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Phylogenetic relationships within cation transporter families of Arabidopsis.

Uptake and translocation of cationic nutrients play essential roles in physiological processes including plant growth, nutrition, signal transduction, and development. Approximately 5% of the Arabidopsis genome appears to encode membrane transport proteins. These proteins are classified in 46 unique families containing approximately 880 members. In addition, several hundred putative transporters have not yet been assigned to families. In this paper, we have analyzed the phylogenetic relationships of over 150 cation transport proteins. This analysis has focused on cation transporter gene families for which initial characterizations have been achieved for individual members, including potassium transporters and channels, sodium transporters, calcium antiporters, cyclic nucleotide-gated channels, cation diffusion facilitator proteins, natural resistance-associated macrophage proteins (NRAMP), and Zn-regulated transporter Fe-regulated transporter-like proteins. Phylogenetic trees of each family define the evolutionary relationships of the members to each other. These families contain numerous members, indicating diverse functions in vivo. Closely related isoforms and separate subfamilies exist within many of these gene families, indicating possible redundancies and specialized functions. To facilitate their further study, the PlantsT database (http://plantst.sdsc.edu) has been created that includes alignments of the analyzed cation transporters and their chromosomal locations.

Antiporters↗

Rapidly Induced Wound Ethylene from Excised Segments of Etiolated Pisum sativum L., cv. Alaska: I. Characterization of the Response.

A rapidly induced, transitory increase in the rate of ethylene synthesis occurred in wounded tissue excised from actively growing regions of etiolated barley, cucumber, maize, oat, pea, tomato, and wheat seedlings. Cutting intact stems or excising 9-mm segments of tissue from near the apex of 7-day-old etiolated Pisum sativum L., cv. Alaska seedlings induced a remarkably consistent pattern of ethylene production. At 25 C, wound-induced ethylene production by segments excised 9 mm below the apical hook increased linearly after a lag of 26 minutes from 2.7 nanoliters per g per hour to the first maxium of 11.3 nanoliters per g per hour at 56 minutes. The rate of production then decreased to a minimum at 90 minutes, increased to a lower second maximum at 131 minutes, and subsequently declined over a period of about 100 minutes to about 4 nanoliters per g per hour. Removal of endogenous ethylene, before the wound response commenced, had no effect on the kinetics of ethylene production. Tissue containing large amounts of dissolved ethylene released it as an exponential decay with no lag period. Rapidly induced wound ethylene is synthesized by the tissue and is not merely the result of facilitated diffusion of ethylene already present in the tissue through the newly exposed cut surfaces. Previously wounded apical sections did not exhibit a second response when rewounded. No significant correlation was found between wound-induced ethylene synthesis and either CO(2) or ethane production.

Journal Article↗

Essential Sulfhydryl Group in the Transport-catalyzing Protein of the Hexose-Proton Cotransport System of Chlorella.

The polyene antibiotic nystatin transforms the sugar-proton contransport system of Chlorella to a mere facilitated diffusion system. This experimental condition was used to test the sugar-translocating unit of the active uptake system for possible essential sulfhydryl groups. It could be shown that the catalyzed translocation of sugar is sensitive to the sulfhydryl-reactive compound N-ethylmaleimide. Sugar flow by passive leak as induced by the detergent Triton X-100 is not affected by sulfhydryl reagents. These results show that the sugar-translocating carrier protein possesses a sulfhydryl group, which is essential for its function.

Journal Article↗

Sulfhydryl Group Involvement in Plasmalemma Transport of HCO(3) and OH in Chara corallina.

