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Detached leaves of ivy (Hedera helix L.) were used as a model for the study of zinc uptake and transport in vascular plants. By the uptake via the surface of fully immersed leaves in 25 % Hoagland nutrient media (HM) spiked with 65ZnCl2 (50 μmol/dm3 ZnCl2), concentration in leaves 4.98 μg Zn/g (dry wt.), i. e. 2.6 μg Zn/dm2 leaf area after 7d exposition were obtained. By the uptake via immersed stalks of not immersed (transpiring) leaves concentrations up to 370 μg Zn/g (dry wt.) were obtained. When Zn enters into detached leaves via the surface of immersed leaf blades, zinc is uniformly distributed in leaf blades and leaf stalks. When zinc enters detached leaves via immersed stalks of non-immersed transpiring leaves, only small part of zinc is transported to leaf blades and the prevailing part remains in leaf stalks. Stalks act as a trap, able to prevent other leaf tissues against inhibitory effects of high Zn concentrations. Mineral nutrient salts in solutions mobilize Zn trapped in leaf stalks and facilitate Zn transport by transpiration stream to leaf blades, what means that Zn in stalks is bound in ion-exchageable forms.

H2 18 O enrichment develops when leaves transpire, but an accurate generalized mechanistic model has proven elusive. We hypothesized that leaf hydraulic architecture may affect the degree to which gradients in H2 18 O develop within leaves, influencing bulk leaf stable oxygen isotope enrichment (ΔL ) and the degree to which the Peclet effect is relevant in leaves. Leaf hydraulic design predicted the relevance of a Peclet effect to ΔL in 19 of the 21 species tested. Leaves with well-developed hydraulic connections between the vascular tissue and the epidermal cells through bundle sheath extensions and clear distinctions between palisade and spongy mesophyll layers (while the mesophyll is hydraulically disconnected) may have velocities of the transpiration stream such that gradients in H2 18 O develop and are expressed in the mesophyll. In contrast, in leaves where the vascular tissue is hydraulically disconnected from the epidermal layers, or where all mesophyll cells are well connected to the transpiration stream, velocities within the liquid transport pathways may be low enough that gradients in H2 18 O are very small. Prior knowledge of leaf hydraulic design allows informed selection of the appropriate ΔL modelling framework.

Premise Water stored in the xylem of woody plants is important for supporting the transpiration stream under prolonged drought, yet the source of stored water within the xylem during drought remains unclear. Insights into xylem water utilization during drought will uncover the adaptation strategies of the test species to stress. Methods To fill the existing knowledge gap, we excised twigs of Abies firma (Japanese fir, conifer), Cercidiphyllum japonicum (katsura tree, diffuse-porous) and Quercus serrata (konara oak, ring-porous) to quantify interspecific variation of water transfer in xylem corresponding with increasing cumulative water release (CWR) using micro x-ray computed tomography and cryo-SEM. Results For all species studied, the main components of water storage within the operating range of water potential were not living cells but cavitation release and capillaries. Abies firma maintained water in the earlywood-like cells, for possible maintenance of the transpiration stream. Cercidiphyllum japonicum maintained water in its vessels over 200 kg m-3 of CWR, while Q. serrata lost most of its water in vessels with increasing CWR up to 100 kg m-3 . Cercidiphyllum japonicum exhibited a higher water storage capacity than Q. serrata. Under high CWR, narrow conduits stored xylem water in C. japonicum and imperforate tracheary elements in Q. serrata. Conclusions Among the species examined, increasing CWR appears to indicate differential utilization of stored water in relation to variation of xylem structure, thereby providing insight into the interspecific responses of tree species to drought.

The present study was carried out to investigate the effect of exogenous amino acids (AAs) on leaf apoplastic retention and symplastic absorption, leaf distribution, translocation, and nutritional status of manganese (Mn) in cucumber plants. Results showed that application of AAs increased the symplastic to apoplastic Mn proportion in leaves of cucumber. Furthermore, a greater part of Mn applied with AAs accumulated in the internal area of newly growing leaves suggesting the transport of Mn mostly in the phloem in the presence of AAs. In contrast, Mn preferentially accumulated in the marginal area of treated leaves indicating the Mn movement in xylem according to the transpiration stream. The AAs used in the present study also increased the relative translocation of Mn from treated leaves to the upper leaves and roots. More time was needed for long-distance translocation of Mn from treated leaves to the upper leaves than that to the roots. Application of AAs mainly increased the transport of Mn in the phloem. The leaf PO (peroxidase) activity was the better marker to index the nutritional status of Mn than PPO (polyphenoloxidase) activity. In conclusion, exogenous AAs especially lysine (Lys) promoted the mobility and bioavailability of applied Mn as foliar spray.

