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Rice is an important source of calories and mineral nutrients for more than half of the world's population. The accumulation of essential and toxic mineral elements in rice grain affects its nutritional quality and safety. However, the patterns and processes by which different elements progressively accumulate during grain filling remain largely unknown. In the present study, we investigated temporal changes in dry matter, elemental concentrations, and the transcriptome in the grain of field-grown rice. We also investigated the effects of seed setting rate and the position of the grain within the rice panicle on element accumulation. Three different patterns of accumulation were observed: (i) elements including K, Mn, B, and Ca showed an early accumulation pattern; (ii) dry matter and elements including N, P, S, Mg, Cu, Zn, Mo, As, and Cd showed a mid accumulation pattern; and (iii) elements such as Fe showed a gradual increase pattern. These different accumulation patterns can be explained by the differences in the biogeochemical behavior of the various elements in the soil, as well as differences in plant nutrient redistribution, gene expression, and the sink–source relationship. These results improve our knowledge of the dynamics of elemental accumulation in rice grain and are helpful for identification of functional genes mediating the translocation of elements to grain. [ABSTRACT FROM AUTHOR]

Trichomes are potentially important for absorption of foliar fertilizers. A study has shown that the non-glandular trichromes (NGTs) of sunflower (Helianthus annuus) accumulated high concentrations of foliar-applied zinc (Zn); however, the mechanisms of Zn accumulation in the NGTs and the fate of this Zn are unclear. Here we investigated how foliar-applied Zn accumulates in the NGTs and the subsequent translocation of this Zn. Time-resolved synchrotron-based X-ray fluorescence microscopy and transcriptional analyses were used to probe the movement of Zn in the NGTs, with the cuticle composition of the NGTs examined using confocal Raman microscopy. The accumulation of Zn in the NGTs is both an initial preferential absorption process and a subsequent translocation process. This preferred absorption is likely because the NGT base has a higher hydrophilicity, whilst the subsequent translocation is due to the presence of plasmodesmata, Zn-chelating ligands, and Zn transporters in the NGTs. Furthermore, the Zn sequestered in the NGTs was eventually translocated out of the trichome once the leaf Zn concentration had decreased, suggesting that the NGTs are also important in maintaining leaf Zn homeostasis. This study demonstrates for the first time that trichomes have a key structural and functional role in the absorption and translocation of foliar-applied Zn. [ABSTRACT FROM AUTHOR]

Background and Aims Synchrotron- and laboratory-based micro-X-ray fluorescence (µ-XRF) is a powerful technique to quantify the distribution of elements in physically large intact samples, including live plants, at room temperature and atmospheric pressure. However, analysis of light elements with atomic number (Z) less than that of phosphorus is challenging due to the need for a vacuum, which of course is not compatible with live plant material, or the availability of a helium environment. Method A new laboratory µ-XRF instrument was used to examine the effects of silicon (Si) on the manganese (Mn) status of soybean (Glycine max) and sunflower (Helianthus annuus) grown at elevated Mn in solution. The use of a helium environment allowed for highly sensitive detection of both Si and Mn to determine their distribution. Key Results The µ-XRF analysis revealed that when Si was added to the nutrient solution, the Si also accumulated in the base of the trichomes, being co-located with the Mn and reducing the darkening of the trichomes. The addition of Si did not reduce the concentrations of Mn in accumulations despite seeming to reduce its adverse effects. Conclusions The ability to gain information on the dynamics of the metallome or ionome within living plants or excised hydrated tissues can offer valuable insights into their ecophysiology, and laboratory µ-XRF is likely to become available to more plant scientists for use in their research. [ABSTRACT FROM AUTHOR]

Background and Aims Signal grass (Urochloa decumbens) is a widely used pasture grass in tropical and sub-tropical areas due to its high aluminiun (Al) resistance. However, the underlying mechanisms conferring this resistance are not clearly understood. Methods The Al concentrations of bulk root tissues and the intracellular compartment were examined, including the impact of a metabolic inhibitor, carbonyl cyanide m -chlorophenyl hydrazone (CCCP). Next, we examined changes in the properties of signal grass root tissues following exposure to toxic levels of Al, including the cell wall cation exchange capacity (CEC), degree of methylation and concentrations of cell wall fractions. Key Results Although signal grass was highly resistant to Al, there was a delay of 24–48 h before the expression of this resistance. We found that this delay in the expression of Al resistance was not related to the total Al concentration in the bulk apical root tissues, nor was it related to changes in the Al bound to the cell wall. We also examined changes in other properties of the cell wall, including the CEC, degree of methylation and changes in the concentration of pectin, hemicellulose and cellulose. We noted that concentrations of intracellular Al decreased by approx. 50 % at the same time that the root elongation rate improved after 24–48 h. Using CCCP as a metabolic inhibitor, we found that the intracellular Al concentration increased approx. 14-fold and that the CCCP prevented the subsequent decrease in intracellular Al. Conclusions Our results indicate that the delayed expression of Al resistance was not associated with the Al concentration in the bulk apical root tissues or bound to the cell wall, nor was it associated with changes in other properties of the cell wall. Rather, signal grass has an energy-dependent Al exclusion mechanism, and this mechanism requires 24–48 h to exclude Al from the intracellular compartment. [ABSTRACT FROM AUTHOR]

