Water Transport In Plants Research Articles

Xylem Embolism Spreads by Single-Conduit Events in Three Dry Forest Angiosperm Stems.

Xylem cavitation resulting in air embolism is a major cause of plant death during drought, yet the spread of embolism throughout the plant water transport system is poorly understood. Our study used optical visualization and x-ray microcomputed tomography imaging to capture the spread of emboli in stems of three drought-resistant angiosperm trees: drooping she-oak (Allocasuarina verticillata), black wattle (Acacia mearnsii), and blue gum (Eucalyptus globulus). These species have similar degrees of xylem network connectivity (vessel grouping) with largely solitary vessels. The high temporal resolution of the optical vulnerability technique revealed that in current year branches, >80% of the cavitation events were discrete, temporally separated events in single vessels. This suggests that in xylem networks with low connectivity, embolism spread between conduits leading to multiple conduit cavitation events is uncommon. A. mearnsii showed both the highest number of multivessel cavitation events and the highest degree of vessel connectivity, suggesting a link between vessel arrangement and embolism spread. Knowledge of embolism spread will help us to uncover the links between xylem anatomy, arrangement, and the path of water flow in the xylem in diverse species to ultimately understand the drivers of cavitation and plant vulnerability to drought.

Exogenous melatonin alleviates PEG-induced short-term water deficiency in maize by increasing hydraulic conductance

BackgroundWater deficiency is likely to become more frequent and intense as a result of global climate change, which may severely impact agricultural production in the world. The positive effects of melatonin (MEL) on alleviation drought or osmotic stress-induced water deficiency in plants has been well reported. However, the underlying mechanism of MEL on the detailed process of plant water uptake and transport under water deficiency condition remains largely unknown.ResultsApplication of 1 μM MEL led to enhanced tolerance to water deficiency stress in maize seedlings, as evidenced by maintaining the higher photosynthetic parameters, leaf water status and plant transpiration rate. The relatively higher whole-plant hydraulic conductance (Kplant) and root hydraulic conductance (Lpr) in MEL-treated seedlings suggest that exogenous MEL alleviated water deficiency stress by promoting root water absorption. HgCl2 (aquaporin inhibitor) treatment inhibit the transpiration rate in MEL-treated plants greater than those of MEL-untreated; after recovery by dithiothreitol (DTT, anti-inhibitor), the transpiration rate in MEL-treated plants increased much higher than those of untreated plants. Moreover, under water deficiency, the transcription level of aquaporin genes was up-regulated by MEL application, and the H2O2 was less accumulated in MEL-treated root.ConclusionsExogenous MEL promoted aquaporin activity, which contributed to the maintaining of Lpr and Kplant under short-term water deficiency. The increased water uptake and transport lead to improved water status and thus increased tolerance to PEG-induced short-term water deficiency in maize seedlings.

Diverging Responses of Two Subtropical Tree Species (Schima superba and Cunninghamia lanceolata) to Heat Waves

The frequency and intensity of heat waves (HWs) has increased in subtropical regions in recent years. The mechanism underlying the HW response of subtropical trees remains unclear. In this study, we conducted an experiment with broad-leaved Schima superba (S. superba) and coniferous Cunninghamia lanceolata (C. lanceolata) seedlings to examine HW (5-day long) effects on stem water transport, leaf water use efficiency (WUE), morphology and growth, and to elucidate differences in the responses of both species. Our results indicated that HWs can significantly reduce hydraulic conductivity in both species. C. lanceolata experienced significant xylem embolism, with the percentage loss of conductivity (PLC) increasing by 40%, while S. superba showed a non-significant increase in PLC (+25%). Furthermore, HW also caused a reduction in photosynthesis rates (An), but transpiration rates (Tr) increased on the 5th day of the HW, together leading to a significant decrease in leaf WUE. From diurnal dynamics, we observed that the HW caused significant decrease of S. superba An only in the morning, but nearly the all day for C. lanceolata. During the morning, with a high vapor pressure deficit (VPD) environment, the HW increased Tr, which contributed a lot to latently cooling the foliage. In comparing the two tree species, we found that HW effects on S. superba were mostly short-term, with leaf senescence but limited or no xylem embolism. The surviving S. superba recovered rapidly, forming new branches and leaves, aided by their extensive root systems. For C. lanceolata, continued seedling growth initially but with subsequent xylem embolism and withering of shoots, led to stunted recovery and regrowth. In conclusion, apart from the direct thermal impacts caused by HW, drought stress was the main cause of significant negative effects on plant water transport and the photosynthetic system. Furthermore, S. superba and C. lanceolata showed clearly different responses to HW, which implies that the response mechanisms of broad-leaved and coniferous tree species to climate change can differ.