Silicon (Si) has never been acknowledged as a vital nutrient though it confers a crucial role in a variety of plants. Si may usually be expressed more clearly in Si-accumulating plants subjected to biotic stress. It safeguards several plant species from disease. It is considered as a common element in the lithosphere of up to 30% of soils, with most minerals and rocks containing silicon, and is classified as a “significant non-essential” element for plants. Plant roots absorb Si, which is subsequently transferred to the aboveground parts through transpiration stream. The soluble Si in cytosol activates metabolic processes that create jasmonic acid and herbivore-induced organic compounds in plants to extend their defense against biotic stressors. The soluble Si in the plant tissues also attracts natural predators and parasitoids during pest infestation to boost biological control, and it acts as a natural insect repellent. However, so far scientists, policymakers, and farmers have paid little attention to its usage as a pesticide. The recent developments in the era of genomics and metabolomics have opened a new window of knowledge in designing molecular strategies integrated with the role of Si in stress mitigation in plants. Accordingly, the present review summarizes the current status of Si-mediated plant defense against insect, fungal, and bacterial attacks. It was noted that the Si-application quenches biotic stress on a long-term basis, which could be beneficial for ecologically integrated strategy instead of using pesticides in the near future for crop improvement and to enhance productivity.

Silica, a mineral reducing the digestibility of grass tissues, is assumed to form spontaneously in planta. We overturn this model, showing that silica deposition occurs during lignification and demands metabolic energy., Silica deposition in plants is a common phenomenon that correlates with plant tolerance to various stresses. Deposition occurs mostly in cell walls, but its mechanism is unclear. Here we show that metabolic processes control the formation of silica aggregates in roots of sorghum (Sorghum bicolor L.), a model plant for silicification. Silica formation was followed in intact roots and root segments of seedlings. Root segments were treated to enhance or suppress cell wall biosynthesis. The composition of endodermal cell walls was analysed by Raman microspectroscopy, scanning electron microscopy and energy-dispersive X-ray analysis. Our results were compared with in vitro reactions simulating lignin and silica polymerization. Silica aggregates formed only in live endodermal cells that were metabolically active. Silicic acid was deposited in vitro as silica onto freshly polymerized coniferyl alcohol, simulating G-lignin, but not onto coniferyl alcohol or ferulic acid monomers. Our results show that root silica aggregates form under tight regulation by endodermal cells, independently of the transpiration stream. We raise the hypothesis that the location and extent of silicification are primed by the chemistry and structure of polymerizing lignin as it cross-links to the wall.

Working in tandem with root exclusion, stems may provide salt-tolerant woody perennials with some additional capacity to restrict sodium (Na) and chloride (Cl) accumulation in leaves. The Pistacia genus, falling at the nexus of salt tolerance and human intervention, provided an ideal set of organisms for studying the influences of both variable root exclusion and potentially variable discontinuities at the bud union on stem processes. In three experiments covering a wide range of salt concentrations (0 to 150 mM NaCl) and tree ages (1, 2 and 10 years) as well as nine rootstock-scion combinations we show that proportional exclusion of both Na and Cl reached up to ~85% efficacy, but efficacy varied by both rootstock and budding treatment. Effective Na exclusion was augmented by significant retrieval of Na from the xylem sap, as evidenced by declines in the Na concentrations of both sap and wood tissue along the transpiration stream. However, while we observed little to no differences between the concentrations of the two ions in leaves, analogous declines in sap concentrations of Cl were not observed. We conclude that some parallel but separate mechanism must be acting on Cl to provide leaf protection from toxicity specific to this ion and suggest that this mechanism is recirculation of Cl in the phloem. The presented findings underline the importance of holistic assessments of salt tolerance in woody perennials. In particular, greater emphasis might be placed on the dynamics of salt sequestration in the significant storage volumes offered by the stems of woody perennials and on the potential for phloem discontinuity introduced with a bud/graft union.