The effect of the sulfhydryl reagents (-SH) p-chloromercuribenzene-sulfonic acid (PCMBS), N-ethylmaleimide (NEM), and inorganic mercury on H(14)CO(3) (-) assimilation in Chara corallina is reported. Commercial grade PCMBS caused severe inhibition of H(14)CO(3) (-) assimilation. Results obtained using purified PCMBS (stock solution passed through a chelating resin) indicated that inhibition observed using unpurified PCMBS was due predominantly to the presence of inorganic mercury (as a contaminant). The inhibitory role of inorganic mercury was verified using HgCl(2). This chemical caused a dramatic inhibition of H(14)CO(3) (-) assimilation, while it had little effect on cellular (14)CO(2) fixation. Reversal of the Hg(2+) inhibition of H(14)CO(3) (-) assimilation (in presence of 1.0 millimolar dithioerythritol) was extremely slow, requiring 2 to 3 hours for the reestablishment of control rates. This slow recovery may reflect de novo synthesis of transport proteins.Almost complete (irreversible) inhibition of H(14)CO(3) (-) assimilation was observed after cells were briefly (2 min) exposed to 0.1 to 0.3 millimolar NEM. At critical concentrations, both inorganic mercury and NEM perturbed the HCO(3) (-) transport system to such an extent that the H(14)CO(3) (-) assimilation versus HCO(3) concentration profiles changed shape. This could reflect a conversion of the transport system to a facilitated diffusion mode. Alternatively, the efficiency of the transport system may have been affected such that HCO(3) (-) efflux increased significantly.The influence of these -SH reagents on the OH(-) efflux system was also investigated. The H(14)CO(3) (-) assimilation and OH(-) efflux results demonstrate the involvement of -SH groups in the plasmalemma transport of both HCO(3) (-) and OH(-) in C. corallina. Greater transport susceptibility to inorganic mercury and NEM compared to purified PCMBS suggested that the sulfhydryl groups were not located on the peripheral surface of the plasmalemma.

Journal Article↗

Potassium transport in corn roots : I. Resolution of kinetics into a saturable and linear component.

Influx isotherms were obtained for (86)Rb(+) uptake into 2-cm corn (Zea mays [A632 x (C3640 x Oh43)] root segments for both low- (0.2 millimolar CaSO(4)) and high-salt (0.2 millimolar CaSO(4) + 5 millimolar KCl) grown roots. Unlike the discontinuous curves usually presented for K(+) influx, our isotherms were smooth, nonsaturating curves that approached linearity at K(+) (Rb(+)) concentrations above 1 millimolar. The kinetics for K(+) transport could be resolved into saturable and linear components. The saturable components yielded K(m) values of 16 and 86 micromolar for low- and high-salt roots, respectively, while V(max) values were 5.62 and 1.85 moles per gram fresh weight per hour. Results of experiments with the penetrating sulfhydryl reagent, N-ethyl maleimide (NEM), and the impermeant reagent, p-chloromercuribenzene sulfonic acid (PCMBS) indicated that the saturable and linear components were independent mechanisms of K(+) transport.Short-term NEM exposures (30 seconds to 5 minutes) selectively inhibited the saturable system, but had little effect on the linear component. Increasing NEM exposures resulted in further inhibition and subsequent abolition of the saturable component; the linear component exhibited limited NEM sensitivity. PCMBS elicited the same general inhibitory trends, although it was less effective as a saturable component inhibitor.The effects of NEM and PCMBS on K(+) efflux were also studied. Short NEM exposures had no effect on cytoplasmic efflux, while inhibiting vacuolar efflux significantly. From these data, it is unclear at which site(s) NEM is acting. A more complex response was obtained with PCMBS, where a monophasic efflux curve was observed. Analysis indicated that the vacuolar efflux was stimulated, while the cytoplasmic component was abolished.The nature of the linear component is discussed, and it is proposed that the mechanism may be more complex than simple facilitated diffusion.

Journal Article↗

Uptake of l-Ascorbate by Intact Spinach Chloroplasts.

Uptake of l-[1-(14)C]ascorbate by intact ascorbate-free spinach (Spinacia oleracea L. cv Vital(r)) chloroplasts has been investigated using the technique of silicone oil filtering. Rates greater than 100 micromoles per milligram chlorophyll per hour (external concentration, 10 millimolar) of ascorbate transport were observed. Ascorbate uptake into the sorbitol-impermeable space (stroma) followed the Michaelis-Menten-type characteristic for substrate saturation. A K(m) of 18 to 40 millimolar was determined. Transport of ascorbate across the chloroplast envelope resulted in an equilibrium of the ascorbate concentrations between stroma and medium. A pH optimum of 7.0 to 7.5 and the lack of alkalization of the medium upon ascorbate uptake suggest that only the monovalent ascorbate anion is able to cross the chloroplast envelope. The activation energy of ascorbate uptake was determined to be 65.8 kilojoules (16 kilocalories) per mole (8 to 20 degrees C). Interference of ascorbate transport with substrates of the phosphate or dicarboxylate translocator could not be detected, but didehydroascorbate was a competitive inhibitor. Preloading of chloroplasts with didehydroascorbate resulted in an increase of V(max) but did not change the K(m) for ascorbate. Millimolar concentrations of the sulfhydryl reagent p-chloromercuriphenyl sulfonate inhibited ascorbate uptake. The data are interpreted in terms of ascorbate uptake into chloroplasts by the mechanism of facilitated diffusion mediated by a specific translocator.

