• Mary Lou GuerinotEmail author
Part of the Plant Cell Monographs book series (CELLMONO, volume 17)


Fe deficiency commonly limits plant growth and crop yields. If the mechanisms of Fe uptake, distribution, and regulation were clearly understood, it might be feasible to engineer plants better able to grow in soils now considered marginal and to increase crop biomass in soils now in cultivation. Furthermore, plants that serve as better sources of this essential element would improve human nutrition because most people rely on plants as their dietary source of Fe. Here we review our current understanding of Fe homeostasis in plants, emphasizing the challenges of safely transporting and storing this essential redox-active metal.


Xylem Exudate Interveinal Chlorosis Iron Transport Protein Ferric Chelate Reductase Mitochondrial Ferritin 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Work in my laboratory is supported by grants from the National Science Foundation (IBN-0344305; IBN-0419695; DBI-0606193), the National Institutes of Health (RO1 GM 078536), the Department of Energy (DE-FG-2-06ER15809) and the National Institute of Environmental Health Sciences (5 P42 ES007373). I thank Christine Palmer for drawing the original figures on which the ones in this chapter are based.


  1. Aoyama T, Kobayashi T, Takahashi M, Nagasaka S, Usuda K, Kakei Y, Ishimaru Y, Nakanishi H, Mori S, Nishizawa NK (2009) OsYSL18 is a rice iron(III)-deoxymugineic acid transporter specificially expressed in reproductive organs and phloem of lamina joints. Plant Mol Biol 70:681–692PubMedGoogle Scholar
  2. Arnaud N, Murgia I, Boucherez J, Briat J-F, Cellier F, Gaymard F (2006) An iron-induced nitric oxide burst precedes ubiquitin-dependent protein degradation for Arabidopsis AtFer1 ferritin gene expression. J Biol Chem 281:23579–23588PubMedGoogle Scholar
  3. Arnaud N, Ravet K, Borlotti A, Touraine B, Boucherez J, Briat JF, Cellier F, Gaymard F (2007) The iron responsive element (IRE)/iron regulatory protein1 (IRP1)-cytosolic aconitase iron-regulatory switch does not operate in plants. Biochem J 405:523–531PubMedGoogle Scholar
  4. Balk J, Lobreaux S (2005) Biogenesis of iron-sulfur proteins in plants. Trends Plant Sci 10:324–331PubMedGoogle Scholar
  5. Bell WD, Bogorad L, McIlrath WJ (1958) Response of the yellow-stripe maize mutant (ys1) to ferrous and ferric iron. Bot Gaz 119:36–39Google Scholar
  6. Briat JF, Curie C, Gaymard F (2007) Iron utilization and metabolism in plants. Curr Opin Plant Biol 10:276–282PubMedGoogle Scholar
  7. Briat J-F, Ravet K, Arnaud N, Duc C, Boucherez J, Touraine B, Cellier F, Gaymard F (2009) New insights into ferritin synthesis and function highlight a link between iron homeostasis and oxidative stress in plants. An Bot (in press) doi: 10.1093/aob/mcp128
  8. Brown JC, Chaney RL, Ambler JE (1971) A new tomato mutant inefficient in the transport of iron. Physiol Plant 25:48–53Google Scholar
  9. Brumbarova T, Bauer P (2005) Iron-mediated control of the basic helix-loop-helix protein FER, a regulator of iron uptake in tomato. Plant Physiol 137:1018–1026PubMedGoogle Scholar
  10. Bughio N, Takahashi M, Yoshimura E, Nishizawa NK, Mori S (1997a) Characteristics of light-regulated iron transport system in barley chloroplasts. Soil Sci Plant Nutr 43:959–963Google Scholar
  11. Bughio N, Takahashi M, Yoshimura E, Nishizawa NK, Mori S (1997b) Light-dependent iron transport into isolated barley chloroplasts. Plant Cell Physiol 38:101–105Google Scholar
