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comparative trace element nutrition 1 trace element uptake and distribution in plants robin d graham2 and james c r stangoulis department of plant science university of adelaide waite campus south ...

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                                           Comparative Trace Element Nutrition
                                                                                                                     1
         Trace Element Uptake and Distribution in Plants
                    Robin D. Graham2 and James C. R. Stangoulis
                    Department of Plant Science, University of Adelaide, Waite Campus, South Australia 5064
                    ABSTRACT Therearesimilarities between mammals and plants in the absorption and transport of trace elements.
                    The chemistry of trace element uptake from food sources in both cases is based on the thermodynamics of
                    adsorption on charged solid surfaces embedded in a solution phase of charged ions and metal-binding ligands                                                       Downloaded from https://academic.oup.com/jn/article/133/5/1502S/4558537 by guest on 05 January 2023
                    together with redox systems in the case of iron and some other elements. Constitutive absorption systems function in
                    nutrient uptake during normal conditions, and inducible turbo systems increase the supply of a particular nutrient
                    during deficiency. Iron uptake is the most studied of the micronutrients, and divides the plant kingdom into two
                    groups: dicotyledonous plants have a turbo system that is an upregulated version of the constitutive system, which
                    consists of a membrane-bound reductase and an ATP-driven hydrogen ion extrusion pump; and monocotyledonous
                    plants have a constitutive system similar to that of the dicots, but with an inducible system remarkably different that
                    usesthemugeneicacidclassofphytosiderophores(PS).ThePSsystemmayinfactbeanimportantportofentryfor
                    iron from an iron-rich but exceedingly iron-insoluble lithosphere into the iron-starved biosphere. Absorption of trace
                    metals in these graminaceous plants is normally via divalent ion channels after reduction in the plasma membrane.
                    Once absorbed, iron can be stored in plants as phytoferritin or transported to active sites by transport-specific
                    ligands. The transport of iron and zinc into seeds is dominated by the phloem sap system, which has a high pH that
                    requires chelation of heavy metals. Loading into grains involves three or four genes each that control chelation,
                    membrane transport and deposition as phytate.                   J. Nutr. 133: 1502S–1505S, 2003.
                    KEYWORDS:  micronutrients  iron  zinc  absorption  transport  plants  animals  genetics
             Themicronutrients that are known to be required by plants                            transport is facilitated when external concentrations are low (1)
         are iron, zinc, copper, manganese, cobalt, nickel, boron,                                as they commonly are in acid soils everywhere.
         molybdenum and chlorine. The last two are present in soils                                   The remaining six micronutrients for higher plants, the
         as anions and undoubtedly require active transport across the                            transition metals, are generally absorbed as divalent ions via
         plasmalemma of plant root cells for uptake. Boron is either an                           divalent ion channels. These channels either have consider-
         anion or neutral molecule in most soils, and the neutral                                 able specificity for each element, or homeostasis is achieved
         molecule is fairly permeable across biological membranes (1).                            by specific active-excretion mechanisms that are controlled
         Whetherboronisactivelytransportedinto plants is a subject of                             by cytoplasmic concentrations (2). Because iron and zinc
         considerable interest in current literature, but new evidence                            deficiencies are extremely widespread in humans and are also
         suggests that although it may enter as a neutral molecule, boron                         common in some farm animals, this article concentrates on
                                                                                                  their uptake, transport and loading into grains that constitute
                                                                                                  the staple foods of most of the human race. The genetics ex-
                                                                                                  hibited by these processes are also addressed because of the
             1 Published in a supplement to The Journal of Nutrition. Presented as part of        interest in breeding new varieties of staple food crops with
         the 11th meeting of the international organization, Trace Elements in Man and          greater micronutrient density. What is known about the
         Animals (TEMA), in Berkeley, California, June 2–6, 2002. This meeting was              uptake, transport and loading of the other transition elements
         supported by grants from the National Institutes of Health and the U.S. Department       is generally analogous to iron and zinc. However, in the case of
         of Agriculture and by donations from Akzo Nobel Chemicals, Singapore; California         manganese, the redox systems that are important are in the soil
         Dried Plum Board, California; Cattlemens Beef Board and National Cattlemens
         Beef Association, Colorado; GlaxoSmithKline, New Jersey; International Atomic            itself and are controlled by the balance of manganese-oxidizing
         Energy Agency, Austria; International Copper Association, New York; International        and -reducing soil microorganisms, which in turn is controlled
         Life Sciences Institute Research Foundation, Washington, D.C.; International Zinc        by soil and environmental conditions as well as by plant root
         Association, Belgium; Mead Johnson Nutritionals, Indiana; Minute Maid Company,
         Texas; Perrier Vittel Water Institute, France; U.S. Borax, Inc., California; USDA/       activities.