International audience; The ability to transport water through tall stems hydraulically limits stomatal conductance (gs), thereby constraining photosynthesis and growth. However, some plants are able to minimize this height‐related decrease in gs, regardless of path length. We hypothesized that kudzu (Pueraria lobata) prevents strong declines in gs with height through appreciable structural and hydraulic compensative alterations. We observed only a 12% decline in maximum gs along 15‐m‐long stems and were able to model this empirical trend. Increasing resistance with transport distance was not compensated by increasing sapwood‐to‐leaf‐area ratio. Compensating for increasing leaf area by adjusting the driving force would require water potential reaching −1.9 MPa, far below the wilting point (−1.2 MPa). The negative effect of stem length was compensated for by decreasing petiole hydraulic resistance and by increasing stem sapwood area and water storage, with capacitive discharge representing 8–12% of the water flux. In addition, large lateral (petiole, leaves) relative to axial hydraulic resistance helped improve water flow distribution to top leaves. These results indicate that gs of distal leaves can be similar to that of basal leaves, provided that resistance is highest in petioles, and sufficient amounts of water storage can be used to subsidize the transpiration stream.

Transpiration is a function of plants to evaporate water from their leaf stomata, which enables plants to take up ground water from their roots to their leaves through the xylem. This water flux fu...

The increasing load of organophosphate flame retardants (OPFRs) has generated wide concerns about their potential residues in aquatic environments. The uptake and translocation of fourteen OPFRs by wheat (Triticum aestivum L.) were studied under hydroponic conditions. The results revealed that OPFRs were removed from hydroponic solution by wheat, and the removal processes followed first-order kinetics. After 10 days, the removal efficiencies were in a range of 57.9 ± 3.8%−63.8 ± 5.6%. The potential for translocation of these OPFRs from the roots to foliage was also assessed. OPFRs with relatively higher hydrophobicity were more likely taken up by roots, and OPFRs with lower hydrophobicity were more prone to be translocated. Root concentration factors (RCFs), transpiration stream concentration factors (TSCFs), and foliage/root concentration factors (FRCFs) were calculated. Furthermore, significant correlations were found between RCF, FRCF or TSCF values of OPFRs and log Kow (p

Striga parasitizes major crops in arid regions, depriving the host crop of nutrients through the transpiration stream and causing vast agricultural damage. Here, we report on the mechanism underlying how Striga maintains high transpiration under drought conditions. We found that Striga did not respond to abscisic acid, the phytohormone responsible for controlling stomatal closure. Protein phosphatase 2C of Striga (ShPP2C1) is not regulated by abscisic acid receptors, and this feature is attributable to specific mutations in its amino acid sequence. Moreover, Arabidopsis transformed with ShPP2C1 showed an abscisic acid-insensitive phenotype, indicating that ShPP2C1 functions as a dominant negative regulator of abscisic acid signal transduction. These findings suggest that ShPP2C1 interrupts abscisic acid signalling in Striga, resulting in high transpiration and subsequent efficient absorption of host nutrients under drought conditions.

Respired CO2 in woody tissues radially diffuses to the atmosphere or it is transported upward with the transpiration stream, making the origin of CO2 in stem CO2 efflux (EA) uncertain, which may confound stem respiration (RS) estimates. An aqueous 13C-enriched solution was infused into stems of Populus tremula L. trees, and real-time measurements of 13C-CO2 and 12C-CO2 in EA were performed via Cavity Ring Down Laser Spectroscopy (CRDS). The contribution of locally respired CO2 (LCO2) and xylem-transported CO2 (TCO2) to EA was estimated from their different isotopic composition. Mean daily values of TCO2/EA ranged from 13% to 38%, evidencing the notable role that xylem CO2 transport plays in the assessment of stem respiration. Mean daily TCO2/EA did not differ between treatments of drought stress and light exclusion of woody tissues, but they showed different TCO2/EA dynamics on a sub-daily time scale. Sub-daily CO2 diffusion patterns were explained by a light-induced axial CO2 gradient ascribed to woody tissue photosynthesis, and the resistance to radial CO2 diffusion determined by bark water content. Here, we demonstrate the outstanding potential of CRDS paired with 13C-CO2 labelling to advance in the understanding of CO2 movement at the plant-atmosphere interface and the respiratory physiology in woody tissues.