  12. Busi MV, Maliandi MV, Valdez H, Clemente M, Zabaleta EJ, Araya A, Gomez-Casati DF (2006) Deficiency of Arabidopsis thaliana frataxin alters activity of mitochondrial Fe-S proteins and induces oxidative stress. Plant J 48:873–882PubMedGoogle Scholar
  13. Carpenter BM, Whitmire JM, Merrell DS (2009) This is not your mother’s repressor: the complex role of Fur in pathogenesis. Infect Immun 77:2590–2601PubMedGoogle Scholar
  14. Chen OS, Crisp RJ, Valachovic M, Bard M, Winge DR, Kaplan J (2004) Transcription of the yeast iron regulon does not respond directly to iron but rather to iron-sulfur cluster biosynthesis. J Biol Chem 279:29513–29518PubMedGoogle Scholar
  15. Chen S, Sanchez-Fernandez R, Lyver ER, Dancis A, Rea PA (2007) Functional characterization of AtATM1, AtATM2 and AtATM3, a subfamily of Arabidopsis half-molecule ABC transporters implicated in iron homeostasis. J Biol Chem 282:21561–21571PubMedGoogle Scholar
  16. Cheng L, Wang F, Shou H, Huang F, Zheng L, He F, Li J, Zhao FJ, Ueno D, Ma JF, Wu P (2007) Mutation in nicotianamine aminotransferase stimulated the Fe(II) acquisition system and led to iron accumulation in rice. Plant Physiol 145:1647–1657PubMedGoogle Scholar
  17. Colangelo EP, Guerinot ML (2004) The essential bHLH protein FIT1 is required for the iron deficiency response. Plant Cell 16:3400–3412PubMedGoogle Scholar
  18. Connolly EL, Fett JP, Guerinot ML (2002) Expression of the IRT1 metal transporter is controlled by metals at the levels of transcript and protein accumulation. Plant Cell 14:1347–1357PubMedGoogle Scholar
  19. Connolly EL, Campbell N, Grotz N, Prichard CL, Guerinot ML (2003) Overexpression of the FRO2 iron reductase confers tolerance to growth on low iron and uncovers post-transcriptional control. Plant Physiol 133:1102–1110PubMedGoogle Scholar
  20. Curie C, Briat J-F (2003) Iron transport and signaling in plants. Annu Rev Plant Biol 54:183–206PubMedGoogle Scholar
  21. Curie C, Alonso JM, Le Jean M, Ecker JR, Briat J-F (2000) Involvement of NRAMP1 from Arabidopsis thaliana in iron transport. Biochem J 347:749–755PubMedGoogle Scholar
  22. Curie C, Panaviene Z, Loulergue C, Dellaporta SL, Briat J-F, Walker EL (2001) Maize yellow stripe1 encodes a membrane protein directly involved in Fe(III) uptake. Nature 409:346–349PubMedGoogle Scholar
  23. Curie C, Cassin G, Counch D, Divol F, Higuchi K, Le Jean M, Misson J, Schikora A, Czernic P, Mari S (2009) Metal movement within the plant: contribution of nictotianamine and yellow stripe 1-like transporters. Ann Bot 103:1–11PubMedGoogle Scholar
  24. Dello Ioio R, Linhares FS, Scacchi E, Casamitjana-Martinez E, Heidstra R, Costantino P, Sabatini S (2007) Cytokinins determine Arabidopsis root-meristem size by controlling cell differentiation. Curr Biol 17:678–682PubMedGoogle Scholar
  25. Dinneny JR, Long TA, Wang JY, Jung JW, Mace D, Pointer S, Barron C, Brady SM, Schiefelbein J, Benfey PN (2008) Cell identity mediates the response of Arabidopsis roots to abiotic stress. Science 320:942–945PubMedGoogle Scholar
  26. Durrett TP, Gassmann W, Rogers EE (2007) The FRD3-mediated efflux of citrate into the root vasculature is necessary for efficient iron translocation. Plant Physiol 144:197–205PubMedGoogle Scholar
  27. Duy D, Wanner G, Meda AR, von Wiren N, Soll J, Philippar K (2007) PIC1, an ancient permease in Arabidopsis chloroplasts, mediates iron transport. Plant Cell 19:986–1006PubMedGoogle Scholar
