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Drought poses a serious threat to the productivity of hop, an important perennial crop. However, the precise physiological mechanisms that make it highly susceptible to drought are not yet fully understood. In this study, we investigated stomatal regulation and xylem vulnerability to embolism, which are important traits closely associated with plant drought resistance. In a glasshouse cultivation experiment, we monitored changes in leaf water potential, stem elongation rates, and leaf gas exchange, including net photosynthetic rates, stomatal conductance, and intrinsic water use efficiency, on relatively young hop plants (traditional Saaz - Osvald's clone 31) exposed to declining soil water availability. The transpiration rate and stem elongation of plants decreased significantly with a small decline in substrate water potential (ΨSUB), indicating a highly sensitive stomata response during early phases of soil dehydration. The stem elongation was completely halted, and the transpiration rate dropped to less than 50% of its maximum at ΨSUB levels below − 0.8 MPa. In well-watered hop plants, xylem in stems operates near the initial point of embolization and is highly vulnerable to embolism, with a water potential corresponding to a 50% loss of xylem conductivity at -1.6 MPa. The sensitive stomatal response to declining ΨSUB likely helps to mitigate the risk of hydraulic failure, albeit at the cost of impaired growth. Scheduled irrigation, particularly during the sensitive stem elongation stage, may be a promising approach to mitigate the detrimental effects of reduced soil water availability on hop growth and yield while also conserving water resources. [ABSTRACT FROM AUTHOR]

The xylem undergoes physiological changes in response to various environmental conditions during the process of plant growth. To understand these physiological changes, it is extremely important to observe the transport of xylem. In this study, the distribution and structure of vascular bundle in Lilium lancifolium were observed using the method of semithin section. Methods for introducing a fluorescent tracer into the xylem of the stems were evaluated. Then, the transport rule of 5(6)-Carboxyfluorescein diacetate (CFDA) in the xylem of the stem of L. lancifolium was studied by fluorescence dye in live cells tracer technology. The results showed that the vascular bundles of L. lancifolium were scattered in the basic tissue, the peripheral vascular bundles were smaller and densely distributed, and the closer to the center, the larger the volume of vascular bundles and the more sparsely distributed. The vascular bundles of L. lancifolium are limited external tenacity vascular bundles, which are composed of phloem and xylem. The most suitable method for CFDA labeling the xylem of isolated stem segments of L. lancifolium was solution soaking for 24 h. The running speed of CF in the isolated stem was 0.3 cm/h, which was consistent with the running speed of the material in the field. CF could be transported between the xylem and parenchyma cells, indicating that the material transport in the xylem could be through the symplastic pathway. The above results laid a foundation for the study of the xylem transport mechanism and the xylem pathogen disease of lily. [ABSTRACT FROM AUTHOR]

Sap exudation is the process whereby trees such as sugar (Acer saccharum Marsh.) and red maple (Acer rubrum L.) generate unusually high positive stem pressure in response to repeated cycles of freeze and thaw. This elevated xylem pressure permits the sap to be harvested over a period of several weeks and hence is a major factor in the viability of the maple syrup industry. The extensive literature on sap exudation documents competing hypotheses regarding the physical and biological mechanisms that drive positive pressure generation in maple, but to date, relatively little effort has been expended on devising mathematical models for the exudation process. In this paper, we utilize an existing model of Graf et al. (J Roy Soc Interface 12:20150665, 2015) that describes heat and mass transport within the multiphase gas–liquid–ice mixture in the porous xylem tissue. The model captures the inherent multiscale nature of xylem transport by including phase change and osmotic transport in wood cells on the microscale, which is coupled to heat transport through the tree stem on the macroscale. A parametric study based on simulations with synthetic temperature data identifies the model parameters that have greatest impact on stem pressure build-up. Measured daily temperature fluctuations are then used as model inputs and the resulting simulated pressures are compared directly with experimental measurements taken from mature red and sugar maple stems during the sap harvest season. The results demonstrate that our multiscale freeze–thaw model reproduces realistic exudation behavior, thereby providing novel insights into the specific physical mechanisms that dominate positive pressure generation in maple trees. [ABSTRACT FROM AUTHOR]

In this study, Cd isotope analysis was conducted on drought-tolerant (cowpea and sesame) and less drought-tolerant vegetables (water spinach, green pepper, and mung bean) to elucidate the mechanisms underlying Cd uptake and transport. Cd isotopes in plants were identical to or lighter than those in the available pool and exhibited negative fractionation from roots to straws (Δ114/110Cd = −0.22 ‰ to −0.17 ‰) in drought-tolerant vegetables, whereas contrasting results were obtained for less drought-tolerant vegetables (Δ114/110Cd = −0.050 ‰ to 0.39 ‰). Positive Cd isotope fractionation from straws to fruits in drought-tolerant vegetables (Δ114/110Cd = 0.33 ‰ ± 0.03 ‰ and 0.10 ‰ ± 0.03 ‰, respectively) was observed, whereas negligible or negative fractionation was found in less drought-tolerant vegetables (Δ114/110Cd = 0.01 ‰ ± 0.04 ‰ and −0.34 ‰ ± 0.02 ‰, respectively). The vast secretion of organic acids might have led to positive available pool-to-roots and negative roots-to-straws isotope fractionation in drought-tolerant vegetables. In contrast, preferential xylem transport resulted in negative straws-to-fruits isotope fractionation in less drought-tolerant vegetables. This study demonstrated that Cd isotope fractionation in the soil-plant system is associated with plant drought tolerance, and drought-tolerant and less-tolerant plants developed a distinct Cd detoxification mechanism, corresponding to a reversed fractionation of Cd isotopes. [Display omitted] • Lower soil pH shifted Cd isotope ratio of available pool toward bulk soil. • Cd isotope fractionation in plants was opposite depending on their drought tolerance. • Cd uptake, root-to-straw transport, and loading into fruit caused a successively reversed fractionation. • Cd uptake and transport mechanisms were differentiated based on drought tolerance. • Isotope effect during straw-to-fruit transport may relate to the transpiration ability. [ABSTRACT FROM AUTHOR]

