transport in the xylem is critical for vascular plants to grow and survive. The drought-induced accumulation of embolism, when gas enters xylem conduits, causes declines in hydraulic conductance (K ) and is ultimately lethal. Several methods can be used to estimate the degree of embolism in xylem, from measuring K in tissues to directly visualising embolism in conduits. One method allowing a direct quantification of embolised xylem area is the optical vulnerability (OV) technique. This method has been used across different organs and has a high spatial and temporal resolution. Here, we review studies using the OV technique, discuss the main advantages and disadvantages of this method, and summarise key advances arising from its use. Vulnerability curves generated by the OV method are regularly comparable to other methods, including the centrifuge and X-ray microtomography. A major advantage of the OV technique over other methods is that it can be simultaneously used to determine in situ embolism formation in leaves, stems and roots, in species spanning the phylogeny of land plants. The OV method has been used to experimentally investigate the spreading of embolism through xylem networks, associate embolism with downstream tissue death, and observe embolism formation in the field.

Ultrasound Pulse Emission Spectroscopy Method to Characterize Xylem Conduits in Plant Stems.

Although it is well known that plants emit acoustic pulses under drought stress, the exact origin of the waveform of these ultrasound pulses has remained elusive. Here, we present evidence for a correlation between the characteristics of the waveform of these pulses and the dimensions of xylem conduits in plants. Using a model that relates the resonant vibrations of a vessel to its dimension and viscoelasticity, we extract the xylem radii from the waveforms of ultrasound pulses and show that these are correlated and in good agreement with optical microscopy. We demonstrate the versatility of the method by applying it to shoots of ten different vascular plant species. In particular, for Hydrangea quercifolia, we further extract vessel element lengths with our model and compare them with scanning electron cryomicroscopy. The ultrasonic, noninvasive characterization of internal conduit dimensions enables a breakthrough in speed and accuracy in plant phenotyping and stress detection.

Evidence for distinct isotopic compositions of sap and tissue water in tree stems: consequences for plant water source identification.

The long-standing hypothesis that the isotopic composition of plant stem water reflects that of source water is being challenged by studies reporting bulk water from woody stems with an isotopic composition that cannot be attributed to any potential water source. The mechanism behind such source-stem water isotopic offsets is still poorly understood. Using a novel technique to extract selectively sap water from xylem conduits, we show that, in cut stems and potted plants, the isotopic composition of sap water reflects that of irrigation water, demonstrating unambiguously that no isotopic fractionation occurs during root water uptake or sap water extraction. By contrast, water in nonconductive xylem tissues is always depleted in deuterium compared with sap water, irrespective of wood anatomy. Previous studies have shown that isotopic heterogeneity also exists in soils at the pore scale in which water adsorbed onto soil particles is more depleted in deuterium than unbound water. Data collected at a riparian forest indicated that sap water matches best unbound soil water from depth below -70 cm, while bulk stem and soil water differ markedly. We conclude that source-stem isotopic offsets can be explained by micrometre-scale heterogeneity in the isotope ratios of water within woody stems and soil micro-pores.

Not all lipids in xylem conduits are artefacts. A reply to Yamagishi et al.

Not all lipids in xylem conduits are artefacts. A reply to Yamagishi et al.

Cell wall polymer distribution in bamboo visualized with in situ imaging FTIR

To better understand the high recalcitrance of bamboo during bioconversion, the fine spatial distribution of polymers in bamboo was studied with Imaging FTIR microscopy under both transmission and ATR modes, combined with PCA data processing. The results demonstrated that lignin, xylan and hydroxycinnamic acid (HCA) were more concentrated in the fibers near the xylem conduit, while cellulose was evenly distributed across the whole fiber sheath. PCA processing produced a clear separation between bamboo fibers and parenchyma cells, indicating that the parenchyma cells contains more pectin and HCA than fibers. It also demonstrated that cellulose, xylan and S-lignin were concentrated most heavily in bamboo fiber secondary cell walls, while G-lignin, pectin and HCA were found more in the compound middle lamella. The revealed information regarding polymer distribution is of great significance for better understanding of the inherent design mechanism of plant cell wall and its efficient utilization.

Solid-electrolyte interphase from nanowood for high-performance Li-metal batteries

Solid-electrolyte interphase from nanowood for high-performance Li-metal batteries

Hydraulic-stomatal coordination in tree seedlings: tight correlation across environments and ontogeny in Acer pseudoplatanus.