  28. Foury F, Roganti T (2002) Deletion of the mitochondrial carrier genes MRS3 and MRS4 suppresses mitochondrial iron accumulation in a yeast frataxin-deficient strain. J Biol Chem 277:24475–24483PubMedGoogle Scholar
  29. Furukawa J, Yamaji N, Wang H, Mitani N, Murata Y, Sato K, Katsuhara M, Takeda K, Ma JF (2007) An aluminum-activated citrate transporter in barley. Plant Cell Physiol 48:1081–1091PubMedGoogle Scholar
  30. Ganz T (2008) Iron homeostasis: fitting the puzzle pieces together. Cell Metab 7:288–290PubMedGoogle Scholar
  31. Gitan RS, Eide DJ (2000) Zinc-regulated ubiquitin conjugation signals endocytosis of the yeast ZRT1 zinc transporter. Biochem J 346:329–336PubMedGoogle Scholar
  32. Graziano M, Lamattina L (2005) Nitric oxide and iron in plants: an emerging and converging story. Trends Plant Sci 10:4–8PubMedGoogle Scholar
  33. Graziano M, Lamattina L (2007) Nitric oxide accumulation is required for molecular and physiological responses to iron deficiency in tomato roots. Plant J 52:949–960PubMedGoogle Scholar
  34. Graziano M, Beligni MV, Lamattina L (2002) Nitric oxide improves internal iron availability in plants. Plant Physiol 130:1852–1859PubMedGoogle Scholar
  35. Green LS, Rogers EE (2004) FRD3 controls iron localization in Arabidopsis thaliana. Plant Physiol 136:2523–2531PubMedGoogle Scholar
  36. Grotz N, Guerinot ML (2006) Molecular aspects of Cu, Fe and Zn homeostasis in plants. Biochim Biophys Acta 1763:595–608PubMedGoogle Scholar
  37. Grusak MA, Pezeshgi S (1996) Shoot-to-root signal transmission regulates root Fe(III) reductase activity in the dgl mutant of pea. Plant Physiol 110:329–334PubMedGoogle Scholar
  38. Guerinot ML (2000) The ZIP family of metal transporters. Biochim Biophys Acta 1465:190–198PubMedGoogle Scholar
  39. Halliwell B, Gutteridge JMC (1992) Biologically relevant metal ion-dependent hydroxyl radical generation. FEBS Lett 307:108–112PubMedGoogle Scholar
  40. Haydon MJ, Cobbett CS (2007) Transporters of ligands for essential metal ions in plants. New Phytol 174:499–506PubMedGoogle Scholar
  41. Heazlewood JL, Tonti-Filippini JS, Gout AM, Day DA, Whelan J, Millar AH (2004) Experimental analysis of the Arabidopsis mitochondrial proteome highlights signaling and regulatory components, provides assessment of targeting prediction programs, and indicates plant-specific mitochondrial proteins. Plant Cell 16:241–256PubMedGoogle Scholar
  42. Inoue H, Kobayashi T, Nozoye T, Takahashi M, Kakei Y, Suzuki K, Nakazono M, Nakanishi H, Mori S, Nishizawa NK (2009) Rice OsYSL15 is an iron-regulated iron(III)-deoxymugineic acid transporter expressed in the roots and is essential for iron uptake in early growth of the seedlings. J Biol Chem 284:3470–3479PubMedGoogle Scholar
  43. Ishimaru Y, Suzuki M, Tsukamoto T, Suzuki K, Nakazono M, Kobayashi T, Wada Y, Watanabe S, Matsuhashi S, Takahashi M, Nakanishi H, Mori S, Nishizawa NK (2006) Rice plants take up iron as an Fe3+-phytosiderophore and as Fe2+. Plant J 45:335–346PubMedGoogle Scholar
  44. Ishimaru Y, Kim SA, Tsukamoto T, Oki H, Kobayashi T, Watanabe S, Matsuhashi S, Takahashi M, Nakanishi H, Mori S, Nishizawa NK (2007) Mutational reconstructed ferric chelate reductase confers enhanced tolerance in rice to iron deficiency in calcareous soil. Proc Natl Acad Sci USA 104:7373–7378PubMedGoogle Scholar