Microplastic (MP, 100 nm−5 mm) may present an attributable risk to ecosystem and human health, and its pollution has become a global environmental concern. Despite a wealth of information on the accumulation of MPs in aquatic species, there is no information on the uptake and accumulation of MPs by higher plants. Terrestrial edible plants are directly exposed to MPs when agricultural soil was applied with organic manure, sewage sludge as fertilizer or plastic mulching. In this paper, the uptake of two sizes of polystyrene (PS) microbeads (0.2 and 1.0 μm) and then their distribution and migration in an edible plant lettuce were firstly investigated based on laboratory experiments. We used fluorescent markers to track PS microbeads in plant tissues and found fluorescence to be a sensitive and reliable detection method. Sections from untreated control lettuce showed no autofluorescence. When roots were treated with fluorescently labeled PS microbeads, the microbeads could be identified by its fluorescence. Our main study investigated the uptake of 0.2 μm beads, as few luminescence signals were observed in lettuce roots for 1.0 μm beads in our experiment. We observed that 0.2 μm fluorescent microbeads were extracellularly trapped in the root cap mucilage (which is a highly hydrated polysaccharide) and a “dark green tip” (which was typical of lettuce roots exposed to label PS beads) was usually visible to the naked eye. Confocal images revealed that the PS luminescence signals were mainly located in the vascular system and on the cell walls of the cortex tissue of the roots, indicated that the beads passed through the intercellular space via the apoplastic transport system. Once inside the central cylinder, the 0.2 μm PS beads were transferred from the roots to the stems and leaves via the vascular system following the transpiration stream. We also observed that the PS beads adhered to one another and self-assembled systematically into “grape-like” and “(chain) string-like” clusters in the intercellular space of the root and stem vascular tissue of lettuce plant. In contrast to the root and stem, PS beads were dispersed in the leaf tissue. Here, for the first time we provide evidence of the adherence, uptake, accumulation, and translocation of submicrometer MPs within an edible plant. Our findings highlight the previously underappreciated human exposure pathway to MPs through the consumption of contaminated crops and emphasize the need for new management strategies to control the release of MPs waste products into the terrestrial environment. Ultimately, the potential impacts of low range sized MPs on food safety of crop plants and human health need to be urgently considered.

We cultivated Brassica rapa var. perviridis in soil mixed with four pesticides (fenobucarb, procymidone, flutolanil, and tolclofos-methyl) at different temperatures, day lengths, and soil water contents. We compared plants' uptake and translocation abilities of the pesticides as affected by growth conditions. The root concentration factor (RCF) of pesticides tended to increase with rising temperature; however, but the influence of temperature on the transpiration stream concentration factor (TSCF) differed for each pesticide. The RCFs and TSCFs of pesticides were high for short days. The soil water content had little or no effect on the uptake and translocation of pesticides. These results showed that it is necessary to consider growth conditions, especially the temperature and day length in plant uptake models for these pesticides.

summary Pathways for water and salts were traced from the vein endings to their sites of exit in leaves of Avicennia marina (Forsk.) Vierh. Investigations were conducted to determine whether the water and solutes separate at any point in the transpiration stream, and to identify their subsequent pathways. Leaves were placed with their cut petioles in solution containing one of two fluorochromes chosen to trace the transpiration stream. The apoplastic tracer was B-hydroxypyrene-1, 3, 6-trisulphonic acid (PTS), and the symplastic tracer was riboflavin buffered to pH 6.8. Leaf pieces were freeze-substituted with special care to keep the specimens dry to prevent redistribution of the water-soluble fluorochromes. Stages of the technique were sensitive to atmospheric moisture, so sections were cut in a room dried to 30% relative humidity, and they were stored in a sealed box containing desiccant. Riboflavin was never seen in the apoplast, and was inside the abaxtal (lower) glands within 30 min indicating a low resistance pathway through the symplast to the gland. The veins terminate in heavily pitted xylem vessels, and at the ends of these pits, the junction between the vein ending and the apoplast of the surrounding mesophyll, were high concentrations of PTS indicating regions of separation of the fluorochrome from water – places where the water crosses from the apoplast to the symplast. PTS diffused slowly in cell walls (measured diffusivity: 4.7 × 10−10 cm2 s−1 reaching the abaxial epidermis after 8 h, which was sufficient time for four complete changes of leaf water. It was concluded that the transpiration stream, probably containing the salt destined for secretion, moved symplastically from the xylem vein endings.