Copper (Cu) is an essential micronutrient for humans, but excessive Cu in rice grains causes health risks. Currently, the mechanisms underlying Cu accumulation in rice are unclear. Here, we identified a novel member of the high-affinity copper transporter (Ctr)-like (COPT) protein family in rice, OsCOPT7, which controls Cu accumulation in rice grains. Mutation in the coding sequence of OsCOPT7 (mutant lc1) leads to inhibition of Cu transport through the xylem, contributing to lower Cu concentrations in the grain of lc1. Knockout or modulation of the expression of OsCOPT7 significantly impacts Cu transportation in the xylem and its accumulation in rice grains. OsCOPT7 localizes at the multi-pass membrane in the cell and the gene is expressed in the exodermis and stele cells, facilitating Cu loading into the xylem. OsCOPT7 expression is upregulated under Cu deficiency and in various organs, implying its contribution to Cu distribution within the rice plant. The variable expression pattern of OsCOPT7 suggests that OsCOPT7 expression responds to Cu stress in rice. Moreover, assays reveal that OsCOPT7 expression level is suppressed by the SQUAMOSA promoter-binding protein-like 9 (OsSPL9) and that OsCOPT7 interacts with Antioxidant Protein1 (OsATX1). This study elucidates the involvement of OsCOPT7 in Cu loading into the xylem, its subsequent distribution within the rice plant, and the potential of this protein in reducing the risk of high Cu concentrations in rice grain grown on Cu-contaminated soil. [Display omitted] ● Inhibition of Cu transport from xylem induces the special phenotype of low Cu accumulation in lc1 mutant. ● OsCOPT7 is the causal gene relative to low Cu accumulation in lc1. ● Changing the activity of OsCOPT7 affects Cu accumulation and transport through xylem in rice. ● OsCOPT7 interacts with OsATX1 involving in Cu loading into xylem in rice. [ABSTRACT FROM AUTHOR]

Determining the relationship between reductions in stomatal conductance (gs) and leaf water transport during dehydration is key to understanding plant drought responses. While numerous studies have analysed the hydraulic function of woody species, minimal research has been conducted on grasses. Here, we sought to characterize hydraulic vulnerability in five widely‐occurring pasture grasses (including both C3 and C4 grasses) and determine whether reductions in gs and leaf hydraulic conductance (Kleaf) during dehydration could be attributed to xylem embolism. Using the optical vulnerability (OV) technique, we found that all species were highly resistant to xylem embolism when compared to other herbaceous angiosperms, with 50% xylem embolism (PX50) occurring at xylem pressures ranging from −4.4 to −6.1 MPa. We observed similar reductions in gs and Kleaf under mild water stress for all species, occurring well before PX50. The onset of xylem embolism (PX12) occurred consistently after stomatal closure and 90% reduction of Kleaf. Our results suggest that factors other than xylem embolism are responsible for the majority of reductions in gs and Kleaf during drought and reductions in the productivity of pasture species under moderate drought may not be driven by embolism. Summary Statement: Pasture grasses are highly resistant to xylem cavitation, with cavitation occurring consistently after significant reductions in stomatal conductance (gs) and leaf hydraulic conductance (Kleaf). Our results suggest that factors other than xylem cavitation are responsible for reductions in gs and Kleaf during moderate drought with cavitation only occurring under extreme drought conditions. [ABSTRACT FROM AUTHOR]

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 differences and connectivity between conduits increasingly contribute to embolism spread. Summary statement: We find that pit membranes are able to prevent embolism spreading between gas and water‐filled conduits until there is a considerable number of embolized conduits. Consequently, conduits appear to have specific thresholds of embolism resistance, and that a small degree of pre‐existing embolism in the xylem may make the remaining xylem apparently more resistant. [ABSTRACT FROM AUTHOR]

The mechanisms by which woody plants recover xylem hydraulic capacity after drought stress are not well understood, particularly with regard to the role of embolism refilling. We evaluated the recovery of xylem hydraulic capacity in young Eucalyptus saligna plants exposed to cycles of drought stress and rewatering. Plants were exposed to moderate and severe drought stress treatments, with recovery monitored at time intervals from 24 h to 6 months after rewatering. The percentage loss of xylem vessels due to embolism (PLV) was quantified at each time point using microcomputed tomography with stem water potential (Ψx) and canopy transpiration (Ec) measured before scans. Plants exposed to severe drought stress suffered high levels of embolism (47.38% ± 10.97% PLV) and almost complete canopy loss. No evidence of embolism refilling was observed at 24 h, 1 week, or 3 weeks after rewatering despite rapid recovery in Ψx. Recovery of hydraulic capacity was achieved over a 6‐month period by growth of new xylem tissue, with canopy leaf area and Ec recovering over the same period. These findings indicate that E. saligna recovers slowly from severe drought stress, with potential for embolism to persist in the xylem for many months after rainfall events. Summary Statement: After a cycle of drought and rehydration, Eucalyptus saligna plants recovered lost xylem hydraulic capacity by growth of new xylem tissue rather than refilling of embolised vessels. [ABSTRACT FROM AUTHOR]