  45. Jakoby M, Wang H-Y, Reidt W, Weissharr B, Bauer P (2004) FRU(BHLH029) is required for induction of iron mobilization genes in Arabidopsis thaliana. FEBS Lett 577:528–534PubMedGoogle Scholar
  46. Jeong J, Cohu C, Kerkeb L, Piln M, Connolly EL, Guerinot ML (2008) Chloroplast Fe(III) chelate reductase activity is essential for seedling viability under iron limiting conditions. Proc Natl Acad Sci USA 105:10619–10624PubMedGoogle Scholar
  47. Kerkeb L, Mukherjee I, Chatterjee I, Lahner B, Salt DE, Connolly EL (2008) Iron-induced turnover of the Arabidopsis IRT1 metal transporter requires lysine residues. Plant Physiol 146:1964–1973PubMedGoogle Scholar
  48. Kim SA, Punshon T, Lanzirotti A, Li L, Alonzo JM, Ecker JR, Kaplan J, Guerinot ML (2006) Localization of iron in Arabidopsis seed requires the vacuolar membrane transporter VIT1. Science 314:1295–1298PubMedGoogle Scholar
  49. Klatte M, Schuler M, Wirtz M, Fink-Straube C, Hell R, Bauer P (2009) The analysis of Arabidopsis nicotianamine synthase mutants reveals functions for nicotianamine in seed iron loading and iron deficiency responses. Plant Physiol 150:257–271PubMedGoogle Scholar
  50. Kobayashi T, Nakayama Y, Itai RN, Nakanishi H, Yoshihara T, Mori S, Nishizawa NK (2003) Identification of novel cis-acting elements, IDE1 and IDE2, of the barley IDS2 gene promoter conferring iron-deficiency-inducible, root-specific expression in heterologous tobacco plants. Plant J 36:780–793PubMedGoogle Scholar
  51. Kobayashi T, Ogo Y, Itai RN, Nakanishi H, Takahashi M, Mori S, Nishizawa NK (2007) The transcription factor IDEF1 regulates the response to and tolerance of iron deficiency in plants. Proc Natl Acad Sci USA 104:19150–19155PubMedGoogle Scholar
  52. Koike S, Inoue H, Mizuno D, Takahashi M, Nakanishi H, Mori S, Nishizawa N (2004) OsYSL2 is a rice metal-nicotianamine transporter that is regulated by iron and expressed in the phloem. Plant J 39:415–424PubMedGoogle Scholar
  53. Kruger C, Berkowitz O, Stephan UW, Hell R (2002) A metal-binding member of the late embryogenesis abundant protein family transports iron in the phloem of Ricinus communis L. J Biol Chem 277:25062–25069PubMedGoogle Scholar
  54. Kumanovics A, Chen OS, Li L, Bagley D, Adkins EM, Lin H, Dingra NN, Outten CE, Keller G, Winge D, Ward DM, Kaplan J (2008) Identification of FRA1 and FRA2 as genes involved in regulating the yeast iron regulon in response to decreased mitochondrial iron-sulfur cluster synthesis. J Biol Chem 283:10276–10286PubMedGoogle Scholar
  55. Kushnir S, Babiychuk E, Storozhenko S, Davey MW, Papenbrock J, De Rucke R, Engler G, Stephan UW, Lange H, Kispai G, Lill R, Van Monagu M (2001) A mutation of the mitochondrial ABC transporter Sta1 leads to dwarfism and chlorosis in the Arabidopsis mutant starik. Plant Cell 13:89–100PubMedGoogle Scholar
  56. Landsberg E (1984) Regulation of iron-stress-response by whole-plant activity. J Plant Nutr 7:609–621Google Scholar
  57. Lanquar V, Lelievre F, Bolte S, Hames C, Alcon C, Neumann D, Vansuyt G, Curie C, Schroder A, Kramer U, Barbier-Brygoo H, Thomine S (2005) Mobilization of vacuolar iron by AtNRAMP3 and AtNRAMP4 is essential for seed germination on low iron. EMBO J 24:4041–4051PubMedGoogle Scholar
  58. Lee S, An G (2009) Over-expression of OsIRT1 leads to increased iron and zinc accumulation in rice. Plant Cell Environ 32:408–416PubMedGoogle Scholar
