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    As(V), arsenate, is an analog of phosphate and is taken up into plant roots by the high-affinity phosphate uptake system (Asher & Reay, 1979).  Arsenic can be toxic

    • directly (as As(V) ) by substituting for phosphate in phosphorylation reactions
    • after reduction to As(III), arsenite.  Arsenite is more toxic due to its high affinity for protein thiolds (Jocelyn, 1972).  Arsenite is also taken up directly via aquaglyceroporins( Meharg and Jardine, 2003; Wysocki et al., 2001)

    Arsenic tolerant plants often sho reduced As(V) uptake, but also accumulate arsenic in their tissues (De Koe, 1994), suggesting some kind of internal sequestration (Bleeker et at, 2006).

    • (Matthew, 2011) Arsenate is taken up via the P transporters in P. vittata (Wang et al., 2002)
    • (Matthews, 2011) The transporters involved in AsIII uptake are unknown in P. vittata, but research in a number of plants and microbes shows that AsIII is taken up via aquaglyceroporins (Meng et al., 2004; Isayenkov and Maathuis, 2008).  Note Meng reference is E. coli., Isayenkov is Arabidopsis, neither is Pteris.
    • (Matthews, 2011) I was able to take up both AsIII and AsV and translocated them from the roots to the fronds, and, regardless of arsenic species supplied, AsV dominated in the roots while AsIII dominated in the fronds. This would indicate the presence of an arsenite oxidizing enzyme in the roots, and arsenate reducing enzyme in the fronds of the fern.  Judging by fig 3-4, AS(III) is taken up much more rapidly and efficiently from the growth medium (which becomes totally As(III) depleted).
    • (Matthews, 2011) arsenate reductase and cytosolic triosephosphate isomerase from the fronds were previously implicated in arsenate reduction (Ellis et al., 2006; Rathinasabapathi et al., 2006), others have reported arsenate reductase activities in protein extracts from the roots with rhizomes (Duan et al., 2005; Liu et al., 2009).
    • (Matthews, 2011) Unlike other plants, hyperaccumulators of As contain AsIII as uncomplexed species due to low phytochelatin content in tissue (Raab et al., 2004). It is believed that P. vittata takes up AsIII and AsV by the roots (Wang et al., 2002; Fayiga et al., 2005), translocates AsIII and AsV from the roots to fronds (Kertulis-Tartar et al., 2005; Singh and Ma, 2006), reduces AsV to AsIII in the fronds (Bondada et al., 2004; Tu et al., 2004b), and transports AsIII into vacuoles for storage (Lombi et al., 2002).

    Detoxification (from Bleeker, 2006)

    • As(V) to As(III) reduction is essential.  As(III) can form complexes with low MW metal -binding thiol peptides such as glutathione (GSH) or phytochelatins (PCs).
    • transporters that move As from cytoplasm to vacuole use AS(III) as substrate as a tri-glutathione complex (Gosh et al., 1999; Rosen, 2002)
    • in plant cells non enzymatic reduction in plant cells is  slow, plant genomes contain a homolog to the yeast As(V) reductase, Acr2. Duan et al. (2005), demostarated enzymatic, i.e., non GSH dependent, reduction of As(V) in Pteris vittata roots.  Enzyme may be induced by exposure to As.
    • PCs are metal binding peptides:(gamma-Glu-Cys)n-Gly (n=2 to 11) synthesized from GSH by phytochelatin synthase, PCS, a gamma-glutamate-cysteine transpeptidase. Phytochelatin synthase-deficient Arabidopsis
      thaliana is hypersensitive to Cd, mercury (Hg) and As(V), showing that PCs are absolutely required for As(V) tolerance in this species (Ha et al., 1999; Howden et al., 1995).
    • Charge neutral,ATP-dependent and DpH-independent transport of the complex suggests transport via an ATP-binding cassette(-)type (ABC) transporter (Bleeker, 2006)

    Prediction(Bleeker, 2006)

    As tolerance should increase with
    • level of expression of As(V) reductase
    • increase in GSH
    • increased PC levels


    "rhodanese/Cdc25-like", homologs present in rice (OsAsr1, OsAsr2), arabidopsis (AtAsr), maize (ZmAsr), and velvetgrass (HlAsr)