Summary: Experimental evidence that nonstomatal limitations to photosynthesis (NSLs) correlate with leaf sugar and/or leaf water status suggests the possibility that stomata adjust to maximise photosynthesis through a trade‐off between leaf CO2 supply and NSLs, potentially involving source–sink interactions. However, the mechanisms regulating NSLs and sink strength, as well as their implications for stomatal control, remain uncertain.We used an analytically solvable model to explore optimal stomatal control under alternative hypotheses for source and sink regulation. We assumed that either leaf sugar concentration or leaf water potential regulates NSLs, and that either phloem turgor pressure or phloem sugar concentration regulates sink phloem unloading.All hypotheses led to realistic stomatal responses to light, CO2 and air humidity, including conservative behaviour for the intercellular‐to‐atmospheric CO2 concentration ratio. Sugar‐regulated and water‐regulated NSLs are distinguished by the presence/absence of a stomatal closure response to changing sink strength. Turgor‐regulated and sugar‐regulated phloem unloading are distinguished by the presence/absence of stomatal closure under drought and avoidance/occurrence of negative phloem turgor. Results from girdling and drought experiments on Pinus sylvestris, Betula pendula, Populus tremula and Picea abies saplings are consistent with optimal stomatal control under sugar‐regulated NSLs and turgor‐regulated unloading.Our analytical results provide a simple representation of stomatal responses to above‐ground and below‐ground environmental factors and sink activity. [ABSTRACT FROM AUTHOR]

Root pressure, also manifested as profusive sap flowing from cut stems, is a phenomenon in some species that has perplexed biologists for much of the last century. It is associated with increased crop production under drought, but its function and regulation remain largely unknown. In this study, we investigated the initiation, mechanisms, and possible adaptive function of root pressure in six genotypes of Sorghum bicolor during a drought experiment in the greenhouse. We observed that root pressure was induced in plants exposed to drought followed by re-watering but possibly inhibited by 100% re-watering in some genotypes. We found that root pressure in drought stressed and re-watered plants was associated with greater ratio of fine: coarse root length and shoot biomass production, indicating a possible role of root allocation in creating root pressure and adaptive benefit of root pressure for shoot biomass production. Using RNA-Seq, we identified gene transcripts that were up- and down-regulated in plants with root pressure expression, focusing on genes for aquaporins, membrane transporters, and ATPases that could regulate inter- and intra-cellular transport of water and ions to generate positive xylem pressure in root tissue.

Root pressure, also manifested as profusive sap flowing from cut stems, is a phenomenon in some species that has perplexed biologists for much of the last century. It is associated with increased crop production under drought, but its function and regulation remain largely unknown. In this study, we investigated the initiation, mechanisms, and possible adaptive function of root pressure in six genotypes of Sorghum bicolor during a drought experiment in the greenhouse. We observed that root pressure was induced in plants exposed to drought followed by re-watering but possibly inhibited by 100% re-watering in some genotypes. We found that root pressure in drought stressed and re-watered plants was associated with greater ratio of fine: coarse root length and shoot biomass production, indicating a possible role of root allocation in creating root pressure and adaptive benefit of root pressure for shoot biomass production. Using RNA-Seq, we identified gene transcripts that were up- and down-regulated in plants with root pressure expression, focusing on genes for aquaporins, membrane transporters, and ATPases that could regulate inter- and intra-cellular transport of water and ions to generate positive xylem pressure in root tissue. [ABSTRACT FROM AUTHOR]

Wintergreen fern Polystichum acrostichoides has fronds that are photosynthetically active year-round, despite diurnal and seasonal changes in soil moisture, air temperature and light availability. This species can fix much of its annual carbon during periods when the deciduous canopy is open. Yet, remaining photosynthetically active year-round requires the maintenance of photosynthetic and hydraulic systems that are vulnerable to freeze–thaw cycles. We aimed to determine the anatomical and physiological strategies P. acrostichoides uses to maintain positive carbon gain, and the coordination between the hydraulic and photosynthetic systems. We found that the first night below 0 °C led to 25 % loss of conductivity (PLC) in stipes, suggesting that winter-induced embolism occurred. Maximum photosynthetic rate and chlorophyll fluorescence declined during winter but recovered by spring, despite PLC remaining high; the remaining hydraulic capacity was sufficient to supply the leaves with water. The onset of colder temperatures coincided with the development of a necrotic hinge zone at the stipe base, allowing fronds to overwinter lying prostrate and maintain a favourable leaf temperature. Our conductivity data show that the hinge zone did not affect leaf hydraulics because of the flexibility of the vasculature. Collectively, these strategies help P. acrostichoides to survive in northeastern forests. [ABSTRACT FROM AUTHOR]