WRKY15 Suppresses Tracheary Element Differentiation Upstream of VND7 During Xylem Formation.

Formation of the vascular cylinder, a structure critical to water and nutrient transport in higher plants, is highly regulated. Here we identify WRKY15 as an important regulator that suppresses tracheary element (TE) differentiation in Arabidopsis (Arabidopsis thaliana). Overexpression of WRKY15 resulted in discontinuous protoxylem vessel files and TEs with reduced spiral wall thickening/lignification. Expression of a dominant-negative WRKY15 variant, WRKY15-EAR, led to extra protoxylem vessels and ectopic TEs with increased spiral wall thickening/lignification. Ectopic TE formation in the root cortex and hypocotyl/leaf epidermis reveals that the suppression of WRKY15 is sufficient to trigger the transdifferentiation of other types of cells to TEs. Expression profiling, RT-qPCR, and reporter analyses revealed that WRKY15 suppresses the expression of VASCULAR-RELATED NAC DOMAIN7 (VND7), a master transcriptional regulator that promotes TE differentiation. We propose that WRKY15 negatively regulates VND7 expression indirectly based on (1) the absence of a W-box in the promoter of VND7 and (2) the observation that WRKY15 and VND7 are expressed in different cells in the vascular cylinder, with WRKY15 expressed in the procambial cells and VND7 in the protoxylem poles of procambium and differentiating TEs. Future research is needed to reveal the details underlying the interaction of WRKY15 and VND7 in plant vascular development.

Investigating Effects of Bordered Pit Membrane Morphology and Properties on Plant Xylem Hydraulic Functions-A Case Study from 3D Reconstruction and Microflow Modelling of Pit Membranes in Angiosperm Xylem.

Pit membranes in between neighboring conduits of xylem play a crucial role in plant water transport. In this review, the morphological characteristics, chemical composition and mechanical properties of bordered pit membranes were summarized and linked with their functional roles in xylem hydraulics. The trade-off between xylem hydraulic efficiency and safety was closely related with morphology and properties of pit membranes, and xylem embolism resistance was also determined by the pit membrane morphology and properties. Besides, to further investigate the effects of bordered pit membranes morphology and properties on plant xylem hydraulic functions, here we modelled three-dimensional structure of bordered pit membranes by applying a deposition technique. Based on reconstructed 3D pit membrane structures, a virtual fibril network was generated to model the microflow pattern across inter-vessel pit membranes. Moreover, the mechanical behavior of intervessel pit membranes was estimated from a single microfibril’s mechanical property. Pit membranes morphology varied among different angiosperm and gymnosperm species. Our modelling work suggested that larger pores of pit membranes do not necessarily contribute to major flow rate across pit membranes; instead, the obstructed degree of flow pathway across the pit membranes plays a more important role. Our work provides useful information for studying the mechanism of microfluid flow transport across pit membranes and also sheds light on investigating the response of pit membranes both at normal and stressed conditions, thus improving our understanding on functional roles of pit membranes in xylem hydraulic function. Further work could be done to study the morphological and mechanical response of bordered pit membranes under different dehydrated conditions, as well as the related microflow behavior, based on our constructed model.

The intersection of nitrogen nutrition and water use in plants: new paths toward improved crop productivity.