    • Asher, D.J. and Reay, P.F. (1979) Arsenic uptake by barley seedlings. J. Plant Physiol. 6, 459–466.
    • Bleeker (2006)
    • Duan, G., Y. Zhu, Y. Tong, C. Cai, and R. Kneer. 2005. Characterization of arsenate reductase in the extract of roots and fronds of Chinese Brake Fern, an arsenic hyperaccumulator. Plant Physiology 138:461-469
    • Ellis, D.R., L. Gumaelius, E. Indriolo, I.J. Pickering, J.A. Banks, and D.E. Salt. 2006. A novel arsenate reductase from the arsenic hyperaccumulating fern Pteris vittata. Plant Physiology 141:1544-1554.
    • Isayenkov, S.V., and F.J.M. Maathuis. 2008. The Arabidopsis thaliana aquaglyceroporin AtNIP7;1 is a pathway for arsenite uptake. FEBS Letters 582:1625-1628.
    • Jocelyn, P.C. (1972) Biochemistry of the SH Group. London: Academic Press.
    • Meharg, A.A. and Jardine, L. (2003) Arsenite transport into paddy rice (Oryza sativa) roots. New Phytol. 157, 39–44.
    • Liu, Y., H.B. Wang, M.H. Wong, and Z.H. Ye. 2009. The role of arsenate reductase and superoxide dismutase in As accumulation in four Pteris speices. Environ Int 35:4.Meng, Y.-L., Z. Liu, and B.P. Rosen. 2004. AsIII and SbIII uptake by GlpF and efflux by ArsB in Escherichia coli. J. Biol. Chem. 279:18334-18341.
    • Rathinasabapathi, B., S. Wu, S. Sundaram, J. Rivoal, M. Srivastava, and L.Q. Ma. 2006. Arsenic resistance in Pteris vittata L: identification of a cytosolic triosephosphate isomerase based on cDNA expression cloning in Escherichia coli. Plant Molecular Biology 62:845-857.
    • Wysocki, R., Chery, C.C., Wawrzycka, D., Van Hulle, M., Cornelis, R.,Thevelein, J.M. and Tamas, M.J. (2001) The glycerol channel Fps1p mediates the uptake of arsenite and antimonite in Saccharomyces cerevisiae. Mol. Microb. 40, 1391–1401.
    • Wang, J., Z. F, M. AA, and R. A. 2002. Mechanisms of arsenic hyperaccumulation in Pteris vittata: uptake kinetics, interactions with phosphate, and arsenic speciation. Plant Physiology 130:1552–1561.