Rates of ion transport and transpiration were measured during the day and night in whole seedlings of Suaeda maritima growing over a range of salinities, in order to calculate concentrations of sodium and potassium in the xylem during these periods. Mean sodium concentration in the xylem was maximal at 56 mol m−3 Na with an external salinity of 200 mol m−3 NaCl. The sodium concentration in the xylem was greater in the dark than in the light at all external salinities investigated. Comparison of the external sodium with that in the xylem indicated that sodium was more strongly excluded from the transpiration stream as salinity increased. The mean concentration of potassium in the xylem declined as external NaCl concentration increased, although selectivity for potassium increased at higher salinities. Results are discussed in relation to osmotic adjustment in S. maritima.

Our objective was to establish whether the xylem vascular connections (architecture) of a genotype of Trifolium repens L. had implications for the intraplant allocation of recently assimilated phosphorus (P). One nodal root of each plan! was isolated and fed 32 P-labelled nutrient solution for 24 h. The fed root was either on the parent axis (11-12 nodes proximal CO the apex) or on a branch at the third or fourth node along the branch when counting from the junction with the parent stolon. Allocation patterns were obtained by dissecting plants and assaying each dissected component for 32 P. Under conditions of undisturbed growth in a controlled environment xylem architecture was found to have an important influence on the intraplant distribution of 32 P from nodal roots. Allocation patterns of 32 P were consistent with those predicted from knowledge of the xylem vascular architecture of the genotype and a predominantly acropetal direction of the transpiration stream. For instance, very little 32 P (< 1 % of exported 32 P from the fed root) was found in branches on the opposite side of the stolon to the fed root, and the strong acropetal direction of the transpiration stream resulted in little allocation of 32 P to leaves that had traces that arose from axial bundles at positions proximal to that of the fed root (the leaf subtending the fed root and the next distal leaf had low 32 P content), to all plant organs proximal to the fed root (< 6%) and to all other roots (

summary Cuticular water permeance was manipulated in Corylus avellana L., Hypericum androsaemum L. and Populus tremula L. by (1) long-term application of low doses of various systemic herbicides inhibiting biosynthesis of cuticular waxes, (2) very short-term application of organic solvents to the leaf surface, and (3) exposure to natural strong winds. Treatment effects were very variable, but increased the natural range of permeances by a factor of 10 or so in undamaged leaves. All species had hypostomatous leaves. Relative change of leaf conductance (g) in response to stepwise increases of leaf-to-air water vapour pressure difference (VPD) was measured for individual leaves (Corylus) or groups of leaves at the shoot or branch tip. Adaxial cuticular water permeance (P) was determined for the same leaves after measurement of the VPD-response. A proportional measure of relative change of g with VPD, d(log e.g)dVPD, was then plotted against P. No increase in the strength of the closing response to increasing VPD was found with increasing P, as would have been expected if water loss through the cuticle was involved in stomatal response to changes in VPD via a direct effect on guard cell turgor. By contrast, high P coincided, most clearly in Corylus, with a reduced strength of the stomatal closing response to increasing VPD, i.e. less negative d(logeg)dVPD. As the responses were non-linear, the value of d(logeg)dVPD changed with VPD. With rising VPD, all three species and a fourth one previously studied showed a decline in the value of [d(logeg)dVPD]/d(log P), reaching negative values in one species. This is interpreted in terms of two independent and antagonistic effects of increased cuticular water permeance on guard cell response to VPD, one acting by reducing the backpressure exerted on guard cells by the epidermis, and the other one possibly causing greater depression of guard cell turgor through delivery of more chemical messengers (such as abscisic acid) to the guard cells with the cuticular transpiration stream.