Understanding the distribution of elements in plants is important for researchers across a broad range of fields, including plant molecular biology, agronomy, plant physiology, plant nutrition, and ionomics. However, it is often challenging to evaluate the applicability of the wide range of techniques available, with each having its own strengths and limitations. Here, we compare scanning/transmission electron microscopy-based energy-dispersive x-ray spectroscopy, x-ray fluorescence microscopy, particle-induced x-ray emission, laser ablation inductively coupled plasma-mass spectrometry, nanoscale secondary ion mass spectroscopy, autoradiography, and confocal microscopy with fluorophores. For these various techniques, we compare their accessibility, their ability to analyze hydrated tissues (without sample preparation) and suitability for in vivo analyses, as well as examining their most important analytical merits, such as resolution, sensitivity, depth of analysis, and the range of elements that can be analyzed. We hope that this information will assist other researchers to select, access, and evaluate the approach that is most useful in their particular research program or application. [ABSTRACT FROM AUTHOR]

Background and Aims X-ray fluorescence microscopy (XFM) is a powerful technique to elucidate the distribution of elements within plants. However, accumulated radiation exposure during analysis can lead to structural damage and experimental artefacts including elemental redistribution. To date, acceptable dose limits have not been systematically established for hydrated plant specimens. Methods Here we systematically explore acceptable dose rate limits for investigating fresh sunflower (Helianthus annuus) leaf and root samples and investigate the time–dose damage in leaves attached to live plants. Key Results We find that dose limits in fresh roots and leaves are comparatively low (4.1 kGy), based on localized disintegration of structures and element-specific redistribution. In contrast, frozen-hydrated samples did not incur any apparent damage even at doses as high as 587 kGy. Furthermore, we find that for living plants subjected to XFM measurement in vivo and grown for a further 9 d before being reimaged with XFM, the leaves display elemental redistribution at doses as low as 0.9 kGy and they continue to develop bleaching and necrosis in the days after exposure. Conclusions The suggested radiation dose limits for studies using XFM to examine plants are important for the increasing number of plant scientists undertaking multidimensional measurements such as tomography and repeated imaging using XFM. [ABSTRACT FROM AUTHOR]

Background and Aims Understanding the speciation of Zn in edible portions of crops helps identify the most effective biofortification strategies to increase the supply of nutrients for improving the health and nutrition of consumers. Methods Kernels of 12 sweetcorn and three maize (Zea mays) varieties were analysed for Zn concentration and content. The speciation of the Zn in the embryos, endosperms and whole kernels at 21, 28 and 56 days after pollination (DAP) was then examined for one maize and one sweetcorn variety using synchrotron-based X-ray absorption spectroscopy (XAS). Key Results Averaged across all sweetcorn and maize varieties at 21 DAP, the embryo contributed 27–29% of the whole kernel Zn whilst the endosperm contributed 71–73 %. While sweetcorn embryos contributed a lower proportion to the total kernel Zn than those of maize, the proportion of total Zn in the embryo increased as kernels aged for both varieties, reaching 33 % for sweetcorn and 49% for maize at 28 DAP. Using XAS, it was predicted that an average of 90 % of the Zn in the embryos was present as Zn-phytate, while in the endosperm the Zn was primarily complexed with an N-containing ligand such as histidine and to a lesser extent with phytate. However, in maize endosperms, it was also observed that the proportion of Zn present as Zn-phytate increased as the kernel matured, thereby also probably decreasing its bioavailability in these mature maize kernels. Conclusions The apparent low bioavailability of Zn supplied in maize at its consumption stage (i.e. mature kernels) probably undermines the effectiveness of biofortification of this crop. Conversely, successful biofortification of Zn in sweetcorn and green maize consumed as immature kernels could potentially provide a good source of bioavailable Zn in human diets. [ABSTRACT FROM AUTHOR]