Journal Article↗

Characterization of amino Acid efflux from isolated soybean cells.

Cells from reproductive soybean (Glycine max [L.] Merr.) plants were isolated using a mechanical-enzymic technique that produced a high yield of uniform, physiologically active cells. Cells were incubated in a pH 6.0 buffered solution and subjected to various treatments in order to determine the nature of net amino acid efflux. Total net amino acid (ninhydrinreactive substances) efflux was not affected by the following conditions: (a) darkness, (b) aeration, (c) K(+) concentrations of 0.1, 1.0, 10, or 100 millimolar and (d) pH 4, 5, 6, 7, or 8. The Q(10) for net amino acid efflux between 10 degrees C and 30 degrees C was 1.6. Thus, it seems that net amino acid efflux requires neither current photosynthetic energy nor a pH/ion concentration gradient. Amino acid analyses of the intra-and extracellular fractions over time showed that each amino acid was exported linearly for at least 210 minutes, but that export rate was not necessarily related to internal amino acid pools. Amino acids that were exported fastest were alanine, lysine, leucine, and glycine. Addition of the inhibitor p-chloromercuriphenyl sulfonic acid, 3(3,4-dichlorophenyl)-1,1-dimethylurea, or carbonylcyanide p-trifluoromethoxyphenylhydrazone increased the rate of total amino acid efflux but had specific effects on the efflux of certain amino acids. For example, p-chloromercuriphenyl sulfonic acid greatly enhanced efflux of gamma-aminobutyric acid, which is not normally exported rapidly even though a high concentration normally exists within cells. The data suggest that net amino acid efflux is a selective diffusional process. Because net efflux is the result of simultaneous efflux and influx, we propose that efflux is a facilitated diffusion process whereas influx involves energy-dependent carrier proteins.

Journal Article↗

Transport Properties of the Tomato Fruit Tonoplast : II. Citrate Transport.

Citrate transport across the membrane of tomato fruit tonoplast vesicles was investigated. In the tonoplast vesicles, [(14)C]methylamine uptake was stimulated 10-fold by MgATP and strongly inhibited by NO(3) (-). Under identical experimental conditions, [(14)C]citrate uptake was inhibited by 5 millimolar free Mg(2+), and this inhibition was reversed in the presence of ATP, presumably by ATP chelation of free Mg(2+). No evidence was obtained in support of energy-linked ATP stimulation of citrate uptake. Citrate uptake showed saturation kinetics, and was inhibited by 4,4'-diisothiocyano-2,2'-stilbenedisulfonic acid and by other organic acids. The pH-dependence of uptake suggested that citrate(3-) was the transported species. Our results indicate that citrate transport across the tomato fruit tonoplast occurs by facilitated diffusion of citrate(3-). The carrier shares some features in common with anion channels in that it is relatively nonspecific for organic acids and is inhibitable by 4,4'-diisothyocyano-2,2'-stilbenedisulfonic acid.

Journal Article↗

Facilitated transport of glucose in isolated Phloem segments of celery.

In isolated phloem segments of celery (Apium graveolens L.), a tissue highly specific for sucrose and mannitol uptake, glucose uptake occurs at very low rates and exhibits biphasic kinetics. Nonpenetrating inhibitors such as parachloromercuribenzene sulfonic acid did not inhibit glucose uptake. However, uptake was greatly inhibited by penetrating inhibitors such as N-ethylmaleimide and carbonylcyanide-m-chlorophenyl hydrazone. Carbonylcyanide-m-chlorophenyl hydrazone inhibition of uptake was reversed by washing and addition of thiol reagents to uptake solutions. Phlorizin, a competitive inhibitor of glucose caused moderate inhibition of uptake only after 3 hours of tissue exposure. Low pH, fusicoccin, and low turgor which enhance H(+)-sugar cotransport did not alter uptake rates. Furthermore, glucose did not induce alkalinization of the uptake media. Efflux analysis indicated that the presence of 50 millimolar unlabeled glucose in the wash media enhanced exchange of the labeled glucose across the tonoplast. Results indicate that the glucose carrier is not located at the plasmalemma but appears to be present at the membrane of an intracellular compartment, most likely the tonoplast. Carrier-mediated glucose transport in this tissue is proposed to be a facilitated diffusion.