Increased abiotic stress along with increasing temperatures, dry periods and forest disturbances may favor biotic stressors such as simultaneous invasion of bark beetle and ophiostomatoid fungi. It is not fully understood how tree desiccation is associated with colonization of sapwood by fungi. A decrease in xylem sap surface tension (σxylem) as a result of infection has been hypothesized to cause xylem embolism by lowering the threshold for air-seeding at the pits between conduits and disruptions in tree water transport. However, this hypothesis has not yet been tested. We investigated tree water relations by measuring the stem xylem hydraulic conductivity (Kstem), σxylem, stem relative water content (RWCstem), and water potential (Ψstem), and canopy conductance (gcanopy), as well as the compound composition in xylem sap in Norway spruce (Picea abies) saplings. We conducted our measurements at the later stage of Endoconidiophora polonica infection when visible symptoms had occurred in xylem. Saplings of two clones (44 trees altogether) were allocated to treatments of inoculated, wounded control and intact control trees in a greenhouse. The saplings were destructively sampled every second week during summer 2016. σxylem, Kstem and RWCstem decreased following the inoculation, which may indicate that decreased σxylem resulted in increased embolism. gcanopy did not differ between treatments indicating that stomata responded to Ψstem rather than to embolism formation. Concentrations of quinic acid, myo-inositol, sucrose and alkylphenol increased in the xylem sap of inoculated trees. Myo-inositol concentrations also correlated negatively with σxylem and Kstem. Our study is a preliminary investigation of the role of σxylem in E. polonica infected trees based on previous hypotheses. The results suggest that E. polonica infection can lead to a simultaneous decrease in xylem sap surface tension and a decline in tree hydraulic conductivity, thus hampering tree water transport. [ABSTRACT FROM AUTHOR]

Grafting is widely used in fruit, vegetable, and flower propagation to improve biotic and abiotic stress resistance, yield, and quality. At present, the systemic changes caused by grafting, as well as the mechanisms and effects of long-distance signal transport between rootstock and scion have mainly been investigated in model plants (Arabidopsis thaliana and Nicotiana benthamiana). However, these aspects of grafting vary when different plant materials are grafted, so the study of model plants provides only a theoretical basis and reference for the related research of grafted vegetables. The dearth of knowledge about the transport of signaling molecules in grafted vegetables is inconsistent with the rapid development of large-scale vegetable production, highlighting the need to study the mechanisms regulating the rootstock-scion interaction and long-distance transport. The rapid development of molecular biotechnology and "omics" approaches will allow researchers to unravel the physiological and molecular mechanisms involved in the rootstock–scion interaction in vegetables. We summarize recent progress in the study of the physiological aspects (e.g., hormones and nutrients) of the response in grafted vegetables and focus in particular on long-distance molecular signaling (e.g., RNA and proteins). This review provides a theoretical basis for studies of the rootstock–scion interaction in grafted vegetables, as well as provide guidance for rootstock breeding and selection to meet specific demands for efficient vegetable production. [ABSTRACT FROM AUTHOR]

Summary: Photosynthetic rate is concurrently limited by stomatal limitations and nonstomatal limitations (NSLs). However, the controls on NSLs to photosynthesis and their coordination with stomatal control on different timescales remain poorly understood. According to a recent optimization hypothesis, NSLs depend on leaf osmotic or water status and are coordinated with stomatal control so as to maximize leaf photosynthesis.Drought and notching experiments were conducted on Pinus sylvestris, Picea abies, Betula Pendula and Populus tremula seedlings in glasshouse conditions to study the dependence of NSLs on leaf osmotic and water status, and their coordination with stomatal control, on timescales of minutes and weeks, to test the assumptions and predictions of the optimization hypothesis.Both NSLs and stomatal conductance followed power‐law functions of leaf osmotic concentration and leaf water potential. Moreover, stomatal conductance was proportional to the square root of soil‐to‐leaf hydraulic conductance, as predicted by the optimization hypothesis.Though the detailed mechanisms underlying the dependence of NSLs on leaf osmotic or water status lie outside the scope of this study, our results support the hypothesis that NSLs and stomatal control are coordinated to maximize leaf photosynthesis and allow the effect of NSLs to be included in models of tree gas‐exchange. [ABSTRACT FROM AUTHOR]

A highly sensitive quantitative method for assaying nicotianamine (NA) and 2'-deoxymugineic acid (DMA) using liquid chromatography electrospray ionization time-of-flight mass spectrometry (LC-ESI-TOF-MS) was developed. This method requires fewer sample preparation steps than high-performance liquid chromatography (HPLC). The amino and hydroxyl groups of NA and DMA were derivatized using 9-fluorenylmethoxycarbonyl chloride, which improved their retention on a C18 reversed-phase column and facilitated separation. The amounts of NA and DMA in 10 μl of xylem sap from rice cultivated under Fe-sufficient and Fe-deficient conditions were quantified without concentration. In Fe-sufficient plants, the concentrations of NA and DMA were almost equal to that of Fe. In Fe-deficient plants, the concentration of NA did not change significantly, whereas that of DMA increased markedly.

Fluorescent dyes are often used to observe transport mechanisms in plant vascular tissues. However, it has been technically challenging to apply fluorescent dyes on roots to monitor xylem transport in vivo. Here, we present a fast, noninvasive, and high-throughput protocol to monitor xylem transport in seedlings. Using the fluorescent dyes 5(6)-carboxyfluorescein diacetate (CFDA) and Rhodamine WT, we were able to observe xylem transport on a cellular level in Arabidopsis thaliana roots. We describe how to apply these dyes on primary roots of young seedlings, how to monitor root-to-shoot xylem transport, and how to measure xylem transport velocity in roots. Moreover, we show that our protocol can also be applied to lateral roots and grafted seedlings to assess xylem (re)connection. Altogether, these techniques are useful for investigating xylem functionality in diverse experimental setups., (© 2024. The Author(s), under exclusive license to Springer Science+Business Media, LLC, part of Springer Nature.)

