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Plant Nutrition 3: Micronutrients and Metals

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Plant Nutrition 3: Micronutrients and Metals Plant Nutrition 3: Micronutrients and Metals Non-metals Essential Metalloids micronutrients Metals D-block metals Non-essential toxic elements (examples) Essential for animals, beneficial for www.plantcell.org/cgi/doi/10.1105/tpc.109.tt1009plants © 2015 American Society of Plant Biologists Lesson outline • Introduction to micronutrients The hydroponics system developed by Hoagland and Arnon for the characterization of micronutrients • Micronutrient transporters and transport • Essential metal micronutrients • Fe, Zn, Cu, Mn, Mo, Ni • Metal tolerance and metal hyperaccumulation • Toxic metals and metalloids • As, Cd, Al • Other micronutrients • B, Cl, Si, Se • Summary and ongoing research Dennis Hoagland and colleagues developed a soil- free system for micronutrient studies. Today, “Hoagland’s Solution” continues to be used as a complete plant nutrient solution. Adapted from Hoagland, D.R., and Arnon, D.I. (1950).The water-culture method for growing plants without soil. Circular. California Agricultural Experiment Station. Volume 347. 2nd edition. © 2015 American Society of Plant Biologists Micronutrients are essential in at least one plant taxon “An element is not considered essential Or… “the plant can be so severely deficient unless a deficiency of it in the element that it exhibits abnormalities makes it impossible for in growth, development, or reproduction, i.e. the plant to complete its "performance," compared to plants with a life cycle” Mn lower deficiency” -Arnon and Stout, 1939 B -Epstein and Bloom, 2003 Zn Cu Fe Mo Cl 1969 Ni 1920 - 1954 1987 1860 Si 1940 (Required for Equisetum) Year when each element was shown to be essential Arnon, D.I., and Stout, P.R. (1939). The essentiality of certain elements in minute quantity for plants with special reference to copper. Plant Physiol. 14: 371 – 375; Epstein, E. and Bloom, A.J. (2003). Mineral Nutrition of Plants: Principles and Perspectives, 2nd Ed., John Wiley & Sons, New York. See also Marschner, P., ed (2012). Marschner’s Mineral Nutrition of Higher Plants. 3rd ed. (London: Academic Press). © 2015 American Society of Plant Biologists Nutrient availability in soil is highly dependent on soil pH Micronutrient availability There are exceptions, but most plants grow best at a slightly acid pH where the availability of nutrients is generally high. In strongly acidic soils, growth is often limited by increased solubility of the abundant but detrimental non-nutritive element aluminum Aluminum Example: Iron is more soluble in acidic soils + 3+ Fe(OH)3 + 3H Fe + 3H2O Strongly Mildly Insoluble Soluble acidic alkaline © 2015 American Society of Plant Biologists Most micronutrients have a narrow optimal concentration range Essential nutrient Non-essential element Optimal range Deficiency Luxury (plants (plants can can store or adjust) detoxify) Limitation Toxicity Tolerance Toxicity Plant dry weight dry Plant Plant dry weight dry Plant Concentration of element Concentration of element Adapted from Lin, Y.-F. and Aarts, M.M. (2012). The molecular mechanism of zinc and cadmium stress response in plants. Cellular and Molecular Life Sciences. 69: 3187-3206 and Merchant, S.S., and Helmann, J.D. (2012). Elemental economy: Microbial strategies for optimizing growth in the face of nutrient limitation. Adv. Microb. Physiol 60: 91 – 210. © 2015 American Society of Plant Biologists Essential micronutrients are normally found in small amounts Element Biologically Concentration in plant relevant form in (mg / kg) plants (These values vary by species, other nutrient levels etc.) Deficiency Normal Toxicity Iron (Fe) Fe2+, Fe3+ < 20 20 – 1000 > 2000 For comparison, Copper (Cu) Cu+, Cu2+ < 10 10 – 25 > 25 concentrations of Zinc (Zn) Zn2+ < 10 10 – 120 > 120 macronutrients Manganese (Mn) Mn2+, Mn3+, Mn4+ < 90 90 – 200 > 200 range from 1000 – Molybdenum Mo4+, Mo6+ (in < 0.1 0.1 – 90 > 90 450,000 mg / kg) (Mo) Moco or FeMoco) Boron B(OH)3 < 10 10 – 80 > 80 Chloride Cl- > 100 100 – 800 > 800 Nickel (Ni) Ni2+ > 0.05 0.05 – 10 > 10 Palmer, C.M. and Guerinot, M.L. (2009). Facing the challenges of Cu, Fe and Zn homeostasis in plants. Nat Chem Biol. 5: 333-340. See also Marschner, P., ed (2012). Marschner’s Mineral Nutrition of Higher Plants. 3rd ed. (London: Academic Press) and Krämer, U. (2010). Metal hyperaccumulation in plants. Annu. Rev. Plant Biol. 61: 517-534. © 2015 American Society of Plant Biologists Plants respond to nutrient deficiency and nutrient excess Limitation Deficiency Luxury Toxicity Optimal range Plant dry Plant weight Concentration of element Increased Efflux to Alternative uptake: apoplast metabolic (upregulated pathways transporters, Storage secretion of (organelle) Recycling siderophores and chelators) Chelation / sequestration Adapted from Lin, Y.