Journal Article↗

Exopolysaccharides Produced by Phytopathogenic Pseudomonas syringae Pathovars in Infected Leaves of Susceptible Hosts.

Bacterial exopolysaccharide (EPS) was extracted from infected leaves of several host plants inoculated with phytopathogenic strains of Pseudomonas syringae pathovars. Extraction was by a facilitated diffusion procedure or by collection of intercellular fluid using a centrifugation method. The extracted EPS was purified and characterized. All bacterial pathogens which induced watersoaked lesions on their host leaves, a characteristic of most members of this bacterial group, were found to produce alginic acid (a polymer consisting of varying ratios of mannuronic and guluronic acids). Only trace amounts of bacterial EPS could be isolated from leaves inoculated with a pathovar (pv. syringae) which does not induce the formation of lesions with a watersoaked appearance. Guluronic acid was either present in very low amounts or absent in the alginic acid preparations. All bacterial alginates were acetylated (7-11%). Levan (a fructan) was apparently not produced as an EPS in vivo by any of the pathogens tested.

Journal Article↗

Competitive Al Inhibition of Net Mg Uptake by Intact Lolium multiflorum Roots : I. Kinetics.

Aluminum impairs uptake of Mg(2+), but the mechanisms of this inhibition are not understood. The depletion technique was used to monitor net Mg(2+) uptake from nutrient solution by intact, 23-day-old plants of ryegrass (Lolium multiflorum Lam., cv Gulf and Wilo). Activities of Mg(2+) and monomeric Al species in nutrient solution were calculated and used as the basis for expressing the results. The kinetics of net Mg(2+) absorption was resolved into (a) a transpiration-dependent uptake component, (b) a metabolically mediated, discontinuous saturable component that is Al(3+) sensitive and p-chloromercuribenzene sulfonic acid (PCMBS) resistant, and (c) a linear, carbonyl cyanide m-chlorophenylhydrazone resistant, Al(3+) sensitive component that might be a type of facilitated diffusion. Lowering the pH from 6.0 to 4.2 exerted a noncompetitive inhibition of net Mg(2+) uptake, while aluminum at 6.6 micromolar Al(3+) activity exerted competitive inhibition of net Mg(2+) uptake at pH 4.2. The Al(3+)-induced effect was obvious after 30 minutes. Cultivar-specific ability to retain a higher affinity for Mg(2+) by postulated transport proteins in the presence of Al(3+) might be one of the mechanisms of differential Al tolerance among ryegrass cultivars.

Journal Article↗

Poplar metal tolerance protein 1 confers zinc tolerance and is an oligomeric vacuolar zinc transporter with an essential leucine zipper motif.

Cation diffusion facilitator (CDF) proteins are a recently discovered family of cation efflux transporters that might play an essential role in metal homeostasis and tolerance. Here, we describe the identification, characterization, and localization of PtdMTP1, a member of the CDF family from the hybrid poplar Populus trichocarpa x Populus deltoides. PtdMTP1 is expressed constitutively and ubiquitously, although at low levels. Heterologous expression in yeast showed that PtdMTP1 was able to complement the hypersensitivity of mutant strains to Zn but not to other metals, including Cd, Co, Mn, and Ni. PtdMTP1 fused to green fluorescent protein localized to the vacuolar membrane both in yeast and in plant cells, consistent with a function of PtdMTP1 in zinc sequestration. Overexpression of PtdMTP1 in Arabidopsis confers Zn tolerance. We show that PtdMTP1, when expressed in yeast and Arabidopsis, forms homooligomers, a novel feature of CDF members. Oligomer formation is disrupted by reducing agents, indicating possible disulfide bridge formation. PtdMTP1 also contains a conserved Leu zipper motif. Although not necessary for oligomer formation, Leu residues within this motif are required for PtdMTP1 functional activity.

Adaptation, Physiological↗

Structure of bovine carbonic anhydrase II at 1.95 A resolution.