It is well known that xylem embolism can be repaired by bark water uptake and that the sugar required for embolism refilling can be provided by corticular photosynthesis. However, the relationship between corticular photosynthesis and embolism repair by bark water uptake is still poorly understood. In this study, the role of corticular photosynthesis in embolism repair was assessed using Salix matsudana branch segments dehydrated to −1.9 MPa (P50, water potential at 50% loss of conductivity). The results indicated that corticular photosynthesis significantly promoted water uptake and nonstructural carbohydrate (NSC) accumulation in the bark and xylem during soaking, thereby effectively enhancing the refilling of the embolized vessels and the recovery of hydraulic conductivity. Furthermore, the influence of the extent of dehydration on the embolism refilling enhanced by corticular photosynthesis was investigated. The enhanced refilling effects were much higher in the mildly dehydrated (−1.5 MPa) and moderately dehydrated (−1.9 MPa) branch segments than in the severely dehydrated (−2.2 MPa) branch segments. This study provides evidence that corticular photosynthesis plays a crucial role in xylem embolism repair by bark water uptake for mildly and moderately dehydrated branches. Corticular photosynthesis promoted water uptake and nonstructural carbohydrate (NSC) accumulation in the bark and xylem of dehydrated Salix matsudana branch segments during soaking. Maintenance of corticular photosynthesis in the wetted branch segments effectively enhanced the refilling of embolized vessels and the recovery of hydraulic conductivity. [ABSTRACT FROM AUTHOR]

The genus Pinus has wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade‐offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species: Pinus echinata, Pinus elliottii, Pinus palustris, Pinus taeda, and Pinus virginiana. We found that increasing needle length contributed to a trade‐off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity. We quantified the variation in leaf trait values (needle size, hydraulic conductance, and photosynthetic capacity) of five co‐occurring southern pine species and assessed the combination of characteristics as solutions to match energy capture by leaf surfaces with water supply to stomata. We found that, from the shortest to the longest needles, mechanical tissue fraction increased by 50% and hydraulic conductance by fourfold. Although the consequences on carbon gain and water use of these various needle structural designs need to be evaluated at higher hierarchical scales, our results suggest that needle size strongly influences their anatomical traits, which, in turn, are reflected in needle mechanical support and physiology. [ABSTRACT FROM AUTHOR]

Summary: Sap molecules are transported by xylem flow throughout the whole plant body. Factors regulating the xylem transport of different molecules remain to be identified.We used fluorophores to visualize xylem transport from roots to leaves in Arabidopsis thaliana. Several previously established Arabidopsis lines with modified xylem cell walls were used to determine the contribution of xylem cell walls to xylem transport.Fluorophores underwent xylem flow‐dependent transport from roots to leaves within 20 min. A comparison of rhodamine, fluorescein and three fluorescently labeled CLV3/ESR‐related (CLE) peptides revealed cargo‐dependent xylem transport patterns in terms of leaf position and vein order. Only minor changes in amino acid sequence were sufficient to alter the xylem transport patterns of the labeled CLE peptides. We found that the xylem transport pattern of fluorescein was affected in Arabidopsis lines with modified AtXYN1,LAC4 or CCoAOMT1 expression. In these lines, application of a defense inducer, pipecolic acid, to roots resulted in altered defense response patterns in leaves, whereas all the lines showed wild‐type‐like responses when pipecolic acid was sprayed onto leaves.The combined results reveal a finely controlled cargo‐dependent xylem transport and suggest that the xylem cell wall structure is crucial for this transport system. [ABSTRACT FROM AUTHOR]

The study of tree rings can reveal long-term records of a tree's response to the environment. This dendroecological approach, when supplemented with finer-scale observations of the xylem anatomy, can provide novel information about a tree's year-to-year anatomical and hydraulic adjustments. Here we use this method in aspen (Populus tremuloides Michx.) to identify xylem response to drought and insect defoliation. Surprisingly, we found that precipitation influenced vessel diameter mostly in the trees' youth, while this correlation was less pronounced at maturity. This is likely due to a reduction in stress the stand experiences as it ages, and reflects an ability to mediate drought stress as trees mature. Defoliation events caused consistent and profound changes in fiber anatomy likely leading to reduced structural support to vessels. We therefore expect that in years of defoliation trees may be vulnerable to drought-induced cavitation when leaf area recovers. This study highlights how the inclusion of cellular level measurements in tree ring studies provides additional information on how stress events may alter tree functioning through alterations in structure. [ABSTRACT FROM AUTHOR]

Abstract: Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known. We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period. The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations. Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed. [ABSTRACT FROM AUTHOR]

Abstract: Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially explicit, analytical water transport model for stems. Our model allows Huber ratio, stem‐saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole‐stem emergent property resulting from non‐linear pressure‐conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits. [ABSTRACT FROM AUTHOR]

