TABLE 1.
Manganese transport proteins reviewed in this article.
| Family/name | Organ(s)/tissue(s) expression | Subcellular localization | Gene expression response | Other substrates | References |
| CaCA | |||||
| AtCAX2 | Root (stele and root tip), stem, leaf, flower, pollen, fruit | Tonoplast | Unaffected by +Cu, +Mn, +Zn | Ca, Cd | Hirschi et al., 2000; Shigaki et al., 2003; Pittman et al., 2004; Edmond et al., 2009 |
| AtCAX4 | Root, stem, leaf, flower, fruit | Tonoplast | Up-regulated by +Mn, +Ni, -Ca | Cd, Ca, Zn | Cheng et al., 2002; Koren’kov et al., 2007; Mei et al., 2009 |
| AtCAX5 | Root, stem, leaf, flower, fruit | Tonoplast | Up-regulated by +Mn down-regulated by +Zn | Ca | Edmond et al., 2009 |
| HvCAX2 | Root, leaf, seed | – | Up-regulated by +Ca, +Na unaffected by +Mn | Ca | Edmond et al., 2009 |
| LeCAX2 | Root, leaf, fruit | – | – | Ca | Edmond et al., 2009 |
| OsCAX1a | Root, shoot, flower, seed | Tonoplast | – | Ca | Kamiya et al., 2005 |
| OsCAX3 | Root, shoot, flower, seed | – | – | Ca | Kamiya et al., 2005 |
| OsCAX4 | Root | – | Up-regulated by +Ca | Ca,Cu | Yamada et al., 2014 |
| VvCAX3 | Root, stem, leaf, fruit | Tonoplast | Up-regulated by +Ca, +Na unaffected by +Mn | Ca, Cu, Li, Na | Martins et al., 2017 |
| AtCCX3 | Root, stem, leaf, flower | Tonoplast, endomembranes | Up-regulated by +K, +Na unaffected by +Mn | K, Na | Morris et al., 2008 |
| BICAT | |||||
| AtBICAT1/PAM71/CCHA1 | Leaf | Chloroplast (thylakoid membrane) | Unaffected by +Mn | Ca | Schneider et al., 2016; Wang et al., 2016; Eisenhut et al., 2018; Frank et al., 2019 |
| AtBICAT2/CMT1 | Root, stem, leaf, flower, fruit | Chloroplast (inner envelope) | Down-regulated by +Mn | Ca, Mg | Eisenhut et al., 2018; Frank et al., 2019; Zhang et al., 2018 |
| CDF/MTP | |||||
| AtMTP8 | Root (epidermis, cortex), seed | Tonoplast | Up-regulated by +Mn -Fe | Fe | Eroglu et al., 2016, 2017 |
| AtMTP9 | – | – | Unaffected by +Mn | – | Delhaize et al., 2007; Chu et al., 2017 |
| AtMTP10 | – | – | Unaffected by +Mn | – | Delhaize et al., 2007; Chu et al., 2017 |
| AtMTP11 | Root (root tip), leaf (guard cells) | Golgi/PVC | Unaffected by +Mn | – | Delhaize et al., 2007; Peiter et al., 2007 |
| ShMTP8 | – | Tonoplast | – | – | Delhaize et al., 2003, 2007 |
| BmMTP10 | Root, leaf | Endomembranes* | Up-regulated by +Mn | – | Erbasol et al., 2013 |
| BmMTP11 | Root, leaf | Endomembranes* | Unaffected by +Mn | Ni | Erbasol et al., 2013 |
| OsMTP8.1 | Root, shoot | Tonoplast | – | – | Chen et al., 2013; Takemoto et al., 2017 |
| OsMTP8.2 | Root, shoot | Tonoplast | – | – | Takemoto et al., 2017 |
| OsMTP9 | Root (endodermis, exodermis) | Plasma membrane | Unaffected by +Mn -Mn | Ueno et al., 2015 | |
| OsMTP11 | Root, shoot | Golgi/TGN | Up-regulated by +Mn +Zn +Cd +Ni | Co, Ni | Zhang and Liu, 2017; Ma et al., 2018; Tsunemitsu et al., 2018 |
| HvMTP8.1 | Root, leaf | Golgi | Down-regulated by -Mn (root) down-regulated by +Mn (shoot) | – | Pedas et al., 2014 |