         ARS Western Human Nutrition Research Center, California and Wyeth-Ayerst
         Global Pharmaceuticals, Pennsylvania. Guest editors for the supplement
         publication were Janet C. King, USDA/ARS WHNRC and the University of
         California at Davis; Lindsay H. Allen, University of California at Davis; James R.       Iron uptake by plant roots
         Coughlin, Coughlin & Associates, Newport Coast, California; K. Michael
         Hambidge, University of Colorado, Denver; Carl L. Keen, University of California             Planet Earth is replete in iron that constitutes much of its
                          ¨
         at Davis; Bo L. Lonnerdal, University of California at Davis and Robert B. Rucker,       molten core, and iron is also the fourth most abundant element
         University of California at Davis.
             2 To whom correspondence should be addressed. E-mail: r.graham@cgiar.                in the earth’s crust. The amount of iron in the soil may be
         org.                                                                                     10,000timesgreaterthaninthevegetationgrowninit,yetiron
         0022-3166/03 $3.00  2003 American Society for Nutritional Sciences.
                                                                                          1502S
                                                              TRACEELEMENTUPTAKEANDDISTRIBUTIONINPLANTS                                                                                1503S
            deficiency is common in crop plants. This anomaly is due to the
            low availability of iron in the presence of oxygen especially at
            moderate and high soil pH values. The solubility product of
            some compounds formed in soil that precipitate iron is on
            the order of 10235. These forms of iron in the soil are only
            solubilized by lowering of the pH value, by complexation of
            ferric iron [Fe(III)] and/or by reduction of Fe(III) to ferrous
            iron [Fe(II)].3 The strategies used by plant roots to access iron
            exploit each of these chemical options, but the mechanisms
            vary between species in such a way as to divide the plant
            kingdom into two groups known as Strategy I and Strategy II
            plants (3). The latter group is the Gramineae, and the former
            includes all dicotyledonous plants together with the non-
            graminaceous monocotyledonous plants.                                                                                                                                               Downloaded from https://academic.oup.com/jn/article/133/5/1502S/4558537 by guest on 05 January 2023
                Both groups have a constitutive system that is adequate to
            supply plants that are grown in fertile soils having plenty of                                   FIGURE 1 Strategy I: upregulation of the constitutive system for
            available forms of iron. The constitutive system consists of                                 iron uptake, which is characteristic of dicotyledonous plants. R, inducible
            a membrane-bound ferric reductase that is linked to a divalent                               reductase; PM, plasma membrane. [Adapted from Romheld (18).]
            ion transporter or channel and an ATP-driven proton-
            extrusion pump. Recently, Rogers et al. (4) showed that
            single–amino acid substitutions in the sequence of this channel                              ligand is separated from the metal by reduction of the latter,
            protein create specificity for the various divalent cations. These                            which is then stored in phytoferritin or transported in the
            two membrane functions are able to supply adequate iron to                                   plant with ferrous-specific ligands such as nicotianamine.
            most plants in a healthy soil. However, in iron-deficient soil,                               Graminaceous species contain the various members of the PS
            iron chlorosis (yellowing) in leaf tissues occurs, and additional                            family (Fig. 3) in unique ratios: generally, the small-grain
            mechanisms of iron acquisition are induced to restore plants’                                cereals such as barley, wheat, oat and rye have the greatest
            iron status. In both strategies, these induced responses are                                 expression, which explains their remarkable adaptation to the
            restricted to the apical zones of the roots and are fully shut                               high-pH soils that are usually found in the semi-arid winter-
            down again within 1 d of restoration of normal iron status.                                  cereal–cropping belts of the world. The PS pathway appears to
                Strategy I plants respond to signals of low iron status by                               be a major vehicle for the entry of iron into the biosphere from
            upregulating the ferric reductase (by deploying a new 70-kDa                                 the lithosphere. Curiously, the release of PS from the roots is
            protein in the membrane) and the proton-extrusion pump.                                      diurnal and peaks a few hours after sunrise. As in Strategy I
            In addition, many Strategy I plants have a mechanism for                                     plants, the synthesis of PS is quickly suppressed when the plants
            excreting iron-binding ligands and soluble reductants, which                                 are restored to adequate iron status, which suggests that these
            are commonly phenols (Fig. 1). All of these changes are de-                                  inducible systems are energetically demanding.