Background and Aims Transport of carbohydrates and water are essential aspects of plant function. The aim of this study was to develop and test the methods for mechanistic modelling of quasi-stationary coupled phloem/ xylem transport in the context of functional--structural plant modelling. Methods The novelty of this approach is in combining analytical and computational methods. The plant structure is modelled at a metamer level with the internodes represented by conduit elements and the lateral organs represented by sources and sinks. Transport equations are solved analytically for each internode and then the solutions are adjusted and 'sewn' together using an iterative computational procedure taking into account concentration-dependent sinks and sources. The model is implemented in L-studio and uses the aspect-oriented modelling approach for phloem/xylem coupling. Key Results To our knowledge, this is the first transport model that provides continuous distributions of the system variables in a complex developing structure. The model takes into account non-linear dependence of phloem resistance and osmotic potential on the local carbohydrate concentration. The model solutions show excellent agreement with the existing results of other analytical and numerical models. These comparisons confirm the validity of the approximations made in the model. Combining analytical and computational methods made it possible to take into account continuous sink/source distribution within internodes without much increase in the complexity of the computational procedure, because the necessary changes in the model were mostly in the analytical part. The results emphasize sensitivity of phloem flux and lateral xylem flux to the presence of distributed sinks and sources along the transport system. Conclusions The presented approach provides a new insight into mechanistic modelling of phloem/xylem transport in growing plants. It will be useful for both fine-scale modelling of carbohydrate dynamics and for creating simpler models at a growth unit level. [ABSTRACT FROM AUTHOR]

Abstract: Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature piñon (Pinus edulis) and juniper (Juniperus monosperma) after 3+ years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation‐exclusion (−45% ambient precipitation), and exclusion‐structure control. Supplemental watering elevated leaf water potential, sapwood‐area specific hydraulic conductivity, and leaf‐area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted KL in piñon but not juniper. Piñon and juniper achieved similar KL under ambient conditions, but juniper matched or outperformed piñon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests piñon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA. [ABSTRACT FROM AUTHOR]

While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source-sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source-sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area. [ABSTRACT FROM AUTHOR]

Drought-induced tree decline is a complex event, and recent hypotheses suggest that hydraulic failure and carbon starvation are co-responsible for this process. We tested the possible role of non-structural carbohydrates (NSC) content on post-drought hydraulic recovery, to verify the hypothesis that embolism reversal represents a mechanistic link between carbon starvation and stem hydraulics. Measurements were performed in laurel plants subjected to similar water stress levels either over short or long term, to induce comparable embolism levels. Plants subjected to mild and prolonged water shortage (S) showed reduced growth, adjustment of turgor loss point driven by changes in both osmotic potential at full turgor and bulk modulus of elasticity, a lower content of soluble NSC and a higher content of starch with respect to control (C) plants. Moreover, S plants showed a lower ability to recover from xylem embolism than C plants, even after irrigation. Our data suggest that plant carbon status might indirectly influence plant performance during and after drought via effects on xylem hydraulic functioning, supporting the view of a possible mechanistic link between the two processes. [ABSTRACT FROM AUTHOR]

Trees must simultaneously balance their CO2 uptake rate via stomata, photosynthesis, the transport rate of sugars and rate of sugar utilization in sinks while maintaining a favourable water and carbon balance. We demonstrate using a numerical model that it is possible to understand stomatal functioning from the viewpoint of maximizing the simultaneous photosynthetic production, phloem transport and sink sugar utilization rate under the limitation that the transpiration-driven hydrostatic pressure gradient sets for those processes. A key feature in our model is that non-stomatal limitations to photosynthesis increase with decreasing leaf water potential and/or increasing leaf sugar concentration and are thus coupled to stomatal conductance. Maximizing the photosynthetic production rate using a numerical steady-state model leads to stomatal behaviour that is able to reproduce the well-known trends of stomatal behaviour in response to, e.g., light, vapour concentration difference, ambient CO2 concentration, soil water status, sink strength and xylem and phloem hydraulic conductance. We show that our results for stomatal behaviour are very similar to the solutions given by the earlier models of stomatal conductance derived solely from gas exchange considerations. Our modelling results also demonstrate how the 'marginal cost of water' in the unified stomatal conductance model and the optimal stomatal model could be related to plant structural and physiological traits, most importantly, the soil-to-leaf hydraulic conductance and soil moisture. [ABSTRACT FROM AUTHOR]

In this review, we address the relationship between stomatal behaviour, water potential regulation and hydraulic transport in plants, focusing on the implications for the iso/anisohydric classification of plant drought responses at seasonal timescales. We first revise the history of the isohydric concept and its possible definitions. Then, we use published data to answer two main questions: (1) is greater stomatal control in response to decreasing water availability associated with a tighter regulation of leaf water potential (ΨL) across species? and (2) is there an association between tighter ΨL regulation (~isohydric behaviour) and lower leaf conductance over time during a drought event? These two questions are addressed at two levels: across species growing in different sites and comparing only species coexisting at a given site. Our analyses show that, across species, a tight regulation of ΨL is not necessarily associated with greater stomatal control or with more constrained assimilation during drought. Therefore, iso/anisohydry defined in terms of ΨL regulation cannot be used as an indicator of a specific mechanism of drought-induced mortality or as a proxy for overall plant vulnerability to drought. [ABSTRACT FROM AUTHOR]

Symbiotic nitrogen fixation in legumes is strongly diminished by mineral nitrogen. Nevertheless, recent work with some tropical legumes revealed low sensitivity to mineral nitrogen or even enhancement of nodulation by ammonium. Thus, in this study, we investigated plant growth and nodulation of Inga edulis over a period of 128 days using different sources of nitrogen, i.e. ammonium (15 mM), nitrate (15 mM) and symbiotic nitrogen fixation. Plant growth was evaluated through dry mass, height, stem diameter and root/shoot ratio as well as nodulation by nodule number and dry mass. Nitrate, ureides and total amino acids were determined in roots, shoots and xylem sap. All three nitrogen sources were found to stimulate growth relative to the noninoculated N-free control, however ammonium was the most effective. Nodulation, both nodule number and mass, was strongly reduced by nitrate but not ammonium. The transport of total amino acids in the xylem sap was stimulated by ammonium without change in the composition with asparagine predominating in all treatments. Inga edulis can benefit from fertilisation with ammonium since even at high concentrations growth was improved without impairing nodulation, a condition appropriate for restoration of soil conditions. [ABSTRACT FROM AUTHOR]