-F. and Aarts, M.M. (2012). The molecular mechanism of zinc and cadmium stress response in plants. Cellular and Molecular Life Sciences. 69: 3187-3206 and Merchant, S.S., and Helmann, J.D. (2012). Elemental economy: Microbial strategies for optimizing growth in the face of nutrient limitation. Adv. Microb. Physiol 60: 91 – 210. © 2015 American Society of Plant Biologists Nutrient deficiency characteristics illustrate nutrient function Zinc is required for growth Iron and Mn are both particularly of young needed for tissues. Deficiency chlorophyll causes stunting and the production. formation of small leaves (“little leaf”) Deficiency of either of these poorly mobile elements causes interveinal chlorosis (yellowing) Copper is required for cell energetics including photosynthesis and respiration. Deficiency symptoms include chlorosis and necrosis of young tissues and leaf tips Photo credits: William M. Ciesla, Forest Health Management International, Bugwood.org; CropNutrition © 2015 American Society of Plant Biologists Metal micronutrients: What’s so special about metals? Metal micronutrients Metals D-block metals © 2015 American Society of Plant Biologists Most metal micronutrients are important cofactors for enzymes Bar height = % of cellular proteins Enzymes that use Fe, Cu or Mo Macro- requiring element as cofactor are mainly oxidoreductases nutrients (blue bars) Micronutrients Enzymes that use Mn, or Zn have more diverse functions: These data are obtained from a Yellow= transferases; purple= protein database representing hydrolase; red= lyase; green= isomerase; grey= ligase multiple organisms Reprinted by permission from Macmillan Publishers Ltd: Waldron, K.J., Rutherford, J.C., Ford, D. and Robinson, N.J. (2009). Metalloproteins and metal sensing. Nature. 460: 823-830. © 2015 American Society of Plant Biologists Many metal nutrients are redox active: necessary but dangerous − Fenton reaction About ¼ of cellular proteins e (n)+ are metalloproteins (not M + H2O2 counting those that make loose associations with Mg) M(n)+ M(n+1)+ M(n+1)+ + HO· + OH− − HO· (hydroxyl radical) is highly reactive e− Several metals can alternate between being electron donors and acceptors, Uncontrolled metal reactions can damage extending the range of cells for example through reactions that can occur in free radical production The apoprotein is biological systems. the polypeptide part without the metal This particularly applies to Fe(II) and Fe (III) and Cu(I) and Cu(II) © 2015 American Society of Plant Biologists Nutrients move into and through the plant through the apo- and symplast and through membrane transporters Root vacuole Shoot vacuole Shoot & other Root apoplast & other apoplast pool organelle pools pool organelle pools Efflux into Uptake (influx) xylem Bioavailable into root apoplast Shoot pool in Root symplast rhizosphere symplast pool Efflux from pool Xylem Influx into root apoplast symplast pool Small molecules Phloem Sink tissues Effects of released from symplast (e.g., seeds) soil root to increase pool microbes nutrient availability Indicates that a transporter is required © 2015 American Society of Plant Biologists Nutrients move by symplastic, transcellular or apoplastic pathways Nutrients must be Membrane transport can transported across Cross membrane consume 1/3 of a cell’s into living cell in metabolic energy (or more) membranes to enter root hair the plant Symplastic or transcellular pathway Casparian strip Bidirectional transport Pumps, channels Apoplastic pathway between xylem parenchyma and carriers are the Cross membrane cells and apoplastic molecular into living cell in transpiration stream mediators of these at endodermis processes © 2015 American Society of Plant Biologists Long-distance transport of nutrients involves xylem and phloem Zn, Cd Mn A nutrient Cu transported solely by transpiration, would accumulate in leaves (sites of most transpiration) Nutrients need to move into phloem (through transporters) for distribution to young growing tissues M(n)+ Metal micronutrients are bound by chelators (small molecules or proteins) during transport Reprinted from Yamaji, N. and Ma, J.F. (2014). The node, a hub for mineral nutrient distribution in graminaceous plants. Trends Plant Sci. 19: 556-563 with permission from Elsevier. © 2015 American Society of Plant Biologists Membrane transporters are crucial for metal element homeostasis Apoplast to cytosol to vacuole and vice versa Soil to root Into and out of apoplastic xylem Reprinted from Clemens, S., Palmgren, M.G. and Krämer, U. (2002). A long way ahead: understanding and engineering plant metal accumulation. Trends Plant Sci. 7: 309-315 with permission from Elsevier. © 2015 American Society of Plant Biologists Metal transporter families Most metal NRAMPs. Metal influx transporters
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