  59. Lee S, Chiecko JC, Kim SA, Walker EL, Lee Y, Guerinot ML, An G (2009) Disruption of OsYSL15 leads to iron inefficiency in rice plants. Plant Physiol 150:786–800PubMedGoogle Scholar
  60. Levi S, Arosio P (2004) Mitochondrial ferritin. Int J Biochem Cell Biol 36:1887PubMedGoogle Scholar
  61. Li L, Kaplan J (2004) A mitochondrial-vacuolar signaling pathway in yeast that affects iron and copper metabolism. J Biol Chem 279:33653–33661PubMedGoogle Scholar
  62. Li L, Chen OS, Ward DM, Kaplan J (2001) CCC1 is a transporter that mediates vacuolar iron storage in yeast. J Biol Chem 276:29515–29519PubMedGoogle Scholar
  63. Lill R (2009) Function and biogenesis of iron-sulfur proteins. Nat Chem Biol 460:831–838Google Scholar
  64. Ling HQ, Koch G, Baulein H, Ganal MW (1999) Map-based cloning of chloronerva, a gene involved in iron uptake of higher plants encoding nicotianamine synthase. Proc Natl Acad Sci USA 96:7098–7103PubMedGoogle Scholar
  65. Ling H-Q, Bauer P, Bereczky Z, Keller B, Ganal M (2002) The tomato fer gene encoding a bHLH protein controls iron-uptake responses in roots. Proc Natl Acad Sci USA 99:13938–13943PubMedGoogle Scholar
  66. Liu J, Magalhaes JV, Shaff JE, Kochian LV (2009) Aluminum-activated citrate and malate transporters from the MATE and ALMT families function independently to confer Arabidopsis aluminum tolerance. Plant J 57:389–399PubMedGoogle Scholar
  67. Lucena C, Waters BM, Romera FJ, Garcia MJ, Morales M, Alcantara E, Perez-Vicente R (2006) Ethylene could influence ferric reductase, iron transporter, and H+-ATPase gene expression by affecting FER (or FER-like) gene activity. J Exp Bot 57:4145–4154Google Scholar
  68. Magalhaes JV, Liu J, Guimarães CT, Lana UG, Alves VM, Wang YH, Schaffert RE, Hoekenga OA, Piñeros MA, Shaff JE, Klein PE, Carneiro NP, Coelho CM, Trick HN, Kochian LV (2007) A gene in the multidrug and toxic compound extrusion (MATE) family confers aluminum tolerance in sorghum. Nat Genet 39:1156–1161PubMedGoogle Scholar
  69. Millar AH, Heazlewood JL (2003) Genomic and proteomic analysis of mitochondrial carrier proteins in Arabidopsis. Plant Physiol 131:443–453PubMedGoogle Scholar
  70. Missirlis F, Holmberg S, Georgieva T, Dunkov BC, Rouault TA, Law JH (2006) Characterization of mitochondrial ferritin in Drosophila. Proc Natl Acad Sci USA 103:5893–5898PubMedGoogle Scholar
  71. Moseley JL, Allinger T, Herzog S, Hoerth P, Wehinger E, Merchant S, Hippler M (2002) Adaptation to Fe-deficiency requires remodeling of the photosynthetic apparatus. EMBO J 21:6709–6720PubMedGoogle Scholar
  72. Muckenthaler MU, Galy B, Hentze MW (2008) Systemic iron homeostasis and the iron-responsive element/iron-regulatory protein (IRE/IRP) regulatory network. Annu Rev Nutr 28:197–213PubMedGoogle Scholar
  73. Muhlenhoff U, Stadler JA, Richhardt N, Seubert A, Eickhorst T, Schweyen RJ, Lill R, Wiesenberger G (2003) A specific role of the yeast mitochondrial carriers Mrs3/4p in mitochondrial iron acquisition under iron-limiting conditions. J Biol Chem 278:40612–40620PubMedGoogle Scholar
  74. Murgia I, Vazzola V, Tarantino D, Cellier F, Ravet K, Briat JF, Soave C (2007) Knock-out of ferritin AtFer1 causes earlier onset of age-dependent leaf senescence in Arabidopsis. Plant Physiol Biochem 45:898–907PubMedGoogle Scholar
  75. Nagasaka S, Takahashi M, Nakanishi-Itai R, Bashir K, Nakanishi H, Mori S, Nishizawa NK (2009) Time course analysis of gene expression over 24 hours in Fe-deficient barley roots. Plant Mol Biol 69:621–631PubMedGoogle Scholar
