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:
[1] Alberts, I. L., K. Nadassy & S. J.Wodak.1998.Analysis of zinc binding sites in
protein crystal structures. Prot. Sci.7: 17001716.
[2] Apel, K. and H. Hirt. 2004. Reactive oxygen species: metabolism, oxidative
stress, and signal transduction. Annu. Rev. Plant Biol. 55: 373399.
[3]Asada, K. and M.Takahashi. 1987. Production and Scavenging of Active Oxygen in
Photosynthesis, Kyle, D. J. C. J. Osmond and C. J. Artzen. Elsevier, Amsterdam, pp:
227-287.
[4]Ayia, Saradhi, P. and P. Mohanty. 1997. Involvement of proline in protecting thylakoid
membranes against free radical-induced photodamage. J. Photochem. Photobiol. B.
38: 253-257.
[5] Baker A. M., P.L.Walker.1990. Ecophysiology of metal uptake by tolerant
plants,heavy metal tolerance in plants.pp.155177.In: A.J. Shaw (ed.), Evolutionary
aspects.CRC, Boca Raton.
[6] Blaudez, D.2000. Cadmium uptake and subcellular compartmentation in the
ectomycorrhizal fungus Paxillusinvolutus. Microbiology-UK146: 1109-1117.
[7] Cobbett, C. and P. Goldsbrough. 2002. Phytochelatins and metallothioneins:
Roles in heavy metal detoxification and homeostasis. Annu. Rev. Plant Biol. 53:
159182.
[8] Clemens, S. 2006. Toxic metal accumulation, responses to exposure and mechanisms
of tolerance in plants. Biochimie. 88: 17071719
[9] Detz K-J, Baier M, Kramer U. 1999. Free radicals and reactive oxygen species as
mediators
of
heavy metals toxicity in Plants. in: Prasad MNV, HagemeyerrJ,
eds. Heavy metal stress
in plants: from molecules to ecosystems. Berlin:
SpringerVerlag.73-97.
[10]Di Vos, C.and H. Schat. 1991. Increased resistance to copperinduced damage of the
root cell plasmalemma in copper tolerant Silene cucubalus. Physiologia
Plantarum.82,523-528.
[11]Ernst, W., J.A. Verkleij, H .Schat.
ActaBotanicaNeerlandica. 41: 229-248.

1992.,

Metal

tolerance in

lants.

[12]

Evans, K. M., J. A. Gatehouse, W. P. Lindsay, J. Shi, A. M. Tommey and A. J.

Robinson.1992.Expression of the pea metallothionein-like gene PsMTA in
Escherichia coli and Arabidopsis thaliana and analysis of trace metal ion
accumulation: implications for PsMTA function. Plant Mol. Biol. 20: 10191028.

[13]

Freisinger, E. 2008. Plant MTs- Long neglected members of the metallothionein

superfamily. Dalton trsns. 47: 66636675

[14]GonzalezChavez, M., C. Carrillo-Gonzalez, R., Wright, S. F and Nichols, K. A.