Background and Aims High exchangeable sodium percentage (ESP) and bulk density of sodic soils can reduce seedling emergence. This study examined variation in seedling coleoptile length and seminal root angle of wheat (Triticum aestivum. L) genotypes to determine whether these traits vary between genotypes that differ in their tolerance to sodic soils. Methods Wheat genotypes were grown in three different experiments. First, four wheat genotypes were grown using soils of three ESPs (4, 10 and 17 %) and secondly in soils of three different bulk densities (1.2, 1.4 and 1.5 g cm–3) and ESP 10 %. Thirdly, seedling coleoptile length and seminal root angle were determined for 16 genotypes grown in a soil of ESP 10 % and bulk density 1.2 g cm–2. Seminal root angle and coleoptile length measurements from the current study were compared with seedling emergence rate and force measured previously. Key Results The seedling coleoptile length of all genotypes decreased with increasing soil ESP and bulk density, but with no significant differences between genotypes. In contrast, seminal root angles differed significantly between genotypes, but were not significantly affected by ESP or bulk density. There was an inverse relationship between the seminal root angle of the 16 genotypes and seedling emergence rate (R 2 = 0.89) and also between seminal root angle and seedling emergence force (R 2 = 0.61). Conclusions Lack of significant variation in coleoptile length between genotypes suggests that this may not be a suitable characteristic to identify wheat tolerance to sodic conditions. However, a narrower seminal root angle was correlated with rate and force of seedling emergence, traits likely to improve establishment. The mechanism underlying this correlation is not yet clear. Genotypes with a narrow root angle had greater root depth. One possible mechanism might be that genotypes with narrow root angles were able to take up more soil moisture at depth, leading to a higher proportion of seedling emergence. [ABSTRACT FROM AUTHOR]

Background and Aims Understanding the spatial distribution of inorganic nutrients within edible parts of plant products helps biofortification efforts to identify and focus on specific uptake pathways and storage mechanisms. Methods Kernels of sweetcorn (Zea mays) variety 'High zeaxanthin 103146' and maize inbred line 'Thai Floury 2' were harvested at two different maturity stages, and the distributions of K, P, S, Ca, Zn, Fe and Mn were examined in situ using synchrotron-based X-ray fluorescence microscopy. Key Results The distribution of inorganic nutrients was largely similar between maize and sweetcorn, but differed markedly depending upon the maturity stage after further embryonic development. The micronutrients Zn, Fe and Mn accumulated primarily in the scutellum of the embryo during early kernel development, while trace amounts of these were found in the aleurone layer at the mature stage. Although P accumulated in the scutellum, there was no direct relationship between the concentrations of P and those of the micronutrients, compared with the linear trend between Zn and Fe concentrations. Conclusions This study highlights the important role of the embryo as a micronutrient reserve for sweetcorn and maize kernels, and the need to understand how biofortification efforts can further increase the inorganic nutrient concentration of the embryo for human consumption. [ABSTRACT FROM AUTHOR]

Background and Aims Salt has been shown to affect Cd translocation and accumulation in plants but the associated mechanisms are unclear. This study examined the effects of salt type and concentration on Cd uptake, translocation and accumulation in Carpobrotus rossii. Methods Plants were grown in nutrient solution with the same Cd concentration or Cd2+ activity in the presence of 25 m m NaNO3, 12.5 m m Na2SO4 or 25 m m NaCl for ≤10 d. Plant growth and Cd uptake were measured and the accumulation of peptides and organic acids, and Cd speciation in plant tissues were analysed. Key Results Salt addition decreased shoot Cd accumulation by >50 % due to decreased root-to-shoot translocation, irrespective of salt type. Synchrotron-based X-ray absorption spectroscopy revealed that, after 10 d, 61–94 % Cd was bound to S-containing ligands (Cd–S) in both roots and shoots, but its speciation was not affected by salt. In contrast, Cd in the xylem sap was present either as free Cd2+ or complexes with carboxyl groups (Cd–OH). When plants were exposed to Cd for ≤24 h, 70 % of the Cd in the roots was present as Cd–OH rather than Cd–S. However, NaCl addition decreased the proportion of Cd–OH in the roots within 24 h by forming Cd–Cl complexes and increasing the proportion of Cd–S. This increase in Cd–S complexes by salt was not due to changes in glutathione and phytochelatin synthesis. Conclusions Salt addition decreased shoot Cd accumulation by decreasing Cd root-to-shoot translocation due to the rapid formation of Cd–S complexes (low mobility) within the root, without changing the concentrations of glutathione and phytochelatins. [ABSTRACT FROM AUTHOR]