| HvMTP8.2 | Root, leaf | Golgi | Down-regulated by +Mn | – | Pedas et al., 2014 |
| PtMTP11.1 | – | TGN | – | – | Peiter et al., 2007 |
| PtMTP11.2 | – | TGN | – | – | Peiter et al., 2007 |
| PbMTP8.1 | – | – | – | Fe | Hou et al., 2019 |
| PbMTP8.2 | – | – | – | Fe | Hou et al., 2019 |
| PbMTP9 | – | – | – | Fe | Hou et al., 2019 |
| PbMTP10 | – | – | – | Fe | Hou et al., 2019 |
| PbMTP11.1 | – | – | – | – | Hou et al., 2019 |
| PbMTP11.2 | – | – | – | – | Hou et al., 2019 |
| NtMTP8.1 | Root, stem, leaf, flower | – | Up-regulated by +Co +Zn unaffected by +Mn | – | Liu et al., 2019 |
| NtMTP8.4 | Stem, leaf, flower | – | Up-regulated by +Cd +Co +Mn (root) up-regulated by +Cd +Co +Mg +Zn (shoot) |
– | Liu et al., 2019 |
| NtMTP11.1 | Root, stem, leaf, flower | – | Up-regulated by +Co +Cd +Mn +Zn | – | Liu et al., 2019 |
| CsMTP9 | Root (endodermis) | Plasma membrane | Up-regulated by +Cd +Mn +Ni down-regulated by –Mn -Zn |
Cd | Migocka et al., 2015 |
| NRAMP | |||||
| AtNRAMP1 | Root (Cortex, endodermis) > > shoot | Plasma membrane | Up-regulated by -Fe -Mn | Cd, Fe | Curie et al., 2000; Thomine et al., 2000; Cailliatte et al., 2010; Castaings et al., 2016 |
| AtNRAMP2 | Root (pericycle, root tip), leaf vasculature, flower, trichome | TGN | Up-regulated by -Mn down-regulated by -Fe | – | Curie et al., 2000; Alejandro et al., 2017; Gao et al., 2018 |
| AtNRAMP3 | Root (stele), leaf vasculature, developing seed | Tonoplast | Up-regulated by -Fe unaffected by -Mn | Cd, Fe | Thomine et al., 2000, 2003; Lanquar et al., 2005, 2010 |
| AtNRAMP4 | Root (stele), leaf vasculature, developing seed | Tonoplast | Up-regulated by -Fe unaffected by -Mn | Cd, Fe | Thomine et al., 2000; Lanquar et al., 2005, 2010 |
| OsNRAMP3 | Node, leaf vasculature | Plasma membrane | Unaffected by +Mn -Mn | – | Yamaji et al., 2013; Yang et al., 2013 |
| OsNRAMP5 | Root (exodermis, endodermis, stele), panicle | Plasma membrane | Up-regulated by -Fe (shoot) up-regulated by -Fe, -Zn (root) unaffected by -Mn | Cd, Fe | Ishimaru et al., 2012; Sasaki et al., 2012; Yang et al., 2014 |
| OsNRAMP6 | Leaves | Plasma membrane | – | Fe | Peris-Peris et al., 2017 |
| HvNRAMP5 | Root (epidermis, stele) | Plasma membrane | Up-regulated by -Fe unaffected by +Mn | Cd | Wu et al., 2016 |
| BnNRAMP1b | Root, shoot | – | Up-regulated by +Cd | Cd, Zn | Meng et al., 2017 |
| TcNRAMP3 | – | Tonoplast | – | Cd, Fe | Oomen et al., 2009 |
| TcNRAMP4 | – | Tonoplast | – | Cd, Fe, Zn | Oomen et al., 2009 |
| TcNRAMP3 (T. cacao) |
Root, shoot | – | Up-regulated by -Fe unaffected by -Mn | Fe | Ullah et al., 2018 |
| TcNRAMP5 (T. cacao) |
Root | – | Up-regulated by -Fe unaffected by -Mn |
Cd, Fe, Zn | Ullah et al., 2018 |
| TcNRAMP6 (T. cacao) |
Root, shoot | – | – | – | Ullah et al., 2018 |
| LeNRAMP1 | Root | Endomembranes* | up-regulated by -Fe | – | Bereczky et al., 2003 |