Water and nitrogen availability limit crop productivity globally more than most other environmental factors. Plant availability of macronutrients such as nitrate is, to a large extent, regulated by the amount of water available in the soil, and, during drought episodes, crops can become simultaneously water and nitrogen limited. In this review, we explore the intricate relationship between water and nitrogen transport in plants, from transpiration-driven mass flow in the soil to uptake by roots via membrane transporters and channels and transport to aerial organs. We discuss the roles of root architecture and of suberized hydrophobic root barriers governing apoplastic water and nitrogen movement into the vascular system. We also highlight the need to identify the signalling cascades regulating water and nitrogen transport, as well as the need for targeted physiological analyses of plant traits influencing water and nitrogen uptake. We further advocate for incorporation of new phenotyping technologies, breeding strategies, and agronomic practices to improve crop yield in water- and nitrogen-limited production systems.

Radial water transport in arbuscular mycorrhizal maize plants under drought stress conditions is affected by indole-acetic acid (IAA) application

Drought stress is one of the most devastating abiotic stresses, compromising crop growth, reproductive success and yield. The arbuscular mycorrhizal (AM) symbiosis has been demonstrated to be beneficial in helping the plant to bear with water deficit. In plants, development and stress responses are largely regulated by a complex hormonal crosstalk. Auxins play significant roles in plant growth and development, in responses to different abiotic stresses or in the establishment and functioning of the AM symbiosis. Despite these important functions, the role of indole-3acetic acid (IAA) as a regulator of root water transport and stress response is not well understood. In this study, the effect of exogenous application of IAA on the regulation of root radial water transport in AM plants was analyzed under well-watered and drought stress conditions. Exogenous IAA application affected root hydraulic parameters, mainly osmotic root hydraulic conductivity (Lo), which was decreased in both AM and non-AM plants under water deficit conditions. Under drought, the relative apoplastic water flow was differentially regulated by IAA application in non-AM and AM plants. The effect of IAA on the internal cell component of root water conductivity suggests that aquaporins are involved in the IAA-dependent inhibition of this water pathway.

The Possible Role of Non-Structural Carbohydrates in the Regulation of Tree Hydraulics.

The xylem is a complex system that includes a network of dead conduits ensuring long-distance water transport in plants. Under ongoing climate changes, xylem embolism is a major and recurrent cause of drought-induced tree mortality. Non-structural carbohydrates (NSC) play key roles in plant responses to drought and frost stress, and several studies putatively suggest their involvement in the regulation of xylem water transport. However, a clear picture on the roles of NSCs in plant hydraulics has not been drawn to date. We summarize the current knowledge on the involvement of NSCs during embolism formation and subsequent hydraulic recovery. Under drought, sugars are generally accumulated in xylem parenchyma and in xylem sap. At drought-relief, xylem functionality is putatively restored in an osmotically driven process involving wood parenchyma, xylem sap and phloem compartments. By analyzing the published data on stem hydraulics and NSC contents under drought/frost stress and subsequent stress relief, we found that embolism build-up positively correlated to stem NSC depletion, and that the magnitude of post-stress hydraulic recovery positively correlated to consumption of soluble sugars. These findings suggest a close relationship between hydraulics and carbohydrate dynamics. We call for more experiments on hydraulic and NSC dynamics in controlled and field conditions.

La arquitectura hidráulica de las plantas vasculares terrestres, una revisión

El transporte de agua en las plantas vasculares terrestres es pasivo y está determinado por la transpiración o pérdida de agua a través de las hojas. La teoría de la cohesión-tensión es la más aceptada para explicar este proceso, y se complementa con la analogía de la ley de Ohm, que analiza el flujo del agua como un proceso catenario. La resistencia al estrés hídrico y a la cavitación está fuertemente asociada con las características anatómicas del xilema, de las punteaduras intervasculares y de sus membranas, estas últimas se alteran en función de las propiedades químicas de la solución acuosa que fluye a través de ellas. Con base en estas premisas, esta revisión aborda el fenómeno del ascenso del agua en las plantas vasculares terrestres y analiza los conceptos, las teorías y los métodos más usados en el estudio de la arquitectura hidráulica. Además, señala las diferencias en la estructura del xilema y el transporte del agua entre dicotiledóneas y monocotiledóneas.