Hydraulic conductance is recognized as a major determinant of gas exchange and productivity. However, whether this also applies to seedlings, a critically important stage for vegetation regeneration, has been largely unknown. We analyzed the hydraulic and stomatal conductance of leaves and shoots for 6-wk-old Acer pseudoplatanus seedlings emerging in different lowland and treeline habitats and under glasshouse conditions, respectively, as well as on 9-, 15- and 18-wk-old plants, and related findings to leaf and xylem anatomical traits. Treeline seedlings had higher leaf area-specific shoot hydraulic conductance (Kshoot-L ), and stomatal conductance (gs ), associated with wider xylem conduits, lower leaf area and higher stomatal density than lowland and glasshouse-grown plants. Across the first 18 wk of development, seedlings increased four-fold in absolute shoot hydraulic conductance (Kshoot ) and declined by half in Kshoot-L , with correlated shifts in xylem and leaf anatomy. Distal leaves had higher leaf hydraulic conductance (Kleaf ) and gs compared to basal leaves. Seedlings show strong variation across growth environments and ontogenetic shifts in hydraulic and anatomical parameters. Across growth sites, ontogenetic stages and leaf orders, gs was tightly correlated with Kshoot-L and Kleaf , balancing hydraulic supply with demand for the earliest stages of seedling establishment.

K+ ions improve hydraulic conductance of grapevine shoots for sucrose loading aimed at manipulation of vine carbohydrate reserves

Introduction – Poor fruitfulness of subtropical table grapevines can reflect a low carbohydrate (CHO) status of the plant during floral bud formation and fruit development. CHO reserves can ameliorate poor fruitfulness. Such supplementation can be achieved by sucrose injection into the grapevine trunk; however, poor hydraulic conductance of stem tissues limits the success of this procedure. The objective of this study was to evaluate the co-addition of KCl to the sucrose solution to improve the hydraulic conductivity of grapevine shoots. Materials and methods – The hydraulic conductance of 15 cm long current season cane cuttings was assessed under a pressure gradient of 7.84 kPa during perfusion with water, KCl (1 to 50 mM) and sucrose solutions (2.5%, 5% and 7.5% w/v) in 0 and 10 mM KCl. Experiments were conducted over both short (about 2 hours) and long (about 78 hours) term. Additionally, transpiration of dye solutions enabled estimation of direct xylem conduits was undertaken. Results and discussion – Hydraulic conductance increased by about 20% with perfusion of ≥10 mM KCl compared to water, with the effect reversible and repeatable. Sucrose solutions caused a decrease in conductance relative to water. Co-addition of KCl mitigated the sucrose-mediated decrease in conductance. There was no evidence of physical vessel blockage such as tyloses or microbiota. Conclusion – The results are consistent with the hypothesis that cations cause shrinkage of pectins, increasing the conductance of pit membranes.

The Widened Pipe Model of plant hydraulic evolution

Shaping global water and carbon cycles, plants lift water from roots to leaves through xylem conduits. The importance of xylem water conduction makes it crucial to understand how natural selection deploys conduit diameters within and across plants. Wider conduits transport more water but are likely more vulnerable to conduction-blocking gas embolisms and cost more for a plant to build, a tension necessarily shaping xylem conduit diameters along plant stems. We build on this expectation to present the Widened Pipe Model (WPM) of plant hydraulic evolution, testing it against a global dataset. The WPM predicts that xylem conduits should be narrowest at the stem tips, widening quickly before plateauing toward the stem base. This universal profile emerges from Pareto modeling of a trade-off between just two competing vectors of natural selection: one favoring rapid widening of conduits tip to base, minimizing hydraulic resistance, and another favoring slow widening of conduits, minimizing carbon cost and embolism risk. Our data spanning terrestrial plant orders, life forms, habitats, and sizes conform closely to WPM predictions. The WPM highlights carbon economy as a powerful vector of natural selection shaping plant function. It further implies that factors that cause resistance in plant conductive systems, such as conduit pit membrane resistance, should scale in exact harmony with tip-to-base conduit widening. Furthermore, the WPM implies that alterations in the environments of individual plants should lead to changes in plant height, for example, shedding terminal branches and resprouting at lower height under drier climates, thus achieving narrower and potentially more embolism-resistant conduits.