Calcium has well-documented roles in plant signaling, water relations and cell wall interactions. Significant research into how calcium impacts these individual processes in various tissues has been carried out; however, the influence of calcium on fruit ripening has not been thoroughly explored. Here, we review the current state of knowledge on how calcium may impact fruit development, physical traits and disease susceptibility through facilitating developmental and stress response signaling, stabilizing membranes, influencing water relations and modifying cell wall properties through cross-linking of de-esterified pectins. We explore the involvement of calcium in hormone signaling integral to ripening and the physiological mechanisms behind common disorders that have been associated with fruit calcium deficiency (e.g. blossom end rot in tomatoes or bitter pit in apples). This review works towards an improved understanding of how the many roles of calcium interact to influence fruit ripening, and proposes future research directions to fill knowledge gaps. Specifically, we focus mostly on grapes and present a model that integrates existing knowledge around these various functions of calcium in fruit, which provides a basis for understanding the physiological impacts of sub-optimal calcium nutrition in grapes. Calcium accumulation and distribution in fruit is shown to be highly dependent on water delivery and cell wall interactions in the apoplasm. Localized calcium deficiencies observed in particular species or varieties can result from differences in xylem morphology, fruit water relations and pectin composition, and can cause leaky membranes, irregular cell wall softening, impaired hormonal signaling and aberrant fruit development. We propose that the role of apoplasmic calcium-pectin crosslinking, particularly in the xylem, is an understudied area that may have a key influence on fruit water relations. Furthermore, we believe that improved knowledge of the calcium-regulated signaling pathways that control ripening would assist in addressing calcium deficiency disorders and improving fruit pathogen resistance.

Sulphur (S) is available to plants as sulphate (SO42-) in the soil to the roots and as gaseous SO2 in the atmosphere to the shoots. Of all the macronutrient, S is perhaps the nutrient which has attracted the most attention in soil science and plant nutrition due to its potential defensive characteristics to pests, good nutritive potentiality to crops and its relative immobility in the soil-plant system. This review highlights the mechanisms of uptake and translocation of S from soil and its transport in xylem and phloem tissues of plants. Moreover, it also throws new insights, have led us to revisit the hypothesis of S chelation in reference to oxidative stress induced by heavy metals sequestration.

Plant vascular systems are constructed by specific cell wall modifications through which cells are highly specialized to make conduits for water and nutrients. Xylem vessels are formed by thickened cell walls that remain after programmed cell death, and serve as water conduits from the root to the shoot. In contrast, phloem tissues consist of a complex of living cells, including sieve tube elements and their neighboring companion cells, and translocate photosynthetic assimilates from mature leaves to developing young tissues. Intensive studies on the content of vascular flow fluids have unveiled that plant vascular tissues transport various types of gene product, and the transport of some provides the molecular basis for the long-distance communications. Analysis of xylem sap has demonstrated the presence of proteins in the xylem transpiration stream. Recent studies have revealed that CLE and CEP peptides secreted in the roots are transported to above ground via the xylem in response to plant-microbe interaction and soil nitrogen starvation, respectively. Their leucine-rich repeat transmembrane receptors localized in the shoot phloem are required for relaying the signal from the shoot to the root. These findings well fit to the current scenario of root-to-shoot-to-root feedback signaling, where peptide transport achieves the root-to-shoot signaling, the first half of the signaling process. Meanwhile, it is now well evidenced that proteins and a range of RNAs are transported via the phloem translocation system, and some of those can exert their physiological functions at their destinations, including roots. Thus, plant vascular systems may serve not only as conduits for the translocation of essential substances but also as long-distance communication pathways that allow plants to adapt to changes in internal and external environments at the whole plant level.

Ecosystem models have difficulty predicting plant drought responses, partially from uncertainty in the stomatal response to water deficits in soil and atmosphere. We evaluate a 'supply-demand' theory for water-limited stomatal behavior that avoids the typical scaffold of empirical response functions. The premise is that canopy water demand is regulated in proportion to threat to supply posed by xylem cavitation and soil drying., The theory was implemented in a trait-based soil-plant-atmosphere model. The model predicted canopy transpiration ( E), canopy diffusive conductance ( G), and canopy xylem pressure ( Pcanopy) from soil water potential ( Psoil) and vapor pressure deficit ( D)., Modeled responses to D and Psoil were consistent with empirical response functions, but controlling parameters were hydraulic traits rather than coefficients. Maximum hydraulic and diffusive conductances and vulnerability to loss in hydraulic conductance dictated stomatal sensitivity and hence the iso- to anisohydric spectrum of regulation. The model matched wide fluctuations in G and Pcanopy across nine data sets from seasonally dry tropical forest and piñon-juniper woodland with < 26% mean error., Promising initial performance suggests the theory could be useful in improving ecosystem models. Better understanding of the variation in hydraulic properties along the root-stem-leaf continuum will simplify parameterization. [ABSTRACT FROM AUTHOR]