Growing-season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity.

Stem xylem-specific hydraulic conductivity (KS ) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in KS has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global-scale patterns of KS and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species-at-site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how KS varied with climatic variables, plant functional types, and biomes. Growing-season temperature and growing-season precipitation drove global variation in KS independently. Both the mean and the variation in KS were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for KS in predicting shifts in community composition in the face of climate change.

In vivo pressure gradient heterogeneity increases flow contribution of small diameter vessels in grapevine

Leaves lose approximately 400 H2O molecules for every 1 CO2 gained during photosynthesis. Most long-distance water transport in plants, or xylem sap flow, serves to replace this water to prevent desiccation. Theory predicts that the largest vessels contribute disproportionately to overall sap flow because flow in pipe-like systems scales with the fourth power of radius. Here, we confront these theoretical flow predictions for a vessel network reconstructed from X-ray μCT imagery with in vivo flow MRI observations from the same sample of a first-year grapevine stem. Theoretical flow rate predictions based on vessel diameters are not supported. The heterogeneity of the vessel network gives rise to transverse pressure gradients that redirect flow from wide to narrow vessels, reducing the contribution of wide vessels to sap flow by 15% of the total. Our results call for an update of the current working model of the xylem to account for its heterogeneity.

Contribution of the arbuscular mycorrhizal symbiosis to the regulation of radial root water transport in maize plants under water deficit

In roots, water flows radially through three parallel pathways: apoplastic, symplastic and transcellular (the last two referred as the cell-to-cell), with a different contribution depending on the environmental conditions. Thus, during drought, the cell-to-cell pathway, which is largely regulated by aquaporins, dominates. While it is accepted that water can flow across roots following the apoplastic, symplastic and transcellular pathways, the relative contribution of these pathways to whole root hydraulic conductivity is not well stablished. In addition, the symbiosis with arbuscular mycorrhizal (AM) fungi was reported to modify root water transport in host plants. This study aims to understand if the AM symbiosis alters radial root water transport in the host plant and whether this modification is due to alteration of plant aquaporins activity or amounts and/or changes in apoplastic barriers. Hence, the combined effect of mycorrhizal fungus, water deficit and application of the aquaporin inhibitor sodium azide (NaN3) on radial root water transport of maize plants was analyzed. The development of Casparian bands in these roots was also assessed. NaN3 clearly inhibited osmotic root hydraulic conductivity (Lo). However, the inhibitory effect of sodium azide on Lo was lower in AM plants than in non-AM plants, which together with their higher relative apoplastic water flow values suggests a compensatory mechanism for aquaporin activity inhibition in AM plants, leading to a higher hydrostatic root hydraulic conductivity (Lpr) compared to non-AM plants. This effect seems to be related to the mycorrhizal regulation of aquaporins activity through posttranslational modifications. The development of Casparian bands increased with drought and AM colonization, although this did not decrease water flow values in AM plants. The work provides new clues on the differential mycorrhizal regulation of root water transport.

Vessel-length determination using silicone and air injection: are there artifacts?