  76. Nakanishi H, Yamaguchi H, Sasakuma T, Nishizawa NK, Mori S (2000) Two dioxygenase genes, Ids3 and Ids2, from Hordeum vulgare are involved in the biosynthesis of mugineic acid family phytosiderophores. Plant Mol Biol 44:199–207PubMedGoogle Scholar
  77. Negishi T, Nakanishi H, Yazaki J, Kishimoto N, Fujii F, Shimbo K, Yamamoto K, Sakata K, Sasaki T, Kikuchi S, Mori S, Nishizawa N (2002) cDNA microarray analysis of gene expression during Fe-deficiency stress in barley suggests that polar transport of vesicles is implicated in phytosiderophore secretion in Fe-deficient barley roots. Plant J 30:83–94PubMedGoogle Scholar
  78. Ogo Y, Itai RN, Nakanishi H, Inoue H, Kobayashi T, Suzuki M, Takahashi M, Mori S, Nishizawa NK (2006) Isolation and characterization of IRO2, a novel iron regulated bHLH transcription factor in graminaceous plants. J Exp Bot 57:2867–2878PubMedGoogle Scholar
  79. Ogo Y, Itai RN, Nakanishi H, Kobayashi T, Takahashi M, Mori S, Nishizawa NK (2007) The rice bHLH protein OsIRO2 is an essential regulator of the genes involved in Fe uptake under Fe-deficient conditions. Plant J 51:366–377PubMedGoogle Scholar
  80. Ogo Y, Kobayashi T, Nakanishi Itai R, Nakanishi H, Kakei Y, Takahashi M, Toki S, Mori S, Nishizawa NK (2008) A novel NAC transcription factor, IDEF2, that recognizes the iron deficiency-responsive element 2 regulates the genes involved in iron homeostasis in plants. J Biol Chem 283:13407–13417PubMedGoogle Scholar
  81. Oki H, Kim S, Nakanishi H, Takahashi M, Yamaguchi H, Mori S, Nishizawa NK (2004) Directed evolution of yeast ferric reductase to produce plants with tolerance to iron deficiency in alkaline soils. Soil Sci Plant Nutr 50:1159–1165Google Scholar
  82. Palmer C, Guerinot ML (2009) Facing the challenges of Cu, Fe and Zn homeostasis in plants. Nat Chem Biol 5:333–340PubMedGoogle Scholar
  83. Raven JA, Evans MCW, Korb RE (1999) The role of trace metals in photosynthetic electron transport in O2-evolving organisms. Photosyn Res 60:111–149Google Scholar
  84. Ravet K, Touraine B, Boucherez J, Briat JF, Gaymard F, Cellier F (2008) Ferritins control interaction between iron homeostasis and oxidative stress in Arabidopsis. Plant J 57:400–412PubMedGoogle Scholar
  85. Ravet K, Touraine B, Kim SA, Cellier F, Thmine S, Guerinot ML, Briat J-F, Gaymard F (2009) Post-translational regulation of AtFER2 ferritin in response to intracellular iron trafficking during fruit development in Arabidopsis. Mol Plant 2:1095–1106PubMedGoogle Scholar
  86. Robinson NJ, Proctor CM, Connolly EL, Guerinot ML (1999) A ferric-chelate reductase for iron uptake from soils. Nature 397:694–697PubMedGoogle Scholar
  87. Romera FJ, Alcantara E (2004) Ethylene involvement in the regulation of Fe-deficiency stress responses by Strategy I plants. Func Plant Biol 31:315–328Google Scholar
  88. Rutherford JC, Ojeda L, Balk J, Mulenhoff U, Lill R, Winge DR (2005) Activation of the iron regulon by yeast Aft1/Aft2 transcription factors depends on mitochondrial but not cytosolic iron-sulfur protein biogenesis. J Biol Chem 280:10135–10140PubMedGoogle Scholar
  89. Santi S, Schmidt W (2009) Dissecting iron deficiency-induced proton extrusion in Arabidopsis roots. New Phytol 183:1072–1084PubMedGoogle Scholar
  90. Schikora A, Schmidt W (2001) Iron stress-induced changes in root epidermal cell fate are regulated independently from physiological responses to low iron availability. Plant Physiol 125:1679–1687PubMedGoogle Scholar