Boron (B) is an essential nutrient for plant growth and development, exhibiting extremely narrow margins between deficiency and toxicity. B toxicity is devastating for productivity and apparent for a continuously increasing part of agricultural land, under the influence of on-going climate change. In this study, the effects of increased B supply (by using H3BO3) were addressed by examining critical physiological responses of young and mature leaves, which were devoid of toxicity symptoms, in two melon varieties (Armenian cucumbers, cantaloupes). B was primarily translocated through the transpiration stream, and secondarily via the active cell membrane transport system. The B distribution pattern was independent of leaf age, and remained rather unchanged under increased B supply. Armenian cucumbers, exhibiting higher leaf B levels, underwent an enhanced adverse impact on (root and shoot) growth, photosynthetic pigment content, cellular membrane integrity, and also exhibited attenuated antioxidant defense stimulation. Notably, and unlike other abiotic stressors, no evidence of B toxicity-induced systemic reaction was apparent. B toxicity greatly enhanced the transcription of the genes coding for borate influx and efflux channels, an effect that was mostly evident in mature leaves. In conclusion, shoot physiological responses to B toxicity are highly localized. Moreover, the obstruction of the diffusion and the B translocation to the aerial organs under increased B supply is genotype-dependent, governing plant physiological responses.

The capacity of trees to release water from storage compartments into the transpiration stream can mitigate damage to hydraulic functioning. However, the location of these ‘transient’ water sources and also the pathways of water movement other than vertical through tree stems still remain poorly understood. We conducted an experiment on two tree species in a common garden in eastern Australia that naturally grow in regions of high (Eucalyptus tereticornis, ‘Red Gum’) and low (Eucalyptus sideroxylon, ‘Ironbark’) annual precipitation rates. Deuterium-enriched water (1350% label strength) was directly introduced into the transpiration stream of three trees per species for four consecutive days. Subsequently, the trees were felled, woody tissue samples were collected from different heights and azimuthal positions of the stems, and stable isotope ratios were determined on the water extracted from all samples. The presence/absence of the tracer along the radial and vertical stem axes in combination with xylem hydraulic properties inferred from sapflow, leaf and stem water potentials, wood moisture contents and anatomical sapwood characteristics elucidated species-specific patterns of short-term stem water storage and movement. The distribution of water isotopes at natural abundance among woody tissues indicated systematic differences with highest values of sapwood water and lower values in inner bark and heartwood. Presence of tracer in water of the inner bark highlighted the importance of this tissue as capacitor. Although injected at the northern side of stems, tracer was also discovered at the southern side, providing empirical evidence for circumferential flow in sapwood, particularly of Ironbark. Greater vertical water transport in Red Gum compared with more radial and circumferential water transport in Ironbark were associated with species-specific sapwood anatomy. Our study highlights the value of combining information from stable isotope tracers and wood anatomy to investigate patterns of water transport and storage of tall trees in situ.

Uptake of seven organic contaminants including bisphenol A, estriol, 2,4-dinitrotoluene, N,N-diethyl-meta-toluamide (DEET), carbamazepine, acetaminophen, and lincomycin by tomato (Solanum lycopersicum L.), corn (Zea mays L.), and wheat (Triticum aestivum L.) was measured. The plants were grown in a growth chamber under recommended conditions and dosed by these chemicals for 19 days. The plant samples (stem transpiration stream) and solution in the exposure media were taken to measure transpiration stream concentration factor (TSCF). The plant samples were analyzed by a freeze-thaw centrifugation technique followed by high performance liquid chromatography-tandem mass spectrometry detection. Measured average TSCF values were used to test a neural network (NN) model previously developed for predicting plant uptake based on physicochemical properties. The results indicated that moderately hydrophobic compounds including carbamazepine and lincomycin have average TSCF values of 0.43 and 0.79, respectively. The average uptake of DEET, estriol, acetaminophen, and bisphenol A was also measured as 0.34, 0.29, 0.22, and 0.1, respectively. The 2,4-dinitrotoluene was not detected in the stem transpiration stream and it was shown to degrade in the root zone. Based on these results together with plant physiology measurements, we concluded that physicochemical properties of the chemicals did predict uptake, however, the role of other factors should be considered in the prediction of TSCF. While NN model could predict TSCF based on physicochemical properties with acceptable accuracies (mean squared error less than 0.25), the results for 2,4-dinitrotoluene and other compounds confirm the needs for considering other parameters related to both chemicals (stability) and plant species (role of lipids, lignin, and cellulose).