In a series of experiments with Ricinus communis, abscisic acid (ABA) concentrations in tissues and transport saps, its de novo biosynthesis, long-distance transport, and metabolism (degradation) were affected by nutritional conditions, nitrogen (N) source, and nutrient limitation, or salt stress. In the present study these data were statistically reevaluated, and new correlations presented that underpin the importance of this universal phytohormone. The biggest differences in ABA concentration were observed in xylem sap. N source had the strongest effect; however, nutrient limitation (particularly phosphorus limitation) and salt also had significant effects. ABA was found in greater concentration in phloem sap compared with xylem sap; however, the effect of treatment on ABA concentration in phloem was lower. In the leaves, ABA concentration was most variable compared with the other tissues. This variation was only affected by the N source. In roots, ABA was significantly decreased by nutrient limitation. Of the compartments in which ABA was quantified, xylem sap ABA concentration was most significantly correlated with leaf stomatal conductance and leaf growth. Additionally, ABA concentration in xylem was significantly correlated to that in phloem, indicating a 6-fold concentration increase from xylem to phloem. The ABA flow model showed that biosynthesis of ABA in roots affected the xylem flow of ABA. Moreover, ABA concentration in xylem affected the degradation of the phytohormone in shoots and also its export from shoots via phloem. The role of phloem transport is discussed since it stimulates ABA metabolism in roots. [ABSTRACT FROM AUTHOR]

Field studies analyzing the stable isotope composition of xylem water are providing important information on ecosystem water relations. However, the capacity of stable isotopes to characterize the functioning of plants in their environment has not been fully explored because of methodological constraints on the extent and resolution at which samples could be collected and analysed. Here, we introduce an in situ method offering the potential to continuously monitor the stable isotope composition of tree xylem water via its vapour phase using a commercial laser-based isotope analyser and compact microporous probes installed into the xylem. Our technique enables efficient high-frequency measurement with intervals of only a few minutes per sample while eliminating the need for costly and cumbersome destructive collection of plant material and laboratory-based processing. We present field observations of xylem water hydrogen and oxygen isotope compositions obtained over several days including a labelled irrigation event and compare them against results from concurrent destructive sampling with cryogenic distillation and mass spectrometric analysis. The data demonstrate that temporal changes as well as spatial patterns of integration in xylem water isotope composition can be resolved through direct measurement. The new technique can therefore present a valuable tool to study the hydraulic architecture and water utilization of trees. [ABSTRACT FROM AUTHOR]

Regulation of water flow in an interconnected xylem vessel network enables plants to survive despite challenging environment changes that can cause xylem embolism. In this study, vulnerability to embolisms of xylem vessels and their water-refilling patterns in vascular bundles of maize leaves were experimentally investigated by employing synchrotron X-ray micro-imaging technique. A vascular bundle in maize consisted of a protoxylem vessel with helical thickenings between two metaxylem vessels with single perforation plates and nonuniformly distributed pits. When embolism was artificially induced in excised maize leaves by exposing them to air, protoxylem vessels became less vulnerable to dehydration compared tometaxylemvessels. After supplying water into the embolized vascular bundles, when water-refilling process stopped at the perforation plates in metaxylem vessels, discontinuous radial water influx occurred surprisingly in the adjacent protoxylem vessels. Alternating water refilling pattern in protoxylem and metaxylem vessels exhibited probable correlation between the incidence location and time of water refilling and the structural properties of xylem vessels. These results imply that the maintenance of water transport and modulation of water refilling are affected by hydrodynamic roles of perforation plates and radial connectivity in a xylem vascular bundle network. [ABSTRACT FROM AUTHOR]

Despite the crucial role of carbon transport in whole plant physiology and its impact on plant-environment interactions and ecosystem function, relatively little research has tried to examine how phloem physiology impacts plant ecology. In this review, we highlight several areas of active research where inquiry into phloem physiology has increased our understanding of whole plant function and ecological processes. We consider how xylem-phloem interactions impact plant drought tolerance and reproduction, how phloem transport influences carbon allocation in trees and carbon cycling in ecosystems and how phloem function mediates plant relations with insects, pests, microbes and symbiotes. We argue that in spite of challenges that exist in studying phloem physiology, it is critical that we consider the role of this dynamic vascular system when examining the relationship between plants and their biotic and abiotic environment. [ABSTRACT FROM AUTHOR]

Background and Aims Tyloses and embolisms have been reported to impair water transport during the development of Pierce's disease ( PD), caused by the xylem-dwelling pathogen X ylella fastidiosa. This work investigates the relative importance of these xylem conduit obstructions in stems of inoculated V itis vinifera plants. Methods and Results After 18 weeks in a greenhouse, internodes from control and needle-inoculated Chardonnay vines were assessed for embolisms using non-destructive magnetic resonance imaging ( MRI). Tyloses were identified by subsequent destructive histology and light microscopy examination. Embolism of individual or small sectors of vessels was detected by MRI analysis of a control shoot, usually in the proximity of the pith; virtually no tyloses were observed in the corresponding histological sections. From a symptomatic (inoculated) shoot, MRIs revealed large areas of embolised vessels extending radially towards the epidermis in most of the internodes; several individual or groups of two to three vessels appeared to be occluded with tyloses. Conclusions In all assessed internodes, MRI-detected embolisms were found more frequently than tyloses, suggesting that over the course of PD embolism formation precedes tylose development. Significance of the Study Embolisms appear to be the first and predominant type of occlusion in stems during early PD progression; nevertheless, their importance for explaining water transport impairment has been overlooked in most studies. [ABSTRACT FROM AUTHOR]