2004. the role of glomalin, a protein produced by arbuscularmycorrhizal fungi, in
sequestrering potentially toxic elements. EnvironPollut. 130:317-323.
[15] Hall, J. L. 2002. Cellular mechanisms for heavy metal detoxification and
tolerance. J. Exp. Bot. 53: 111.
[16]Halliwell.B.1987. Oxidative damage ,lipid peroxidation and antioxidant protection
in chloroplast. Chem. Phys. Lipids,44:327-340.
[17]Hendry, G. A., Baker, F. 1992. Cadmium tolerance and toxicity,oxygen radical
processes and molecular damage in cadmium-sensitive clones of Holcuslanatus
L,Acta Bot Neerl. 41:271-281.
[18] Hsieh, H.M., W.K. Liu, P.C. Huang.1995. A novel stress-inducible metallothioneinlike gene from rice. Plant Mol Biol 28:381389.
[19] Hyun, H., Chang, A.C., Parker, D.R., and Page, A.L. 1998. Cadmium solubility
and phytoavailability in sludge treated soils: effect of soil organic carbon. Journal
of Environmental Quality 27: 329-334.
[20]Jentschke,
G.,
Godbold.
DL.
2000.
Metal
toxicity
and
ectomycorrhizas.PhysiologiaPlantarum 109:107-116.
[21] Katzung, B.G. 2004. Basic & Clinical Pharmacology. 9th ed., Naslefara.,Tehran.
[22]Lewis, S., R.D. Handy, Depledge, M.H. 1999. Sressed proteins (HSPs): methods of
detection and their use as an environmental biomarker. Ecotoxicology 8:351-368.
[23]Ma, J.F.,Delhaize, E. 2001. Aluminium tolerance in plants and the complexing role
of organic acids. Trends in Plant Science. 6:273-278.
[24]Ma, J.F., Mastsumoto H. 1997. Detoxiifyying aluminium with buckwheat. Nature
390:569-570.
[25] Margoshes, M. and B. L.Vallee .1957. A cadmiumprotein from equine kidney
cortex. J. Chem. Soc. 79: 48134819.
[26]Marschner, H. 1995. Mineral nutrition of higher plants,2ndedn.London:Academic
Press.
[27] Memon A.R., M. Yatazawa .1984 . Nature of manganese complexes in Mn
accumulator plantAcanthopanax sciadophylloides. J. Plant Nutr. 7:961974.
[28]Mittler, R. 2002. Oxidative stress antioxidants and stress tolerance. Trends Plant
Sci.7:405-410.
[29]Neumann, D., O. Lichtenberger.1994. Heat- shock proteins induce heavy metal
concentrations? Journal of Plant Physiology .146:704-717.

[29] Pal, M., E. Horvath, T. Janda, E. Paldi, and G. Szalai. 2006. Physiological
changes and defense mechanisms induced by cadmium stress in maize, J. Plant
Nutr. Soil Sci., 169:239-246.

[30]Persans,M. 1999. Molecular dissection of the role of histidin in nickel

hyperaccumulation in Thlaspigoesingense (Halacsy).plant physiology. 121,11171126.
[31] Pruvot, C. & F. Douay.2006.Heavy metals in soil, Crops and Grass as a source of
Human Exposure in the Former Mining Areas. Journal of Soil Sediments. 6: 215220.
[32]Rauser, W. E. 1999. Structure and function of metal chelators produced by plants
the case for organic acids, amino acids, phytin and metallothioneins. Cell
Biochemistry and Biophysics. 31: 19-48.
[33]Robinson NJ, A.M. Tommey, C. Kuske, Jackson PJ.1993. Plant metallothioneins.
Biochemical Journal. 295: 1-10.
[34] Rye, L., S. Paul, D. Bera, P. Chattopadhyay.2005. Bioaccumulation of Pb(II) from
Aqueous Solutions by Bacillus Cereus M116. Journal of Hazardous Substance
Research 5:1-21.
[35] Shuklasr, p.2005. Adsorption of Cu(II ), Ni(II ),and Zn (II) on modified jute
fibers.Bioresource Technology. 6:1430-38.
[36] Strange J, Macnair MR. 1991. Evidence for a role for the cell membrane in
copper tolerance of Mimulusguttatus Fischer ex DC. New Phytologist .119, 383388.
[37] Vijayaraghavan K,Y.Yun.2008. Bacterial Biosorbents and Biosorption..Journal of
Biotechnology Advances 26:266-291.
[38]Weissenhorn, I.1995. Arbuscularmycorrhizal contribution to heavy-metal uptake by
maize (Zea mays L) in pot culture with contaminated soil. Mycorrhiza. 5: 245-251.
[39] Zazouli M.A., A.M. Bandpei, A. Maleki, M. H. Saberian.2010.Determination of
cadmium and lead contents in black tea and tea liquor from Iran .Asia Journal
of Chemistry.22:1387- 93.