            signed to solubilize iron by each of the processes mentioned,                                    PS also bind zinc, copper and manganese and can
            but the processes are only expressed in the apical zones of the                              enhance their absorption along with that of iron. However,
            roots where the adaptations are associated with changes in root                              with the possible exception of zinc, the mechanism is not
            morphology and the appearance of transfer cells with in-                                     induced by deficiency of these other transition metals in
            vaginated membranes. The reductase is stimulated by low pH                                   the plant. The constitutively expressed extrusion of protons,
            level and thereby by the proton-extrusion pump such that its                                 reductants and metal-binding ligands will enhance the absorp-
            function is effectively inhibited by bicarbonate in high-pH soils.                           tion of all the divalent cations. Inducible systems for upregu-
            This is the basis for the severe iron chlorosis that is seen in                              lated absorption of micronutrients are best understood for iron,
            dicotyledonous plants from high-pH soils.                                                    and indeed, although the existence of an inducible system in
                Insensitivity to bicarbonate is a feature of Strategy II plants,                         the gut of humans is generally accepted, its nature is not as
            which induce an entirely new mechanism of mobilizing iron                                    clearly understood as that in plants and bacteria. The latter
            under iron stress. Rather than upregulate the constitutive                                   haveaninduciblesystemthatinvolvesthesynthesisofmembers
            system, Strategy II plants synthesize and release to the soil                                of the hydroxamate group of ferric-binding ligands.
            nonprotein amino acids known as phytosiderophores (PS) or
            phytometallophores; the latter term recognizes that these
            amino acids are able to chelate most of the transition metals
            and not just iron (Fig. 2). These form strong soluble chelates
            with ferric ions in the soil, and because they are soluble and less
            positively charged, they are free to diffuse toward the root in
            soil-water films. Additionally, Strategy II plants have constitu-
            tively a highly specific transporter protein [the genes encoding
            for this transporter most likely belong to the natural resistance-
            associated macrophage protein (NRAMP) family (5,6) or the
            interferon-g–responsive transcript (IRT-1) family (7)]. This
            highly specific transporter protein, which is not present in
            Strategy I plants, recognizes and transports its specific ferric
            chelate across the membrane (Fig. 2). In the cytoplasm, the
                                                                                                             FIGURE 2 StrategyII:ahighlyefficientinducible-uptakesystemfor
                                                                                                         iron in graminaceous plants. X, enhanced release of phytosiderophores;
                3 Abbreviations used: Fe(II), ferrous iron; Fe(III), ferric iron; PS, phytosider-        P, specific uptake system for Fe(III) phytosiderophores. Both were
            ophore.                                                                                      induced under iron deficiency. [Adapted from Romheld (18).]
       1504S                                                      SUPPLEMENT
                                                                                                                                            Downloaded from https://academic.oup.com/jn/article/133/5/1502S/4558537 by guest on 05 January 2023
          FIGURE 3 Knownphytosiderophores in root exudates from graminaceous plants (19).
       Genetics                                                            Loading genes
          Thegeneticsofthemembrane-boundinduciblereductaseof                  The movement of iron from the vegetative plant into the
       dicotyledonous plants were first studied by Weiss (8) using one      grain is another major barrier. In rice, this barrier is extreme,
       of the iron-inefficient mutants that show up in soybean-             withconcentrationsinleavesasmuchas100timesgreaterthan
       breeding programs from time to time. In a series of elegant         in polished rice.