Carbonic anhydrase (CA) is a zinc-containing enzyme that catalyzes the reversible hydration of CO2 to HCO3-. In eukaryotes, the enzyme plays a role in various physiological functions, including interconversion between CO2 and HCO3- in intermediary metabolism, facilitated diffusion of CO2, pH homeostasis and ion transport. The structure of bovine carbonic anhydrase II (BCA II) has been determined by molecular replacement and refined to 1.95 A resolution by simulated-annealing and individual B-factor refinement. The final R factor for the BCA II structure was 19.4%. BCA II has a C-terminal knot structure similar to that observed in human CA II. It contains one zinc ion in the active site coordinated to three histidines and one putative water molecule in a tetrahedral geometry. The structure of BCA II reveals a probable alternative proton-wire pathway that differs from that of HCA II.

Animals↗

Dipyridamole potentiates the growth-inhibitory action of methotrexate and 5-fluorouracil in human keratinocytes in vitro.

Human keratinocytes transport extracellular thymidine across the plasma membrane and incorporate it into DNA. Data presented here show that dipyridamole, a well-known inhibitor of facilitated diffusion of nucleosides, blocks the transport of thymidine into human keratinocytes in vitro. Dipyridamole (1.0 microM) inhibited the transport of 3H-thymidine (0.2 microM) into intracellular material by 75% and its subsequent salvage and incorporation into DNA by 48%. Dipyridamole (1 microM) did not affect the growth of keratinocytes in vitro but did potentiate the growth inhibition caused by methotrexate (MTX) or 5-fluorouracil (5-FU). The growth of keratinocytes exposed to 0.1 microM MTX for 8 d was inhibited by 32%. However, in combination with a noninhibitory concentration of dipyridamole (1 microM), this concentration of MTX (0.01 microM) inhibited the growth of keratinocytes by 93%. Thymidine in culture medium reversed the cytotoxicity of MTX. However, in the presence of dipyridamole, thymidine in the culture medium did not reverse the action of MTX. The synergistic interaction between MTX and dipyridamole was also observed with 5-FU and dipyridamole. 5-FU (0.5 microM) inhibited cell growth by 30% but in combination with dipyridamole (1 microM), inhibited cell growth by 86%. These data are consistent with the theory that inhibiting thymidine salvage by blocking transport of extracellular thymidine potentiates the growth inhibitory action of inhibitors of de novo pyrimidine biosynthesis in human keratinocytes. Combination chemotherapy, such as methotrexate plus dipyridamole, might be efficacious in the treatment of hyperproliferative diseases of the epidermis.

Cell Division↗

Nitrite transport into pig erythrocytes and its potential biological role.

AIM: To study nitrite transport and its oxygenation dependency in pig erythrocytes, as this is fundamental to the possible participation of nitrite in blood flow regulation via its reduction to nitric oxide by deoxygenated haemoglobin (Hb). METHODS: Pig red blood cells (RBCs) were tonometer-equilibrated to physiological pCO2 in oxygenated and deoxygenated states. Nitrite was added and the kinetics of NO2- influx and methaemoglobin (metHb) formation were assessed at variable temperature and haematocrit. RESULTS: Nitrite quickly permeated and equilibrated across the membrane, and then continued to enter RBCs as a consequence of its intracellular removal (via reactions with Hb to form nitrate and metHb in oxygenated cells, and NO and metHb in deoxygenated cells). The membrane permeation as such showed little oxygenation dependency, but as metHb formation was significantly higher in oxygenated than deoxygenated RBCs, nitrite transport tended to be largest into oxygenated RBCs. This contrasts with a preferential permeation of deoxygenated RBCs in some fish species. Nitrite transport showed low temperature sensitivity but was speeded up at low haematocrit via more rapid intracellular nitrite removal (metHb formation). Nitrite influx was not affected by inhibitors of facilitated diffusion (DIDS, phloretin and PCMB) and may occur via conductive transport. Extracellular pH was stable during nitrite transport. CONCLUSION: Nitrite extensively permeates both oxygenated and deoxygenated pig RBCs, which may enable a dual function of nitrite entry: viz. conversion to NO at low pO2 to promote blood flow and detoxification to non-toxic nitrate at inappropriate high nitrite levels.

Animals↗

Micro-organisms and dust exposure in an Italian grain mill.