Perspectives and design considerations of capillary-driven artificial trees for fast dewatering processes

Recent progresses on nanocapillary-driven water transport under metastable conditions have substantiated the potential of artificial trees for dewatering applications in a wide pressure range. This paper presents a comprehensive performance analysis of artificial trees encompassing the principle for negative capillary pressure generation; impacts of structural, compositional, and environmental conditions on dewatering performance; and design considerations. It begins by delineating functionalities of artificial trees for evaporation (leaves), conduction (xylem), and filtration (root) of water, in the analogy to natural trees. The analysis revealed that the magnitude of (negative) capillary pressure in the artificial leaves and xylem must be sufficiently large to overcome the osmotic pressure of feed at the root. The required magnitude can be reduced by increasing the osmotic pressure in the artificial xylem conduits, which reduces the risk of cavitation and subsequent blockage of water transport. However, a severe concentration polarization that can occur in long xylem conduits would negate such compensation effect of xylem osmotic pressure, leading to vapor pressure depression at the artificial leaves and therefore reduced dewatering rates. Enhanced Taylor dispersions by increasing xylem conduit diameters are found to alleviate the concentration polarization, allowing for water flux enhancement directly by increasing leaf-to-root membrane area ratio.

Reduced spatial resolution MRI suffices to image and quantify drought induced embolism formation in trees

BackgroundMagnetic resonance imaging (MRI) is uniquely suited to non-invasively and continuously monitor embolism formation in trees. Depending on the MRI method used, quantitative parameter maps of water content and MRI signal relaxation behavior can be generated. The ability to measure dynamic differences in water content and relaxation behavior can be used to detect xylem embolism formation, even if xylem conduits are too small to be spatially resolved. This is especially advantageous when using affordable small-scale low-field MRI scanners. The amount of signal that can be obtained from an object strongly depends on the strength of the magnetic field of the imager’s magnet. Imaging at lower resolutions thus would allow to reduce the cost, size and weight of the MRI scanner and to shorten image acquisition times.ResultsWe investigated how much spatial resolution can be sacrificed without losing the ability to monitor embolism formation in coniferous softwood (spruce, Picea abies) and diffuse porous beech (Fagus sylvatica). Saplings of both species were bench dehydrated, while they were continuously imaged at stepwise decreasing spatial resolutions. Imaging was done by means of a small-scale MRI device, utilizing image matrix sizes of 128 × 128, 64 × 64 and 32 × 32 pixels at a constant FOV of 19 and 23 mm, respectively. While images at the lowest resolutions (pixel sizes 0.59 × 0.59 mm and 0.72 × 0.72 mm) were no longer sufficient to resolve finer details of the stem anatomy, they did permit an approximate localization of embolism formation and the generation of accurate vulnerability curves.ConclusionsWhen using MRI, spatial resolution can be sacrificed without losing the ability to visualize and quantify embolism formation. Imaging at lower spatial resolution to monitor embolism formation has two advantages. Firstly, the acquisition time per image can be reduced dramatically. This enables continuous imaging at high time resolution, which may be beneficial to monitor rapid dynamics of embolism formation. Secondly, if the requirements for spatial resolution are relaxed, much simpler MRI devices can be used. This has the potential to make non-invasive MR imaging of embolism formation much more affordable and more widely available.

Diversity in conduit and pit structure among extant gymnosperm taxa.

Gymnosperm taxa are incredibly diverse in many traits, including taxa with evergreen or deciduous leaves; broad, compound, needle-like or scale-like leaves; trees, shrubs, and lianas; and taxa with seed cones that range from fleshy to woody. Although less appreciated, xylem conduits are also diverse in structure among extant gymnosperm taxa. Within the xylem of gymnosperms, axial transport occurs predominantly via tracheids, although 10-40% of gymnosperm taxa, particularly within the Gnetophyta and Cycadophyta, also contain vessels. Gymnosperm taxa vary greatly in their inter-conduit pit structure, with different types of pit membranes and pitting arrangements. While some taxa display torus-margo bordered pits (60%), many others do not contain tori (40%), and at least some taxa without tori occur within each of the four extant phyla (Coniferophyta, Cycadophyta, Ginkgophyta, and Gnetophyta). Pit membrane types vary within families but appear relatively conserved within genera. Woody species with torus-bearing pit membranes occur in colder environments (lower mean annual temperature) compared to those without tori; but occurrence does not differ with mean annual precipitation. Detailed descriptions of pit membrane types are lacking for many species and genera, indicating a need for increased anatomical study. Increased knowledge of these traits could provide a unique experimental context in which to study the evolution of conduit networks, the hydraulic implications of conduit and pit structure, and the diverse structural and functional strategies utilized by gymnosperms. There are myriad potential study questions and research opportunities within this unique and diverse group of plants.