Background and Aims The pathways whereby foliar-applied nutrients move across the leaf surface remain unclear. The aim of the present study was to examine the pathways by which foliar-applied Zn moves across the sunflower (Helianthus annuus) leaf surface, considering the potential importance of the cuticle, stomata and trichomes. Methods Using synchrotron-based X-ray florescence microscopy and nanoscale secondary ion mass spectrometry (NanoSIMS), the absorption of foliar-applied ZnSO4 and nano-ZnO were studied in sunflower. The speciation of Zn was also examined using synchrotron-based X-ray absorption spectroscopy. Key Results Non-glandular trichomes (NGTs) were particularly important for foliar Zn absorption, with Zn preferentially accumulating within trichomes in ≤15 min. The cuticle was also found to have a role, with Zn appearing to move across the cuticle before accumulating in the walls of the epidermal cells. After 6 h, the total Zn that accumulated in the NGTs was approx. 1.9 times higher than in the cuticular tissues. No marked accumulation of Zn was found within the stomatal cavity, probably indicating a limited contribution of the stomatal pathway. Once absorbed, the Zn accumulated in the walls of the epidermal and the vascular cells, and trichome bases of both leaf sides, with the bundle sheath extensions that connected to the trichomes seemingly facilitating this translocation. Finally, the absorption of nano-ZnO was substantially lower than for ZnSO4, with Zn probably moving across the leaf surface as soluble Zn rather than nanoparticles. Conclusions In sunflower, both the trichomes and cuticle appear to be important for foliar Zn absorption. [ABSTRACT FROM AUTHOR]

Foliar application of zinc (Zn) to crops is an effective way to increase the grain concentration of Zn. However, the development of more efficient foliar Zn fertilizers is limited by a lack of knowledge regarding the distribution, mobility, and speciation of Zn in leaves once it is taken up by the plant. We performed an experiment using radiolabelled Zn (65Zn), and in situ time-resolved elemental imaging using synchrotron X-ray fluorescence microscopy (XFM), to investigate the behaviour of two commonly used Zn foliar fertilizers (Zn sulphate and ZnEDTA) in wheat (Triticum aestivum) leaves. Both experiments showed that Zn had limited mobility in leaves, moving <25 mm from the application point after 24 h. Although limited, the translocation of Zn occurred quickly for both treatments; moving more between 3 h and 12 h after application than between 12 h and 24 h. Speciation analysis using synchrotron-based X-ray absorption near-edge structure (XANES) showed that ZnEDTA was in fact taken up in chelated form and not as ionic Zn (Zn2+). The XANES data also showed that Zn, from both treatments, was then complexed by ligands in the leaf (e.g. phytate and citrate), potentially in response to localized Zn toxicity. The results of the present study provide important insights into the behaviour of commonly used foliar-applied Zn fertilizers, and can be used to optimize current fertilization strategies and contribute to the development of more efficient foliar Zn fertilizers. [ABSTRACT FROM AUTHOR]

The present study investigated the role of trichomes in absorption of foliar-applied zinc fertilizers in soybean and tomato. Using synchrotron-based X-ray fluorescence microscopy for in situ analyses of hydrated leaves, we found that upon foliar application of ZnSO4, Zn accumulated within 15 min in some non-glandular trichomes in soybean, but not in tomato. However, analyses of cross-sections of soybean leaves did not show any marked accumulation of Zn in tissues surrounding trichomes. Furthermore, when near-isogenic lines of soybean differing 10-fold in trichome density were used to compare Zn absorption, it was found that foliar Zn absorption was not related to trichome density. Therefore, it is suggested that trichomes are not part of the primary pathway through which foliar-applied Zn moves across the leaf surface in soybean and tomato. However, this does not preclude trichomes being important in other plant species, as they are known to be highly diverse. We also compared the absorption of Zn when supplied as either ZnSO4, nano-ZnO, or bulk-ZnO, and found that absorption from ZnSO4 was about 10-fold higher than from nano- and bulk-ZnO, suggesting that it was mainly absorbed as soluble Zn. This study improves our understanding of the absorption of foliar-applied nutrients. [ABSTRACT FROM AUTHOR]