| LeNRAMP3 | Root, shoot | Endomembranes* | Up-regulated by -Fe | – | Bereczky et al., 2003 |
| AhNRAMP1 | Root, stem | – | Up-regulated by -Mn, -Zn | Zn | Wang et al., 2019 |
| MbNRAMP1 | Root | Endomembranes* | Up-regulated by -Fe | Fe | Xiao et al., 2008 |
| P-TYPE ATPASE | |||||
| AtECA1 | Root, stem, leaf, guard cells, trichome | ER | – | Ca, Zn | Liang et al., 1997; Wu et al., 2002; Li et al., 2008 |
| AtECA3 | Root (stele), stem, leaf vasculature, guard cells, flower, fruit | Golgi | Unaffected by -Mn | Ca, Zn | Mills et al., 2007; Li et al., 2008 |
| LeLCA1 | – | ER? | – | Ca | Johnson et al., 2009 |
| VIT | |||||
| AtVIT1 | Root, cotyledon, developing seed | Tonoplast | – | Fe | Kim et al., 2006 |
| OsVIT1 | Leaf > > root, stem, panicle, embryo | Tonoplast | Down-regulated by -Fe | Fe, Zn | Zhang et al., 2012; Wang et al., 2017 |
| OsVIT2 | Leaf, stem, panicle, embryo | Tonoplast | Up-regulated by +Fe down-regulated by -Fe | Fe, Zn | Zhang et al., 2012 |
| TaVIT2 | Root, shoot, seed | Tonoplast | – | Fe | Connorton et al., 2017 |
| AtMEB1 | – | ER bodies | – | Fe | Yamada et al., 2013 |
| AtMEB2 | – | ER bodies | – | Fe | Yamada et al., 2013 |
| YSL | |||||
| OsYSL2 | Root (phloem), leaf, vascular bundle, developing seed | Plasma membrane | Up-regulated by -Fe down-regulated by -Mn | Fe | Koike et al., 2004; Yang et al., 2014 |
| OsYSL6 | Root, shoot | Plasma membrane | Unaffected by +Mn -Mn | – | Sasaki et al., 2011 |
| HvYSL2 | Root (endodermis), shoot | – | Up-regulated by -Fe | Fe, Zn, Co, Ni, Cu | Araki et al., 2011 |
| ZIP | |||||
| AtIRT1 | Root (epidermis, cortex) > > shoot | Plasma membrane | Up-regulated by -Fe | Fe, Zn, Co, Ni | Korshunova et al., 1999; Curie et al., 2000; Thomine et al., 2000; Vert et al., 2002 |
| AtZIP1 | Root (stele), leaf vasculature | Tonoplast | Up-regulated by -Fe -Zn (root) up-regulated by -Mn (shoot) down-regulated by -Zn (shoot) | Zn | Milner et al., 2013 |
| AtZIP2 | Root (stele) | Plasma membrane | Down-regulated by -Fe -Mn | Zn | Milner et al., 2013 |
| AtZIP5 | Root, shoot | – | – | – | Milner et al., 2013 |
| AtZIP6 | Root, shoot | – | – | – | Milner et al., 2013 |
| AtZIP7 | Shoot > > root | – | – | Fe, Zn | Milner et al., 2013; Fu et al., 2017 |
| AtZIP9 | Root, shoot | – | – | – | Milner et al., 2013 |
| HvIRT1 | Root (epidermis, cortex**, endodermis**, pericycle), seed | Plasma membrane | Up-regulated by -Fe, -Mn | Fe, Zn | Pedas et al., 2008; Long et al., 2018 |
| LeIRT1 | Root, flower | – | – | Cd, Fe, Zn | Eckhardt et al., 2001 |
| LeIRT2 | Root | – | – | Cd, Fe, Zn | Eckhardt et al., 2001 |
| MtZIP4 | Root***, leaf | – | Down-regulated by -Fe -Mn up-regulated by +Zn | – | López-Millán et al., 2004 |
| MtZIP7 | Leaf | – | Unaffected by -Mn | – | López-Millán et al., 2004 |
| PtIRT1 | Root, leaf | – | Down-regulated by -Mn | Fe, Zn | Fu et al., 2017 |
+ excess, − deficiency, * in yeast cells, ** only under +Fe, *** only under +Zn.