Physiological and histological responses of peach plants grafted onto different rootstocks under calcium deficiency conditions

Calcium (Ca) is a crucial element in plant growth and development. Studies focused on the Ca deficiency in Prunus are still limited that triggered us to examine tolerance of peach plants against Ca deficiency using physiological and histological methods. Peach cultivar Rich May plants grafted onto GF 677 and Garnem rootstocks were planted in 10 L pots containing perlite. Plants were irrigated with Hoagland’s nutrient solution. After growth for two months in nutrient solution, plants were exposed to Ca deficient Hoagland solution (except control). Plants were exposed to mineral deficient conditions for 4 months. End of the study, Ca deficiency depressed plant growth, damaged cell membranes and impaired chlorophyll biosynthesis. Concentrations of chlorophyll and chlorophyll precursors were significantly decreased by Ca deficiency in both rootstocks. Cortical cell diameter decreased in both rootstocks under Ca deficiency conditions. However, cortex thickness increased in Garnem and decreased in GF 677. Increments in xylem thickness, xylem conduits length and number of xylem conduits were found in Ca deficient Garnem plants and the parameters decreased in GF 677 under Ca deficiency. Pith autolysis in midrib tissue was not observed in the current experiment. Garnem was found more tolerant rootstock than GF 677 to Ca deficiency. We consider that larger cortical cells and hyperplasy and hypertrophy of xylem vessel elements increased stomatal conductance to uptake more Ca that mitigated Ca starvation in Garnem.

Positive pressure in xylem and its role in hydraulic function.

Although transpiration-driven transport of xylem sap is well known to operate under absolute negative pressure, many terrestrial, vascular plants show positive xylem pressure above atmospheric pressure on a seasonal or daily basis, or during early developmental stages. The actual location and mechanisms behind positive xylem pressure remain largely unknown, both in plants that show seasonal xylem pressure before leaf flushing, and those that show a diurnal periodicity of bleeding and guttation. Available evidence shows that positive xylem pressure can be driven based on purely physical forces, osmotic exudation into xylem conduits, or hydraulic pressure in parenchyma cells associated with conduits. The latter two mechanisms may not be mutually exclusive and can be understood based on a similar modelling scenario. Given the renewed interest in positive xylem pressure, this review aims to provide a constructive way forward by discussing similarities and differences of mechanistic models, evaluating available evidence for hydraulic functions, such as rehydration of tissues, refilling of water stores, and embolism repair under positive pressure, and providing recommendations for future research, including methods that avoid or minimise cutting artefacts.

Lipids in xylem sap of woody plants across the angiosperm phylogeny.

Lipids have been observed attached to lumen-facing surfaces of mature xylem conduits of several plant species, but there has been little research on their functions or effects on water transport, and only one lipidomic study of the xylem apoplast. Therefore, we conducted lipidomic analyses of xylem sap from woody stems of seven plants representing six major angiosperm clades, including basal magnoliids, monocots and eudicots, to characterize and quantify phospholipids, galactolipids and sulfolipids in sap using mass spectrometry. Locations of lipids in vessels of Laurus nobilis were imaged using transmission electron microscopy and confocal microscopy. Xylem sap contained the galactolipids di- and monogalactosyldiacylglycerol, as well as all common plant phospholipids, but only traces of sulfolipids, with total lipid concentrations in extracted sap ranging from 0.18 to 0.63nmolml-1 across all seven species. Contamination of extracted sap from lipids in cut living cells was found to be negligible. Lipid composition of sap was compared with wood in two species and was largely similar, suggesting that sap lipids, including galactolipids, originate from cell content of living vessels. Seasonal changes in lipid composition of sap were observed for one species. Lipid layers coated all lumen-facing vessel surfaces of L. nobilis, and lipids were highly concentrated in inter-vessel pits. The findings suggest that apoplastic, amphiphilic xylem lipids are a universal feature of angiosperms. The findings require a reinterpretation of the cohesion-tension theory of water transport to account for the effects of apoplastic lipids on dynamic surface tension and hydraulic conductance in xylem.