* Background and Aims Foliar fertilization to overcome nutritional deficiencies is becoming increasingly widespread. However, the processes of foliar nutrient absorption and translocation are poorly understood. The present study aimed to investigate how cuticular leaf properties affect the absorption of foliar-applied nutrients in leaf tissues. * Methods Given that methyl jasmonate (MeJA) can cause alterations in leaf properties, we applied 1 mm MeJA to sunflower (Helianthus annuus), tomato (Solanum lycopersicum) and soybean (Glycine max) to assess changes in leaf properties. Using traditionally analytical approaches and synchrotron-based X-ray fluorescence microscopy, the effects of these changes on the absorption and translocation of foliar-applied Zn, Mn and Fe were examined. * Key Results The changes in leaf properties caused by the application of MeJA increased foliar absorption of Zn, Mn and Fe up to 3- to 5-fold in sunflower but decreased it by 0⋅5- to 0⋅9-fold in tomato, with no effect in soybean. These changes in the foliar absorption of nutrients could not be explained by changes in overall trichome density, which increased in both sunflower (86 %) and tomato (76 %) (with no change in soybean). Similarly, the changes could be not attributed to changes in stomatal density or cuticle composition, given that these properties remained constant. Rather, the changes in the foliar absorption of Zn, Mn and Fe were related to the thickness of the cuticle and epidermal cell wall. Finally, the subsequent translocation of the absorbed nutrients within the leaf tissues was limited (<1⋅3 mm) irrespective of treatment. * Conclusions The present study highlights the potential importance of the combined thickness of the cuticle and epidermal cell wall in the absorption of foliar-applied nutrients. This information will assist in increasing the efficacy of foliar fertilization. [ABSTRACT FROM AUTHOR]

Nitrogen fertilization could improve the efficiency of Cd phytoextraction in contaminated soil and thus shorten the remediation time. However, limited information is available on the effect of N form on Cd phytoextraction and associated mechanisms in plants. This study examined the effect of N form on Cd accumulation, translocation, and speciation in Carpobrotus rossii and Solanum nigrum. Plants were grown in nutrient solution with 5-15 μM Cd in the presence of 1000 μM NH4+ or NO3-. Plant growth and Cd uptake were measured, and Cd speciation was analyzed using synchrotron-based X-ray absorption spectroscopy. Shoot Cd accumulation was 30% greater with NH4 + than NO3- supply. Carpobrotus rossii accumulated three times more Cd than S. nigrum. However, Cd speciation in the plants was not influenced by N form, but it did vary with species and tissues. In C. rossii, up to 91% of Cd was bound to S-containing ligands in all tissues except the xylem sap where 87-95% were Cd-OH complexes. Furthermore, the proportion of Cd-S in shoots was substantially lower in S. nigrum (44-69%) than in C. rossii (60-91%). It is concluded that the application of NH4+ (instead of NO3-) increased shoot Cd accumulation by increasing uptake and transloca tion, rather than changing Cd speciation, and is potentially an effective approach for increasing Cd phytoextraction. [ABSTRACT FROM AUTHOR]

Acid soils with elevated levels of soluble aluminium (Al) comprise ~40% of the world's arable land, but there remains much uncertainty regarding the mechanisms by which Al is rhizotoxic. This review examines the kinetics of the toxic effects of Al on the root elongation rate (RER), its effects on root tissues, and its location at a subcellular level. Depending upon the concentration and plant species, soluble Al decreases the RER in a median time of 73 min, but in as little as 5 min in soybean. This is initially due to a decreased rate at which cells expand anisotropically in the elongation zone. Thereafter, rhizodermal and outer cortical cells rupture through decreased cell wall relaxation. It is in this region where most Al accumulates in the apoplast. Subsequently, Al impacts root growth at a subcellular level through adverse effects on the plasma membrane (PM), cytoplasm, and nucleus. At the PM, Al alters permeability, fluidity, and integrity in as little as 0.5 h, whilst it also depolarizes the PM and reduces H+-ATPase activity. The Al potentially crosses the PM within 0.5 h where it is able to bind to the nucleus and inhibit cell division; sequestration within the vacuole is required to reduce the toxic effects of Al within the cytoplasm. This review demonstrates the increasing evidence of the importance of the initial Al-induced inhibition of wall loosening, but there is evidence also of the deleterious effects of Al on other cellular processes which are important for long-term root growth and function. [ABSTRACT FROM AUTHOR]

Plant species differ in response to high available manganese (Mn), but the mechanisms of sensitivity and tolerance are poorly understood. In solution culture, greater than or equal to 30 mM Mn decreased the growth of soybean (Glycine max), but white lupin (Lupinus albus), narrow-leafed lupin (Lupin angustifolius), and sunflower (Helianthus annuus) grew well at 100 mM Mn. Differences in species' tolerance to high Mn could not be explained simply by differences in root, stem, or leaf Mn status, being 8.6, 17.1, 6.8, and 9.5 mmol kg-1 leaf fresh mass at 100 mM Mn. Furthermore, x-ray absorption near edge structure analyses identified the predominance of Mn(II), bound mostly to malate or citrate, in roots and stems of all four species. Rather, differences in tolerance were due to variations in Mn distribution and speciation within leaves. In Mn-sensitive soybean, in situ analysis of fresh leaves using x-ray fluorescence microscopy combined with x-ray absorption near edge structure showed high Mn in the veins, and manganite [Mn(III)] accumulated in necrotic lesions apparently through low Mn sequestration in vacuoles or other vesicles. In the two lupin species, most Mn accumulated in vacuoles as either soluble Mn(II) malate or citrate. In sunflower, Mn was sequestered as manganite at the base of nonglandular trichomes. Hence, tolerance to high Mn was ascribed to effective sinks for Mn in leaves, as Mn(II) within vacuoles or through oxidation of Mn(II) to Mn(III) in trichomes. These two mechanisms prevented Mn accumulation in the cytoplasm and apoplast, thereby ensuring tolerance to high Mn in the root environment. [ABSTRACT FROM AUTHOR]