Xylem vessels are used by most angiosperm plants for long-distance water and nutrient transport. Vessel length is one of the key functional traits determining plant water-transport efficiency. Additionally, determination of maximum vessel length is necessary for correct sample collection and measurements in hydraulic studies to avoid open-vessel and cutting-under-tension artifacts. Air injection and silicone injection (BLUESIL RTV141A and B mixtures) are two widely used methods for maximum vessel length determination. However, the validity of both methods needs to be carefully tested for species with different vessel lengths. In this study, we tested the air-injection and silicone-injection methods using eight species with different vessel lengths: short (<0.5m), medium (0.5-1m) and long (>1m). We employed a novel approach using RTV141A injection without the RTV141B hardener as a reference method because RTV141A cannot penetrate inter-vessel pit membranes and is not prone to hardening/solidification effects during the injection process. The results revealed that the silicone-injection method substantially underestimated the maximum vessel length of all eight species. However, the air-injection method tended to overestimate the maximum vessel length in five out of eight species. The ratio of underestimation of the silicone-injection method was higher for species with longer vessels, but the overestimation of the air-injection method was independent of the vessel length. Moreover, air injection with different pressures-ranging from 40 to 300kPa-resulted in comparable results. We conclude that the conventional silicone-injection method can underestimate the vessel length, whereas the air-injection method can overestimate the maximum vessel length, particularly for long-vessel led species. We recommend RTV141A-only injection for determining the maximum vessel length, and it can also be used to validate the use of the air-injection and conventional silicone-injection methods for a given species.

A new approach for modelling water transport in fossil plants

ABSTRACTThe origin of xylem in the Silurian was a major step in plant evolution, leading to diverse growth forms with various mechanical and hydraulic properties. In the fossil record, these properties can only be investigated using models based on extant plant physiology. Regarding hydraulics, previous studies have considered either the properties of a single tracheid or of a set of independent tubes. Here, we use the analogy between the flow of water under tension in a plant and an electrical circuit to develop an extension of Wilson’s single tracheid model to the tissue scale. Upscaling to the tissue-level allows considering wood as a heterogeneous tissue by taking into account differences in tracheid density and the presence of rays. The new model provides a more biologically accurate representation of fossil wood hydraulic properties. The single tracheid and new tissue models are applied to two conspecific specimens of Callixylon (Progymnospermopsida, Archeopteridales) from the Late Devonian of Morocco. Differences are shown at the tissue level that cannot be suspected at the single tracheid level. Callixylon represents the first trees with a conifer-like wood and is a major component of Late Devonian floras world-wide. Our results show that the anatomical disparity of its wood might have led to hydraulic plasticity, allowing growth in various environmental conditions. More generally, the new tissue-model suggests that the various combinations of tracheid and ray sizes present in Palaeozoic plants might have led to a higher variety of ecophysiologies than suspected based solely on the properties of individual tracheids.

Surveillance of cell wall diffusion barrier integrity modulates water and solute transport in plants

The endodermis is a key cell layer in plant roots that contributes to the controlled uptake of water and mineral nutrients into plants. In order to provide such functionality the endodermal cell wall has specific chemical modifications consisting of lignin bands (Casparian strips) that encircle each cell, and deposition of a waxy-like substance (suberin) between the wall and the plasma membrane. These two extracellular deposits provide control of diffusion enabling the endodermis to direct the movement of water and solutes into and out of the vascular system in roots. Loss of integrity of the Casparian strip-based apoplastic barrier is sensed by the leakage of a small peptide from the stele into the cortex. Here, we report that such sensing of barrier integrity leads to the rebalancing of water and mineral nutrient uptake, compensating for breakage of Casparian strips. This rebalancing involves both a reduction in root hydraulic conductivity driven by deactivation of aquaporins, and downstream limitation of ion leakage through deposition of suberin. These responses in the root are also coupled to a reduction in water demand in the shoot mediated by ABA-dependent stomatal closure.

Functional Status of Xylem Through Time.

Water transport in vascular plants represents a critical component of terrestrial water cycles and supplies the water needed for the exchange of CO2 in the atmosphere for photosynthesis. Yet, many fundamental principles of water transport are difficult to assess given the scale and location of plant xylem. Here we review the mechanistic principles that underpin long-distance water transport in vascular plants, with a focus on woody species. We also discuss the recent development of noninvasive tools to study the functional status of xylem networks in planta. Limitations of current methods to detect drought-induced xylem blockages (e.g., embolisms) and quantify corresponding declines in sap flow, and the coordination of hydraulic dysfunction with other physiological processes are assessed. Future avenues of research focused on cross-validation of plant hydraulics methods are discussed, as well as a proposed fundamental shift in the theory and methodology used to characterize and measure plant water use.