Plasticity of Leaf Hydraulic Conductance in Maize in Response to Varying Nitrogen and Water Supplies

The availabilities of nitrogen (N) and water are two important environmental factors affecting crop growth and productivity in the dryland farming, how they affected leaf water transport capabilities, a bottleneck of whole-plant water transport, was seldom studied. So, the effects of different N and water supplies on leaf hydraulic conductance (Kleaf) in maize (Zea mays L.) were studied by a pot experiment, Kleaf, leaf gas exchange, vein traits and other physio-anatomical traits were measured. We found Kleaf was strongly coupled to leaf gas exchange (net photosynthetic rate, Pn, and stomatal conductance, gs) and whole-plant performance under varying N and water supplies. Long-term N supply had a greater impact than irrigation on Kleaf, Pn and gs due to the type and duration of stress, and the N and water treatments did not interact with these physiological parameters. Kleaf was related positively with the xylem conduit diameter of large longitudinal veins, the proportion of minor-vein length and interveinal distance of large and intermediate longitudinal veins, and negatively with the density of the major veins, and the interveinal distance of the transverse veins. The higher Kleaf under ambient N and water supply was also associated with larger leaf area, leaf mass per area and N content per leaf area and with an improved status of leaf water, which partly enhanced the vein traits responsible for the higher Kleaf. These results are helpful to understand leaf hydraulic strategy in adapting to long-term resource deficiency.

Tip-to-base xylem conduit widening as an adaptation: causes, consequences, and empirical priorities.

In the stems of terrestrial vascular plants studied to date, the diameter of xylem water-conducting conduits D widens predictably with distance from the stem tip L approximating D∝Lb , with b≈0.2. Because conduit diameter is central for conductance, it is essential to understand the cause of this remarkably pervasive pattern. We give reason to suspect that tip-to-base conduit widening is an adaptation, favored by natural selection because widening helps minimize the increase in hydraulic resistance that would otherwise occur as an individual stem grows longer and conductive path length increases. Evidence consistent with adaptation includes optimality models that predict the 0.2 exponent. The fact that this prediction can be made with a simple model of a single capillary, omitting much biological detail, itself makes numerous important predictions, e.g. that pit resistance must scale isometrically with conduit resistance. The idea that tip-to-base conduit widening has a nonadaptive cause, with temperature, drought, or turgor limiting the conduit diameters that plants are able to produce, is less consistent with the data than an adaptive explanation. We identify empirical priorities for testing the cause of tip-to-base conduit widening and underscore the need to study plant hydraulic systems leaf to root as integrated wholes.

Sperry’s packing rule affects the spatial proximity but not clustering of xylem conduits: the case of Fagus sylvatica L.

Abstract Sperry’s packing rule predicts the optimum packing of xylem conduits in woody plants, where the frequency of xylem conduits varies approximately inversely with the square of the conduit radius. However, it is well established that such anatomical disposition does not remain fixed but is subject to a suite of adaptations induced by physiological constraints driven by both ontogenetic development and environmental characteristics. Here we challenge the hypothesis that increasing frequency of xylem conduits, concomitant with the decrease in their lumen area along the xylem pathway, would affect the spatial distribution of vessels inside tree-rings and their aggregation. To this end, we measured the vessels’ anatomical characteristics inside each tree-ring along with a complete radial series taken at different stem heights of Fagus sylvatica L. trees. Point pattern analysis indicated a significant effect of the distance from the tree base and a weak effect of cambial age on the nearest neighbour distance among xylem vessels, suggesting that vessels were closer to each other near the apex, and became progressively more distant toward the base. The spatial pattern of xylem vessels violated the assumption of complete spatial randomness, vessel spatial arrangement followed a uniform distribution at different distances from the tree base. Although there was an increase in the intensity and proximity among vessels, we demonstrated that no patterns of aggregation between vessels were found in sampled F. sylvatica trees. Rather, point pattern profiles clearly highlighted a lack of aggregation of vessels in the face of a regular spatial distribution in the annual growth rings along the stems.