Knowledge of the distribution of selenium (Se) species within plant tissues will assist in understanding the mechanisms of Se uptake and translocation, but in situ analysis of fresh and highly hydrated plant tissues is challenging. Using synchrotron-based fluorescence X-ray absorption near-edge spectroscopy (XANES) imaging to provide laterally resolved data, the speciation of Se in fresh roots and leaves of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) supplied with 1 nM of either selenate or selenite was investigated. For plant roots exposed to selenate, the majority of the Se was efficiently converted to C-Se-C compounds (i.e. methylselenocysteine or selenomethionine) as selenate was transported radially through the root cylinder. Indeed, even in the rhizodermis which is exposed directly to the bulk solution, only 12-31% of the Se was present as uncomplexed selenate. The C-Se-C compounds were probably sequestered within the roots, whilst much of the remaining uncomplexed Se was translocated to the leaves--selenate accounting for 52-56% of the total Se in the leaves. In a similar manner, for plants exposed to selenite, the Se was efficiently converted to C-Se-C compounds within the roots, with only a small proportion of uncomplexed selenite observed within the outer root tissues. This resulted in a substantial decrease in translocation of Se from the roots to leaves of selenite-exposed plants. This study provides important information for understanding the mechanisms responsible for the uptake and subsequent transformation of Se in plants. [ABSTRACT FROM AUTHOR]

Despite the rhizotoxicity of aluminum (Al) being identified over 100 years ago, there is still no consensus regarding the mechanisms whereby root elongation rate is initially reduced in the approximately 40% of arable soils worldwide that are acidic. We used high-resolution kinematic analyses, molecular biology, rheology, and advanced imaging techniques to examine soybean (Glycine max) roots exposed to Al. Using this multidisciplinary approach, we have conclusively shown that the primary lesion of Al is apoplastic. In particular, it was found that 75 mM Al reduced root growth after only 5 min (or 30 min at 30 mM Al), with Al being toxic by binding to the walls of outer cells, which directly inhibited their loosening in the elongation zone. An alteration in the biosynthesis and distribution of ethylene and auxin was a second, slower effect, causing both a transient decrease in the rate of cell elongation after 1.5 h but also a longer term gradual reduction in the length of the elongation zone. These findings show the importance of focusing on traits related to cell wall composition as well as mechanisms involved in wall loosening to overcome the deleterious effects of soluble Al. [ABSTRACT FROM AUTHOR]

Background and Aims Globally, zinc deficiency is one of the most important nutritional factors limiting crop yield and quality. Despite widespread use of foliar-applied zinc fertilizers, much remains unknown regarding the movement of zinc from the foliar surface into the vascular structure for translocation into other tissues and the key factors affecting this diffusion.Methods Using synchrotron-based X-ray fluorescence microscopy (µ-XRF), absorption of foliar-applied zinc nitrate or zinc hydroxide nitrate was examined in fresh leaves of tomato (Solanum lycopersicum) and citrus (Citrus reticulatus).Key Results The foliar absorption of zinc increased concentrations in the underlying tissues by up to 600-fold in tomato but only up to 5-fold in citrus. The magnitude of this absorption was influenced by the form of zinc applied, the zinc status of the treated leaf and the leaf surface to which it was applied (abaxial or adaxial). Once the zinc had moved through the leaf surface it appeared to bind strongly, with limited further redistribution. Regardless of this, in these underlying tissues zinc moved into the lower-order veins, with concentrations 2- to 10-fold higher than in the adjacent tissues. However, even once in higher-order veins, the movement of zinc was still comparatively limited, with concentrations decreasing to levels similar to the background within 1–10 mm.Conclusions The results advance our understanding of the factors that influence the efficacy of foliar zinc fertilizers and demonstrate the merits of an innovative methodology for studying foliar zinc translocation mechanisms. [ABSTRACT FROM PUBLISHER]