       studies, Weiss (8) cross-grafted scions and rootstocks of              Hitherto this discussion has concerned the absorption of
       efficient and inefficient soybean lines and showed that the           micronutrient cations from the soil into the root and or
       trait is expressed in the roots but the phenotype is expressed in   vegetative parts of the plant. Movement of micronutrients into
       the shoot. Later, this dominant major gene was shown to con-        grain (and from shoot to root or from leaf to leaf) involves the
       trol the ferric reductase activity of the membrane. However,        phloem, the secondary circulatory system of the plant, which
       in breeding programs, all useful breeding material is wild type     utilizes the movement of living-cell sap from cell to cell of the
       and iron efficient at this locus. Subsequent work identified          phloem sieve tubes. To be soluble and transportable in living-
       some 20 genes of minor effect that can enhance the iron ef-         cell sap at a pH of 7.5–8.5, the transition metal cations must be
       ficiency of soybean; this was significant in adapting this crop       strongly complexed. Many natural ligands in plants have been
       to the higher-pH soils of the midwestern U.S. In the same crop,     proposed for this role including di- and tricarboxylic acids,
       several genes were identified with zinc efficiency (9) and are        amino acids, amides and amines and especially nicotianamine,
       likely to be additive (10).                                         which is also an intermediate in PS synthesis. Steps in the
          Fromthebiosynthetic pathway, the genetics of PS synthesis        process include loading into the phloem, unloading, transport
       are potentially quite complex, but Mori and co-workers              across the mother plant/daughter plant barrier and deposition
       (11,12) have elucidated the biochemistry of this pathway,           in the aleurone layer as monoferric phytate. Lonergan (13)
       and the steps have been linked to particular chromosome             identified three loci associated with the loading of zinc into
       segments in barley. A locus on chromosome 4HS appears to be         barley grain: two from one parent of a doubled haploid
       particularly important. Lonergan (13) found that this locus         population and one from the other parent. Each locus ef-
       controls leaf zinc concentration in a doubled haploid pop-          fectively accounted for about one-third of the increase in
       ulation from the cross of Sahara and Clipper barleys. This locus    grain zinc content; together an increase of ;80% was observed
       controls the synthesis of mugineic acid from 29-deoxymu-            in those genotypes with favorable alleles at all three loci
       gineic acid (11). It is also closely linked to a gene of major ef-  compared to those with no favorable alleles. In a rice pop-
       fect that confers manganese efficiency in barley (14) as well as     ulation in which the parents differed in iron concentration by
       to a homeologous region of rye that confers not only part of the    a factor of two, four loci (quantitative trait loci) were in-
       zinc efficiency trait but also carries a major gene with a dom-      volved (16), and in beans, a similar number was reported by
       inant effect for copper efficiency (15). Homeologous genes           Beebe et al. (17). In both cases, there was a locus in common
       in durum wheat are also in this region. Manganese effici-            with those loci encoding the loading of zinc into grain, whereas
       ency in barley and durum wheat involves at least two loci           other loci were unrelated. It is of interest to know whether the
       between efficient and inefficient advanced breeding lines.            locus in common controls the concentration of nicotianamine
       Thus with the exception of a major gene in rye for copper           or some other ligand that is capable of stabilizing these metal
       efficiency, agronomic iron, zinc and manganese efficiency             ions at high pH values.
       in cereals (and in the few dicots studied) appears to be poly-         Comparisonsbetweenmammalianandplantsystemsintheir
       genic.                                                              uptake of trace elements are possible. Inducible high-affinity
                                                             TRACEELEMENTUPTAKEANDDISTRIBUTIONINPLANTS                                                                               1505S
            uptake in plants subjected to nutrient-deficient conditions is                                     7. Eide, D., Broderius, M., Fett, J. & Guerinot, M. L.  (1996)   Anovel iron-
            well documented, and in the case of iron, this process is well                              regulated metal transporter from plants identified by functional expression in yeast.
            understood. Even more sophisticated systems can be expected                                 Proc. Natl. Acad. Sci. U.S.A. 93: 5624–5628.
                                                                                                              8. Weiss, M. G.     (1943)   Inheritance and physiology of efficiency in iron
            in mammals, but these do not appear to be as well understood,                               utilization in soybeans. Genetics 28: 253–268.
            and the need for further research activity in this area is re-                                    9. Hartwig, E. E., Jones, W. F. & Kilen, T. C.      (1991)   Identification and
            quired. Sequence homology among micronutrient cation trans-                                 inheritance of inefficient zinc absorption in soybean. Crop Sci. 31: 61–63.
                                                                                                             10. Majumder, M. D., Rakshit, S. C. & Borthakur, D. N.         (1990)   Genetic
            porter proteins across the plant/animal divide must justify more                            effects on uptake of selected nutrients in some rice (Oryza sativa L.) varieties in
            studies of analogous systems in plants, animals and humans.                                 phosphorus deficient soil. Plant Soil 123: 117–120.