  91. Schwacke R, Schneider A, van der Graaff E, Fischer K, Catoni E, Desimone M, Frommer WB, Flugge U-I, Kunze R (2003) ARAMEMNON, a novel database for Arabidopsis Integral membrane proteins. Plant Physiol 131:16–26PubMedGoogle Scholar
  92. Seguela M, Briat J-F, Vert G, Curie C (2008) Cytokinins negatively regulate the root iron uptake machinery in Arabidopsis through a growth-dependent pathway. Plant J 55:289–300PubMedGoogle Scholar
  93. Shaw GC, Cope JJ, Li L, Corson K, Hersey C, Ackermann GE, Gwynn B, Lambert AJ, Wingert RA, Traver D, Trede NS, Barut BA, Zhou Y, Minet E, Donovan A, Brownlie A, Balzan R, Weiss MJ, Peters LL, Kaplan J, Zon LI, Paw BH (2006) Mitoferrin is essential for erythroid iron assimilation. Nature 440:96–100PubMedGoogle Scholar
  94. Shikanai T, Muller-Moule P, Munekage Y, Niyogi KK, Pilon M (2003) PAA1, a P-type ATPase of Arabidopsis, functions in copper transport in chloroplasts. Plant Cell 15:1333–1346PubMedGoogle Scholar
  95. Shingles R, North M, McCarty RE (2001) Direct measurement of ferrous ion transport across membranes using a sensitive fluorometric assay. Anal Biochem 296:106–113PubMedGoogle Scholar
  96. Shingles R, North M, McCarty RE (2002) Ferrous iron transport across chloroplast inner envelope membranes. Plant Physiol 128:1022–1030PubMedGoogle Scholar
  97. Stacey MG, Koh S, Becker J, Stacey G (2002) AtOPT3, a member of the oligopeptide transporter family, is essential for embryo development in Arabidopsis. Plant Cell 14:2799–2811PubMedGoogle Scholar
  98. Stacey MG, Osawa H, Patel A, Gassmann W, Stacey G (2006) Expression analyses of Arabidopsis oligopeptide transporters during seed germination. Plant J 223:291–305Google Scholar
  99. Stacey MG, Patel A, McClain WE, Mathieu M, Remley M, Rogers EE, Gassmann W, Blevins DG, Stacey G (2008) The Arabidopsis AtOPT3 protein functions in metal homeostasis and movement of iron to developing seeds. Plant Physiol 146:589–601PubMedGoogle Scholar
  100. Takahashi MT, Nakanishi H, Kawasaki S, Nishizawa NK, Mori S (2001) Enhanced tolerance of rice to low iron availability in alkaline soils using barley nicoianamine aminotransferase genes. Nat Biotechnol 19:466–469PubMedGoogle Scholar
  101. Teng YS, Su YS, Chen LJ, Lee YJ, Hwang I, Li HM (2006) Tic21 is an essential translocon component for protein translocation across the chloroplast inner envelope membrane. Plant Cell 18:2247–2257PubMedGoogle Scholar
  102. Terry N, Abadia J (1986) Function of iron in chloroplasts. J Plant Nutr 9:609–646Google Scholar
  103. Thomine S, Lelievre F, Debarbieux E, Schroeder JI, Barbier-Bygoo H (2003) AtNRAMP3, a multispecific vacuolar metal transporter involved in plant responses to iron deficiency. Plant J 34:685–695PubMedGoogle Scholar
  104. Varotto C, Maiwald D, Pesaresi P, Jahns P, Francesco S, Leister D (2002) The metal ion transporter IRT1 is necessary for iron homeostasis and efficient photosynthesis in Arabidopsis thaliana. Plant J 31:589–599PubMedGoogle Scholar
  105. Vasconcelos M, Eckert H, Arahana V, Graef G, Grusak MA, Clemente T (2006) Molecular and phenotypic characterization of transgenic soybean expressing the Arabidopsis ferric chelate reductase gene, FRO2. Planta 224:1116–1128PubMedGoogle Scholar
  106. Vazzola V, Losa A, Soave C, Murgia I (2007) Knockout of frataxin gene causes embryo lethality in Arabidopsis. FEBS Lett 581:667–672PubMedGoogle Scholar