    All pubmed As and Pteris

    1: Watanabe T, Kouho R, Katayose T, Kitajima N, Sakamoto N, Yamaguchi N, Shinano 
    T, Yurimoto H, Osaki M. Arsenic alters uptake and distribution of sulphur in
    Pteris vittata. Plant Cell Environ. 2013 Apr 23. doi: 10.1111/pce.12124. [Epub
    ahead of print] PubMed PMID: 23611758.
    2: Lessl JT, Ma L. Sparingly-soluble phosphate rock induced significant plant
    growth and arsenic uptake by Pteris vittata from three contaminated soils.
    Environ Sci Technol. 2013 Apr 22. [Epub ahead of print] PubMed PMID: 23607730.
    3: Wu F, Deng D, Wu S, Lin X, Wong MH. Arsenic tolerance, uptake, and
    accumulation by nonmetallicolous and metallicolous populations of Pteris vittata 
    L. Environ Sci Pollut Res Int. 2013 Mar 14. [Epub ahead of print] PubMed PMID:
    4: Müller K, Daus B, Mattusch J, Vetterlein D, Merbach I, Wennrich R. Impact of
    arsenic on uptake and bio-accumulation of antimony by arsenic hyperaccumulator
    Pteris vittata. Environ Pollut. 2013 Mar;174:128-33. doi:
    10.1016/j.envpol.2012.10.024. Epub 2012 Dec 17. PubMed PMID: 23257262.
    5: Lessl JT, Ma LQ, Rathinasabapathi B, Guy C. Novel Phytase from Pteris vittata 
    Resistant to Arsenate, High Temperature, and Soil Deactivation. Environ Sci
    Technol. 2013 Feb 20. [Epub ahead of print] PubMed PMID: 23379685.
    6: Hue NV. Arsenic chemistry and remediation in Hawaiian soils. Int J
    Phytoremediation. 2013;15(2):105-16. PubMed PMID: 23487989.
    7: Wan XM, Lei M, Liu YR, Huang ZC, Chen TB, Gao D. A comparison of arsenic
    accumulation and tolerance among four populations of Pteris vittata from habitats
    with a gradient of arsenic concentration. Sci Total Environ. 2013 Jan
    1;442:143-51. doi: 10.1016/j.scitotenv.2012.10.056. Epub 2012 Nov 22. PubMed
    PMID: 23178774.
    8: Mandal A, Purakayastha TJ, Patra AK, Sanyal SK. Phytoremediation of arsenic
    contaminated soil by Pteris vittata L. I. Influence of phosphatic fertilizers and
    repeated harvests. Int J Phytoremediation. 2012 Dec;14(10):978-95. PubMed PMID:
    9: Wei C, Zheng H, Yu J. Arsenic in the rhizosphere soil solution of ferns. Int J
    Phytoremediation. 2012 Dec;14(10):950-65. PubMed PMID: 22908657.
    10: Gonzaga MI, Ma LQ, Pacheco EP, dos Santos WM. Predicting arsenic
    bioavailability to hyperaccumulator Pteris vittata in arsenic-contaminated soils.
    Int J Phytoremediation. 2012 Dec;14(10):939-49. PubMed PMID: 22908656.
    11: Wang X, Rathinasabapathi B, de Oliveira LM, Guilherme LR, Ma LQ.
    Bacteria-mediated arsenic oxidation and reduction in the growth media of arsenic 
    hyperaccumulator Pteris vittata. Environ Sci Technol. 2012 Oct
    16;46(20):11259-66. doi: 10.1021/es300454b. Epub 2012 Oct 4. PubMed PMID:
    12: Forino LM, Ruffini Castiglione M, Bartoli G, Balestri M, Andreucci A,
    Tagliasacchi AM. Arsenic-induced morphogenic response in roots of arsenic
    hyperaccumulator fern Pteris vittata. J Hazard Mater. 2012 Oct 15;235-236:271-8. 
    doi: 10.1016/j.jhazmat.2012.07.051. Epub 2012 Aug 6. PubMed PMID: 22906843.
    13: Niazi NK, Singh B, Van Zwieten L, Kachenko AG. Phytoremediation of an
    arsenic-contaminated site using Pteris vittata L. and Pityrogramma calomelanos
    var. austroamericana: a long-term study. Environ Sci Pollut Res Int. 2012
    Sep;19(8):3506-15. doi: 10.1007/s11356-012-0910-4. Epub 2012 Apr 22. PubMed PMID:
    14: Mandal A, Purakayastha TJ, Patra AK, Sanyal SK. Phytoremediation of arsenic
    contaminated soil by Pteris vittata L. II. Effect on arsenic uptake and rice
    yield. Int J Phytoremediation. 2012 Jul;14(6):621-8. PubMed PMID: 22908631.
    15: Yan X, Zhang M, Liao X, Tu S. Influence of amendments on soil arsenic
    fractionation and phytoavailability by Pteris vittata L. Chemosphere. 2012
    Jun;88(2):240-4. doi: 10.1016/j.chemosphere.2012.03.015. Epub 2012 Mar 29. PubMed
    PMID: 22463947.
    16: Niazi NK, Kachenko AG. Letter to the editor regarding, "first evidence on
    different transportation modes of arsenic and phosphorus in arsenic
    hyperaccumulator Pteris vittata" by Lei et al. (2012). Environ Pollut. 2012
    Jun;165:167; author reply 168. doi: 10.1016/j.envpol.2011.11.031. Epub 2011 Dec
    9. PubMed PMID: 22169523.
    17: Drava G, Roccotiello E, Minganti V, Manfredi A, Cornara L. Effects of cadmium
    and arsenic on Pteris vittata under hydroponic conditions. Environ Toxicol Chem. 
    2012 Jun;31(6):1375-80. doi: 10.1002/etc.1835. Epub 2012 May 1. PubMed PMID:
    18: Lei M, Wan XM, Huang ZC, Chen TB, Li XW, Liu YR. First evidence on different 
    transportation modes of arsenic and phosphorus in arsenic hyperaccumulator Pteris
    vittata. Environ Pollut. 2012 Feb;161:1-7. doi: 10.1016/j.envpol.2011.09.017.
    Epub 2011 Oct 21. PubMed PMID: 22230060.
    19: Yang Q, Tu S, Wang G, Liao X, Yan X. Effectiveness of applying arsenate
    reducing bacteria to enhance arsenic removal from polluted soils by Pteris
    vittata L. Int J Phytoremediation. 2012 Jan;14(1):89-99. PubMed PMID: 22567697.
    20: de la Paix MJ, Lanhai L, de Dieu HJ, John MN. Plant algae method for arsenic 
    removal from arsenic contaminated groundwater. Water Sci Technol.
    2012;65(5):927-31. doi: 10.2166/wst.2012.875. PubMed PMID: 22339029.
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    accumulation by plants in an old antimony mine, China. Biol Trace Elem Res. 2011 
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    PMID: 21547400.
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    phenanthrene on their uptake and antioxidative response in Pteris vittata L.
    Environ Pollut. 2011 Dec;159(12):3398-405. doi: 10.1016/j.envpol.2011.08.045.
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    Bioresour Technol. 2011 Oct;102(20):9827-32. doi: 10.1016/j.biortech.2011.08.017.
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    arsenic in the growth media and increased growth of arsenic hyperaccumulator
    Pteris vittata L. Bioresour Technol. 2011 Oct;102(19):8756-61. doi:
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    Pityrogramma calomelanos var. austroamericana and Pteris vittata L. grown at a
    highly variable arsenic contaminated site. Int J Phytoremediation. 2011
    Oct;13(9):912-32. PubMed PMID: 21972513.
    30: Kumari A, Lal B, Pakade YB, Chand P. Assessment of bioaccumulation of heavy
    metal by Pteris vittata L. growing in the vicinity of fly ash. Int J
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    stress in Pteris vittata roots colonized or not by arbuscular mycorrhizal
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    arsenic and of metal uptake by aboveground tissues of Pteris vittata and Cyperus 
    involucratus growing in copper- and cobalt-rich tailings of the Zambian
    copperbelt. Arch Environ Contam Toxicol. 2011 Aug;61(2):228-42. doi:
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