            With the application of molecular techniques, advances in our                                    11. Mori, S. & Nishizawa, N.     (1989)   Identification of barley chromosome
            understanding of trace element transport in animals should be                               4H, possible encoder of genes of mugineic acid synthesis from 29-deoxymugineic
                                                                                                        acid using wheat-barley addition lines. Plant Cell Physiol. 30: 1057–1061.
            rapid.                                                                                           12. Mori, S., Kishi-Nishizawa, N. & Fujigaki, J.   (1990)   Identification of rye
                                                                                                        chromosome 5R as the carrier of the gene for mugineic acid synthase and
                                                                                                        3-hydroxymugineicacidsynthaseusingwheat-ryeadditionlines.Jpn.J.Genet.65:
                                                                                                        343–352.
                                                                                                             13. Lonergan, P. F.    (2001)   Genetic characterization and QTL mapping of Downloaded from https://academic.oup.com/jn/article/133/5/1502S/4558537 by guest on 05 January 2023
                                                                                                        zinc nutrition in barley (Hordeum vulgare). Ph.D. thesis, University of Adelaide,
                                    LITERATURECITED                                                     Australia.
                                                                                                             14. Pallotta, M. A., Graham, R. D., Langridge, P., Sparrow, D. H. B. & Barker,
                 1. Stangoulis, J. C. R., Reid, R. J., Brown, P. H. & Graham, R. D.                     S. J.   (2000)   RFLP mapping of manganese efficiency in barley. Theor. Appl.
            (2001)   Kinetic analysis of boron transport in Chara. Planta 213: 142–146.                 Genet. 101: 1100–1108.
                 2. Welch, R. M.     (1995)   Micronutrient nutrition of plants. Crit. Rev. Plant            15. Graham, R. D. (1984) Breeding for nutritional characteristics in
            Sci. 14: 49–82.                                                                             cereals. Advances in Plant Nutrition, vol. 1 (Tinker, B. and Lauchli, A., eds.), pp.
                 3. Marschner, H.     (1995)   Mineral Nutrition of Higher Plants, 2nd ed.              57–102. Praeger Publishing, New York.
            Academic Press, London.                                                                          16. Gregorio, G. B., Senadhira, D., Htut, T. & Graham, R. D.    (2000)   Breed-
                 4. Rogers, E. E., Eide, D. & Guerinot, M. L.     (2000)   Altered selectivity in       ing for trace mineral density in rice. Food Nutr. Bull. 21: 382–386.
            an Arabidopsis metal transporter. Proc. Natl. Acad. Sci. U.S.A. 97: 12356–12360.                 17. Beebe, S., Gonzalez, A. V. & Rengifo, J.       (2000)   Research on trace
                 5. Curie, C., Alonso, J. M., Le Jean, M., Ecker, J. R. & Briat, J.-F.                  minerals in common bean. Food Nutr. Bull. 21: 387–391.
            (2000)   Involvement of Nramp1 from Arabidopsis thaliana in iron transport.                      18. Romheld, V. (1987) Different strategies for iron acquisition in higher
            Biochem. J. 347: 749–755.                                                                   plants. Physiol. Plant 70: 231–234.
                 6. Thomine, S., Wang, R., Ward, J. M., Crawford, N. M. & Schroeder,                         19. Wheal, M. S., Heller, L. I., Norvell, W. A. & Welch, R. M.     (2001)   Re-
            J. I.  (2000)   Cadmium and iron transport by members of a plant metal                      verse-phase liquid chromatographic determination of phytometallophores from
            transporter family in Arabidopsis with homology to Nramp genes. Proc. Natl.                 Strategy II Fe-uptake species by 9-fluorenylmethyl chloroformate fluorescence.
            Acad. Sci. U.S.A. 97: 4991–4996.                                                            J. Chromatogr. A 942: 177–183.
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...Comparative trace element nutrition uptake and distribution in plants robin d graham james c r stangoulis department of plant science university adelaide waite campus south australia abstract therearesimilarities between mammals the absorption transport elements chemistry from food sources both cases is based on thermodynamics adsorption charged solid surfaces embedded a solution phase ions metal binding ligands downloaded https academic oup com jn article s by guest january together with redox systems case iron some other constitutive function nutrient during normal conditions inducible turbo increase supply particular deciency most studied micronutrients divides kingdom into two groups dicotyledonous have system that an upregulated version which consists membrane bound reductase atp driven hydrogen ion extrusion pump monocotyledonous similar to dicots but remarkably different usesthemugeneicacidclassofphytosiderophores ps thepssystemmayinfactbeanimportantportofentryfor rich exceeding...

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