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. 2015 Sep 14;6:716. doi: 10.3389/fpls.2015.00716

Table 1.

Description of mutant alleles exhibiting de-regulated iron (Fe) signaling.

Organism/
Gene (allele)
Function Cellular localization/tissue expression Expression of FRO2/IRT1 Fe concentration in the mutant vs. wild-type Treatments that rescue Fe deficiency symptoms Reference
Arabidopsis thaliana /FRD3
(frd3)
Citrate efflux Plasma membrane/
Root pericycle
Constitutive expression of FRO2 and IRT1 High Fe accumulation in apoplast and vasculature, but less Fe in cytosol Foliar application of Fe; grafting frd3 shoots onto Wt roots; citrate addition to growth media Rogers and Guerinot, 2002; Green and Rogers, 2004; Garcia et al., 2013
A. thaliana
/NAS1, NAS2
NAS3, NAS4
(nas4x-1, nas4x-2)
nicotianamine (NA) synthesis Cytosol/
Roots (NAS1/2) and shoots (NAS1/2/3/4)
Elevated expression of FRO2, IRT1, and FIT in Fe-sufficient plants in the nas4x-1 Elevated Fe concentration in the whole rosette of the nas4x-1 mutant compared at the reproductive but not at the vegetative stage. Higher Fe in older leaves vs. younger leaves in the nas4x-1 mutant Foliar application of Fe or NA rescues interveinal leaf chlorosis of nas4x-2 Bauer et al., 2004; Klatte et al., 2009; Schuler et al., 2012
A. thaliana
/OPT3
(opt3-2 and
opt3-3)
Fe influx Plasma membrane/
Phloem companion cells
Constitutive expression of FRO2 and IRT1 High Fe accumulation in shoots
Higher Fe in older leaves vs. younger leaves
Grafting Wt shoots onto opt3 roots
Expression of OPT3 in leaves of the opt3-2
Stacey et al., 2008; Mendoza-Cozatl et al., 2014; Zhai et al., 2014
A. thaliana
/YSL1, YSL2, YSL3(ysl1,ysl2 ysl1ysl3)
Fe–NA transport Plasma membrane/
Vascular parenchyma cells (YSL1 and YSL3), xylem parenchyma cells in roots (YSL2)
Lower or wild-type FRO2 activity in ysl1ysl3 mutants, not tested in ysl1 or ysl2 mutants No difference in ysl2-1 mutant
Less Fe in seeds of ysl1 single mutant
Less Fe in roots and shoots of ysl1ysl3 double mutants
Foliar application of Fe increasing Fe in growth media for detached roots DiDonato et al., 2004; Jean et al., 2005; Waters et al., 2006; Chu et al., 2010
Solanum lycopersicon NAS (chln) NA synthesis Cytosol/
Roots and shoots
Constitutive expression of SlFRO1 and SlIRT1 High Fe accumulation in shoots
Higher Fe in older leaves vs. younger leaves
Foliar application of Fe or NA Scholz et al., 1985; Garcia et al., 2013
Unknown (brz) Unknown Unknown Constitutive expression of PsFRO1 and PsIRT1 High Fe accumulation in shoots
Higher Fe in older leaves vs. younger leaves
Foliar application of Fe Kneen et al., 1990; Garcia et al., 2013
Unknown (dgl) Unknown Unknown Constitutive expression of PsFRO1 and PsIRT1 High Fe accumulation in older leaves Grafting Wt shoots onto dgl roots Grusak and Pezeshgi, 1996; Garcia et al., 2013