AIMS: In order to assess possible occupational risk for workers in a grain mill, we evaluated aerial microbiological contamination in different areas of the mill and at different points of the production line. We also measured the concentration of aerodispersed dust particles. METHODS AND RESULTS: An assessment of microbiological contamination levels based on a Global Index of Microbial Contamination per cubic metre (GIMC per m3), an Index of Mesophilic Bacterial Contamination, and an Amplification Index is proposed. The indices were obtained from total and fungal counts. The cleaning sector is the most contaminated area of the mill: the mean GIMC per m3 was 17,213.6. In this area, the average microbial contamination was 11.41 times higher than that in the external environment. The highest concentrations of aerodispersed dust (inhalable 2.763 mg m(-3); respirable 1.400 mg m(-3)) were found in the cleaning area. CONCLUSIONS: The proposed microbiological indices and the concentrations of aerodispersed dust particles show that the most hazardous section of the mill is the cleaning area. The large variation in the data does not depend on seasonal factors, but rather on not easily identifiable conditions of the internal environment which facilitate diffusion and/or proliferation of the micro-organisms. SIGNIFICANCE AND IMPACT OF THE STUDY: The proposed microbiological contamination indices and the evaluation of the concentration of dust particles allow the identification of critical positions during the production cycle so that suitable measures to prevent the aerial contamination can be taken.

Air Pollutants, Occupational↗

Genetic and physiological responses of Bacillus subtilis to metal ion stress.

Metal ion homeostasis is regulated principally by metalloregulatory proteins that control metal ion uptake, storage and efflux genes. We have used transcriptional profiling to survey Bacillus subtilis for genes that are rapidly induced by exposure to high levels of metal ions including Ag(I), Cd(II), Cu(II), Ni(II) and Zn(II) and the metalloid As(V). Many of the genes affected by metal stress were controlled by known metalloregulatory proteins (Fur, MntR, PerR, ArsR and CueR). Additional metal-induced genes are regulated by two newly defined metal-sensing ArsR/SmtB family repressors: CzrA and AseR. CzrA represses the CadA efflux ATPase and the cation diffusion facilitator CzcD and this repression is alleviated by Zn(II), Cd(II), Co(II), Ni(II) and Cu. CadA is the major determinant for Cd(II) resistance, while CzcD protects the cell against elevated levels of Zn(II), Cu, Co(II) and Ni(II). AseR negatively regulates itself and AseA, an As(III) efflux pump which contributes to arsenite resistance in cells lacking a functional ars operon. Our results extend the range of identified effectors for the As(III)-sensor ArsR to include Cd(II) and Ag(I) and for the Cu-sensor CueR to include Ag(I) and, weakly, Cd(II) and Zn(II). In addition to systems dedicated to metal homeostasis, specific metal stresses also strongly induced pathways related to cysteine, histidine and arginine metabolism.

Adenosine Triphosphatases↗

High expression in leaves of the zinc hyperaccumulator Arabidopsis halleri of AhMHX, a homolog of an Arabidopsis thaliana vacuolar metal/proton exchanger.

Zn hyperaccumulator plants sequester Zn into their shoot vacuoles. To date, the only transporters implicated in Zn sequestration into the vacuoles of hyperaccumulator plants are cation diffusion facilitators (CDFs). We investigated the expression in Arabidopsis halleri of a homolog of AtMHX, an A. thaliana tonoplast transporter that exchanges protons with Mg, Zn and Fe ions. A. halleri has a single copy of a homologous gene, encoding a protein that shares 98% sequence identity with AtMHX. Western blot analysis with vacuolar-enriched membrane fractions suggests localization of AhMHX in the tonoplast. The levels of MHX proteins are much higher in leaves of A. halleri than in leaves of the non-accumulator plant A. thaliana. At the same time, the levels of MHX transcripts are similar in leaves of the two species. This suggests that the difference in MHX levels is regulated at the post-transcriptional level. In vitro translation studies indicated that the difference between AhMHX and AtMHX expression is not likely to result from the variations in the sequence of their 5' untranslated regions (5'UTRs). The high expression of AhMHX in A. halleri leaves is constitutive and not significantly affected by the metal status of the plants. In both species, MHX transcript levels are higher in leaves than in roots, but the difference is higher in A. halleri. Metal sequestration into root vacuoles was suggested to inhibit hyperaccumulation in the shoot. Our data implicate AhMHX as a candidate gene in metal accumulation or tolerance in A. halleri.

Amino Acid Sequence↗