The speciation and spatial distribution of selenium (Se) in hydrated plant tissues is not well understood. Using synchrotron-based x-ray absorption spectroscopy and x-ray fluorescence microscopy (two-dimensional scanning [and associated mathematical model] and computed tomography), the speciation and distribution of toxic Se were examined within hydrated roots of cowpea (Vigna unguiculata) exposed to either 20 µM selenite or selenate. Based upon bulk solution concentrations, selenate was 9-fold more toxic to the roots than selenite, most likely due to increased accumulation of organoselenium (e.g. selenomethionine) in selenate-treated roots. Specifically, uptake of selenate (probably by sulfate transporters) occurred at a much higher rate than for selenite (apparently by both passive diffusion and phosphate transporters), with bulk root tissue Se concentrations approximately 18-fold higher in the selenate treatment. Although the proportion of Se converted to organic forms was higher for selenite (100%) than for selenate (26%), the absolute concentration of organoselenium was actually approximately 5-fold higher for selenate-treated roots. In addition, the longitudinal and radial distribution of Se in roots differed markedly: the highest tissue concentrations were in the endodermis and cortex approximately 4 mm or more behind the apex when exposed to selenate but in the meristem (approximately 1 mm from the apex) when exposed to selenite. The examination of the distribution and speciation of Se in hydrated roots provides valuable data in understanding Se uptake, transport, and toxicity. [ABSTRACT FROM AUTHOR]

Arsenic (As) is considered to be the environmental contaminant of greatest concern due to its potential accumulation in the food chain and in humans. Using novel synchrotron-based x-ray fluorescence techniques (including sequential computed tomography), short-term solution culture studies were used to examine the spatial distribution of As in hydrated and fresh roots of cowpea (Vigna unguiculata 'Red Caloona') seedlings exposed to 4 or 20 µm arsenate [As(V)] or 4 or 20 µm arsenite. For plants exposed to As(V), the highest concentrations were observed internally at the root apex (meristem), with As also accumulating in the root border cells and at the endodermis. When exposed to arsenite, the endodermis was again a site of accumulation, although no As was observed in border cells. For As(V), subsequent transfer of seedlings to an As-free solution resulted in a decrease in tissue As concentrations, but growth did not improve. These data suggest that, under our experimental conditions, the accumulation of As causes permanent damage to the meristem. In addition, we suggest that root border cells possibly contribute to the plant's ability to tolerate excess As(V) by accumulating high levels of As and Limiting its movement into the root. [ABSTRACT FROM AUTHOR]

The focus of the present study was to investigate the mechanisms for the alleviation of Cu toxicity in plants by coexistent cations (e.g. Al3+, Mn2+, Ca2+, Mg2+, H+, Na+, and K+) and the development of an electrostatic model to predict 50% effect activities (EA50s) accurately. The alleviation of Cu2+ toxicity was evaluated in several plants in terms of (i) the electrical potential at the outer surface of the plasma membrane (PM) (Ψ0°) and (ii) competition between cations for sites at the PM involved in the uptake or toxicity of Cu2+, the latter of which is invoked by the Biotic Ligand Model (BLM) as the sole explanation for the alleviation of toxicity. The addition of coexistent cations into the bulk-phase medium reduces the negativity of Ψ0° and hence decreases the activity of Cu2+ at the PM surface. Our analyses suggest that the alleviation of toxicity results primarily from electrostatic effects (i.e. changes in both the Cu2+ activity at the PM surface and the electrical driving force across the PM), and that BLM-type competitive effects may be of lesser importance in plants. Although this does not exclude the possibility of competition, the data highlight the importance of electrostatic effects. An electrostatic model was developed to predict Cu2+ toxicity thresholds (EA50s), and the quality of its predictive capacity suggests its potential utility in risk assessment of copper in natural waters and soils. [ABSTRACT FROM PUBLISHER]

Manganese (Mn) is an essential micronutrient for plant growth but is often toxic in acid or waterlogged soils. Using cowpea (Vigna unguiculata L. Walp.) grown with 0.05–1500 μM Mn in solution, two short-term (48 h) solution culture experiments examined if the effects of cations (Ca, Mg, Na, Al, or H) on Mn nutrition are related to the root cells’ plasma membrane (PM) surface potential, ψ00 . When grown in solutions containing levels of Mn that were toxic, both relative root elongation rate (RRER) and root tissue Mn concentration were more closely related to the activity of Mn2+ at the outer surface of the PM, {Mn2+}00 (R2=0.812 and 0.871) than to its activity in the bulk solution, {Mn2+}b (R2=0.673 and 0.769). This was also evident at lower levels of Mn (0.05–10 μM) relevant to studies investigating Mn as an essential micronutrient (R2=0.791 versus 0.590). In addition, changes in the electrical driving force for ion transport across the PM influenced both RRER and the Mn concentration in roots. The {Mn2+}b causing a 50% reduction in root growth was found to be c. 500 to >1000 μM (depending upon solution composition), whilst the corresponding value was 3300 μM when related to {Mn2+}00 . Although specific effects such as competition are not precluded, the data emphasize the importance of non-specific electrostatic effects in the Mn nutrition of cowpea seedlings over a 1×105-fold range of Mn concentration in solution. [ABSTRACT FROM AUTHOR]