  107. Vert G, Grotz N, Dedaldechamp F, Gaymard F, Guerinot ML, Briat J-F, Curie C (2002) IRT1, an Arabidopsis transporter essential for iron uptake from the soil and plant growth. Plant Cell 14:1223–1233PubMedGoogle Scholar
  108. Vert G, Briat J-F, Curie C (2003) Dual regulation of the Arabidopsis high affinity root iron uptake system by local and long-distance signals. Plant Physiol 132:796–804PubMedGoogle Scholar
  109. Vert G, Barberon M, Zelazny E, Seguela M, Briat J-F, Curie C (2009) Arabidopsis IRT2 cooperates with the high affinity iron uptake system to maintain iron homeostasis in root epidermal cells. Planta 229:1171–1179PubMedGoogle Scholar
  110. von Wiren N, Klair S, Bansal S, Briat J-F, Khodr H, Shioiri T, Leigh RA, Hider RC (1999) Nicotianamine chelates both Fe(III) and Fe(II). Implications for metal transport in plants. Plant Physiol 119:1107–1114Google Scholar
  111. von Wirén N, Mori S, Marschner H, Römheld V (1994) Iron inefficiency in maize mutant ys1 (Zea mays L. cv yellow-stripe) is caused by a defect in uptake of iron phytosiderophores. Plant Physiol 106:71–77Google Scholar
  112. Walker EL, Connolly EL (2008) Time to pump iron: iron-deficiency-signaling mechanisms of higher plants. Curr Opin Plant Biol 11:530–535PubMedGoogle Scholar
  113. Wang H-Y, Klatte M, Jacoby M, Baumlein H, Weisshaar B, Bauer P (2007) Iron deficiency-mediated stress regulation of four subgroup Ib bHLH genes in Arabidopsis thaliana. Planta 226:897–908PubMedGoogle Scholar
  114. Waters BM, Chu HH, Didonato RJ, Roberts LA, Eisley RB, Walker EL (2006) Mutations in Arabidopsis Yellow Stripe-Like1 and Yellow Stripe-Like3 reveal their roles in metal ion homeostasis and loading of metal ions in seeds. Plant Physiol 141:1446–1458Google Scholar
  115. Waters BM, Lucena C, Romera FJ, Jester GG, Wynn AN, Rojas CL, Alcántara E, Pérez-Vicente R (2007) Ethylene involvement in the regulation of the H(+)-ATPase CsHA1 gene and of the new isolated ferric reductase CsFRO1 and iron transporter CsIRT1 genes in cucumber plants. Plant Physiol Biochem 45:293–301Google Scholar
  116. Yokosho K, Yamaji N, Ueno D, Mitani N, Ma JF (2009) OsFRDL1 is a citrate transporter required for efficient translocation of iron in rice. Plant Physiol 149:297–305PubMedGoogle Scholar
  117. Yuan YX, Zhang J, Wang DW, Ling HQ (2005) AtbHLH29 of Arabidopsis thaliana is a functional ortholog of tomato FER involved in controlling iron acquisition in strategy I plants. Cell Res 15:613–621PubMedGoogle Scholar
  118. Yuan Y, Wu H, Wang N, KLi J, Zhao W, Du J, Wang D, Ling H-Q (2008) FIT interacts with AtbHLH38 and AtbHLH39 in regulating iron uptake gene expression for iron homeostasis in Arabidopsis. Cell Res 18:385–397PubMedGoogle Scholar
  119. Zancani M, Peresson C, Biroccio A, Federici G, Urbani A, Murgia I, Soave C, Micali F, Vianello A, Macr F (2004) Evidence for the presence of ferritin in plant mitochondria. Eur J Biochem 271:3657–3664PubMedGoogle Scholar
  120. Zhang Y, Lyver ER, Knight SAB, Pain D, Lesuisse E, Dancis A (2006) Mrs3p, Mrs4p, and frataxin provide iron for Fe-S cluster synthesis in mitochondria. J Biol Chem 281:22493–22502PubMedGoogle Scholar

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© Springer-Verlag Berlin Heidelberg 2010

Authors and Affiliations

  1. 1.Department of Biological SciencesDartmouth CollegeHanoverUSA

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