Preparation and characterization of epicuticular wax films

Dipping films from epicuticular wax (EW) were prepared as model systems of epicuticular wax films found in plants. In these films, the growth uniformity, surface morphology, and hydrophobicity were examined. It was observed growth uniformity (linear growth) only from the fifth layer onwards because of the influence of substrate. The surface morphology of the films was found to be composed of pores formed by aggregates of EW molecules, both with a fractal form. An increase in the number of film layers resulted in the increase of the number of pores up to a maximum value followed by a decrease. Such increase was assigned to the growth of aggregates whereas the decrease was explained by the increase of pore sizes, because during the growth of the aggregates, the small pores are replaced by the large pores. Hydrophobicity increased with the number of layers, which was associated with the increase of irregularities on the surface caused by the pores and aggregates. In addition, it was observed that the number of pores increased with temperature. This was explained by the increase in the mobility of EW molecules, which led to a larger amount of EW molecules deposited. Based on our results and the advantages offered by dipping films – including the control of thickness and structure – this type of film is feasible as a model for studies of cuticular water transport in plants.

Water transport, perception, and response in plants.

Sufficient water availability in the environment is critical for plant survival. Perception of water by plants is necessary to balance water uptake and water loss and to control plant growth. Plant physiology and soil science research have contributed greatly to our understanding of how water moves through soil, is taken up by roots, and moves to leaves where it is lost to the atmosphere by transpiration. Water uptake from the soil is affected by soil texture itself and soil water content. Hydraulic resistances for water flow through soil can be a major limitation for plant water uptake. Changes in water supply and water loss affect water potential gradients inside plants. Likewise, growth creates water potential gradients. It is known that plants respond to changes in these gradients. Water flow and loss are controlled through stomata and regulation of hydraulic conductance via aquaporins. When water availability declines, water loss is limited through stomatal closure and by adjusting hydraulic conductance to maintain cell turgor. Plants also adapt to changes in water supply by growing their roots towards water and through refinements to their root system architecture. Mechanosensitive ion channels, aquaporins, proteins that sense the cell wall and cell membrane environment, and proteins that change conformation in response to osmotic or turgor changes could serve as putative sensors. Future research is required to better understand processes in the rhizosphere during soil drying and how plants respond to spatial differences in water availability. It remains to be investigated how changes in water availability and water loss affect different tissues and cells in plants and how these biophysical signals are translated into chemical signals that feed into signaling pathways like abscisic acid response or organ development.

Modelling water fluxes in plants: from tissues to biosphere.

Contents Summary 1207 I. Introduction 1207 II. A brief history of modelling plant water fluxes 1208 III. Main components of plant water transport models 1208 IV. Stand-scale water fluxes and coupling to climate and soil 1213 V. Water fluxes in terrestrial biosphere models and feedbacks to community dynamics 1215 VI. Outstanding challenges in modelling water fluxes in the soil-plant-atmosphere continuum 1217 Acknowledgements 1218 References 1218 SUMMARY: Models of plant water fluxes have evolved from studies focussed on understanding the detailed structure and functioning of specific components of the soil-plant-atmosphere (SPA) continuum to architectures often incorporated inside eco-hydrological and terrestrial biosphere (TB) model schemes. We review here the historical evolution of this field, examine the basic structure of a simplified individual-based model of plant water transport, highlight selected applications for specific ecological problems and conclude by examining outstanding issues requiring further improvements in modelling vegetation water fluxes. We particularly emphasise issues related to the scaling from tissue-level traits to individual-based predictions of water transport, the representation of nonlinear and hysteretic behaviour in soil-xylem hydraulics and the need to incorporate knowledge of hydraulics within broader frameworks of plant ecological strategies and their consequences for predicting community demography and dynamics.

Effects of gravity direction on water transport in sweetpotato plants

Effects of gravity direction on water transport in sweetpotato plants