The phytotoxicity of trace metals is of global concern due to contamination of the landscape by human activities. Using synchrotron-based x-ray fluorescence microscopy and x-ray absorption spectroscopy, the distribution and speciation of copper (Cu), nickel (Ni), and zinc (Zn) was examined in situ using hydrated roots of cowpea (Vigna unguiculata) exposed to 1.5 µM Cu, 5 µM Ni, or 40 µM Zn for 1 to 24 h. After 24 h of exposure, most Cu was bound to polygalacturonic acid of the rhizodermis and outer cortex, suggesting that binding of Cu to walls of cells in the rhizodermis possibly contributes to the toxic effects of Cu. When exposed to Zn, cortical concentrations remained comparatively low with much of the Zn accumulating in the meristematic region and moving into the stele; approximately 60% to 85% of the total Zn stored as Zn phytate within 3 h of exposure. While Ni concentrations were high in both the cortex and meristem, concentrations in the stele were comparatively low. To our knowledge, this is the first report of the in situ distribution and speciation of Cu, Ni, and Zn in hydrated (and fresh) plant tissues, providing valuable information on the potential mechanisms by which they are toxic. [ABSTRACT FROM AUTHOR]

Electrical properties of plasma membranes (PMs), partially controlled by the ionic composition of the exposure medium, play significant roles in the distribution of ions at the exterior surface of PMs and in the transport of ions across PMs. The effects of coexisting cations (commonly Al3+, Ca2+, Mg2+, H+, Na+) on the uptake and toxicity of these and other ions (such as Cu2+, Zn2+, Ni2+, Cd2+, H2AsO4-) to plants were studied in terms of the electrical properties of PMs. Increased concentrations of cations or decreased pH in rooting media, whether in solution culture or in soils, reduced the negativity of the electrical potential at the PM exterior surface (ψ0°). This reduction decreased the activities of metal cations at the PM surface and increased the activities of anions such as H2AsO4-. Furthermore, the reduced ψ0° negativity increased the surface-to-surface transmembrane potential difference, thus increasing the electrical driving force for cation uptake and decreasing the driving force for anion uptake across PMs. Analysis of measured uptake and toxicity of ions using electrostatic models provides evidence that uptake and toxicity are functions of the dual effects of ψ0° (i.e. altered PM surface ion activity and surface-to-surface transmembrane potential difference gradient). This study provides novel insights into the mechanisms of plant-ion interactions and extends current theory to evaluate ion bioavailability and toxicity, indicating its potential utility in risk assessment of metal(loid)s in natural waters and soils. [ABSTRACT FROM AUTHOR]

Solution culture has been used extensively to determine the phytotoxic effects of trace metals. A review of the literature from 1975 to 2009 was carried out to evaluate the effects of As(V), Cd(II), Co(II), Cu(II), Hg(II), Mn(II), Ni(II), Pb(II), and Zn(II) on plants grown in solution. A total of 119 studies was selected using criteria that allowed a valid comparison of the results; reported toxic concentrations varied by five orders of magnitude. Across a range of plant species and experimental conditions, the phytotoxicity of the trace metals followed the trend (from most to least toxic): Pb≈Hg >Cu >Cd≈As >Co≈Ni≈Zn >Mn, with median toxic concentrations of (μM): 0.30 Pb, 0.47 Hg, 2.0 Cu, 5.0 Cd, 9.0 As, 17 Co, 19 Ni, 25 Zn, and 46 Mn. For phytotoxicity studies in solution culture, we suggest (i) plants should be grown in a dilute solution which mimics the soil solution, or that, at a minimum, contains Ca and B, (ii) solution pH should be monitored and reported (as should the concentrations of the trace metal of interest), (iii) assessment should be made of the influence of pH on solution composition and ion speciation, and (iv) both the period of exposure to the trace metal and the plant variable measured should be appropriate. Observing these criteria will potentially lead to reliable data on the relationship between growth depression and the concentration of the toxic metal in solution. [ABSTRACT FROM PUBLISHER]