Function and Regulation of Ammonium Transporters in Plants
Abstract
:1. Introduction
2. The Physiological Roles of AMTs in Plants
2.1. AMT Mediates Ammonium Acquisition from Soil Solution
2.2. AMT Mediates Root-to-Shoot Transport of Ammonium
2.3. AMT Mediates Ammonium Transport in Leaves
2.4. AMT Mediates Ammonium Acquisition in the Reproductive Organs
2.5. AMT Mediates Ammonium Transport from Symbiotic Fungi to Host Plants
2.6. AMT Is Required for Root Development
2.7. The Role of AMT in Plant Disease Defense
3. Substrate Transport Mechanisms in AMTs
3.1. NH4+ Uniporter
3.2. NH3/H+ Symporter
3.3. NH3 Gas Channel
3.4. NH4+/H+ Symporter
4. Functional Regulations of AMTs
4.1. Regulation by Transcription Factors
4.2. Regulation by pH
4.3. Regulation by Phosphorylation
4.4. Regulation by Internalization and Heterotrimerization
5. Conclusion and Prospect
Funding
Conflicts of Interest
Appendix A
References
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Protein Name | Location | Evidenced by Plant KO/OE Lines | Physiological Roles | References |
---|---|---|---|---|
Ammonium uptake from soils | ||||
AtAMT1;1 | Root, rhizodermis, and root hairs | Yes | Symplastic transport of ammonium, accounts for 30%–35% of total ammonium uptake in roots | [19,21,22,23,24,25,26] |
AtAMT1;2 | Root, endothelial cells | Yes | Apoplastic transport of ammonium, accounts for 18%–26% of total ammonium influx in roots | [21,26,29] |
AtAMT1;3 | Root, rhizodermis, and root hairs | Yes | Symplastic transport of ammonium, accounts for 30%–35% of total ammonium uptake in roots | [21,25,26,30] |
AtAMT1;5 | Root, rhizodermis, and root hairs | No | Potential ammonium uptake in roots | [26] |
OsAMT1;1 | Root, epidermis | Yes | Contributes 25% of ammonium uptake in roots | [32,33,34] |
NtAMT1.3 | Root | Yes | Ammonium uptake in roots | [35] |
ZmAMT1.1a | Root, epidermal cells | Yes | Ammonium uptake in roots | [36] |
ZmAMT1.3 | Root, epidermal cells | Yes | Ammonium uptake in roots | [36] |
GhAMT1;3 | Root | Yes | Ammonium uptake in roots | [37] |
PutAMT1;1 | Root | Yes | Ammonium uptake in roots | [38] |
OsAMT1;2 | Root, exodermis cells | No | Potential ammonium uptake in roots | [1] |
LjAMT1;1-1;3 | Root | No | Potential ammonium uptake in roots | [39] |
PtrAMT1;2 | Root | No | Potential ammonium uptake in roots | [40] |
LeAMT1;1 | Root | No | Potential ammonium uptake in roots | [41,46,49] |
MtAMT1;1, MtAMT2;1 | Root, rhizodermal cells | No | Potential ammonium uptake in roots | [42] |
PpAMT1.3, PpAMT2;3 | Root | No | Potential ammonium uptake in roots | [43] |
OsAMT1;3 | Root | Yes | Potential ammonium uptake in roots | [44] |
PbAMT1;3 | Root | No | Potential ammonium uptake in roots | [45] |
LeAMT1;2 | Root | No | Potential ammonium uptake in roots | [47,48] |
TaAMT1.1 | Root | No | Potential ammonium uptake in roots | [50] |
PvAMT1;1 | Root | No | Potential ammonium uptake in roots | [51] |
Root-to-shoot translocation | ||||
AtAMT2 | Root, pericycle | Yes | Ammonium root-to-shoot translocation | [56] |
OsAMT1;1 | Root, stele | Yes | Ammonium root-to-shoot translocation | [32,33,34] |
ZmAMT1.3 | Root, pericycle cell layer | No | Ammonium root-to-shoot translocation | [36] |
OsAMT1;2 | Root, pericycle cells | No | Ammonium root-to-shoot translocation | [1] |
Ammonium transport in leaves | ||||
LeAMT1;2 | Leaf | No | Potential retrieval of photorespiration ammonium escaping from mitochondria and import of ammonium in the xylem | [61] |
LeAMT1;3 | Leaf | No | Potential compensation for ammonium losses across the plasma membrane caused by de- and transamination processes | [61] |
LjAMT1;1 | Leaf | No | Potential retrieval of photorespiration ammonium escaping from mitochondria and import of ammonium in the xylem | [39] |
BnAMT1;2 | Leaf | No | Potential retrieval of photorespiration ammonium escaping from mitochondria and import of ammonium in the xylem | [62] |
PtrAMT1;6, PtrAMT3;1 | Senescent leaf | No | Potential ammonium transport in leaf | [40] |
PtrAMT2;2, PtrAMT1;1 | Petioles | No | Potential ammonium transport in leaf | [40] |
OsAMT1;1 | Leaf mesophyll cells | No | Potential ammonium transport in leaf | [32,33,34] |
NtAMT1.3 | Leaf | Yes | Ammonium transport in leaf | [35] |
ZmAMT1.1a | Leaf | No | Potential ammonium transport in leaf | [36] |
ZmAMT1.3 | Leaf | No | Potential ammonium transport in leaf | [36] |
GhAMT1;3 | Leaf | No | Potential ammonium transport in leaf | [37] |
Ammonium acquisition in the reproductive organs | ||||
AtAMT1;4 | Flower | Yes | Ammonium acquisition in flower | [27] |
AtAMT1;1 | Flower | No | Potential ammonium acquisition in flower | [64] |
PtrAMT1;5 | Stamen | No | Potential ammonium acquisition in flower | [40] |
PtrAMT1;6 | Female flower | No | Potential ammonium acquisition in flower | [40] |
SbAMT1;1, SbAMT1;2, SbAMT2;1, SbAMT3;1, and SbAMT3;3 | Pistils and stamens | No | Potential ammonium acquisition in flower | [65] |
SbAMT2;2 and SbAMT3;2 | Pistils | No | Potential ammonium acquisition in flower | [65] |
LjAMT1;1-1;3 | Flower | No | Potential ammonium acquisition in flower | [39] |
ZmAMT1.1a | Seeds | No | Potential ammonium acquisition in seeds | [36] |
PutAMT1;1 | Anther | No | Potential ammonium acquisition in flower | [38] |
Ammonium transport from symbiotic fungi to host plants | ||||
SbAMT3;1 | Cortex cells containing developing arbuscules | Yes | Transferring ammonium to host plant | [65,69] |
LjAMT2;2 | Mycorrhizal roots, arbusculated cells | No | Potentially transferring ammonium to host plant | [70] |
GmAMT4;1 | Arbuscular cortex cells | No | Potentially transferring ammonium to host plant | [71] |
LeAMT4, LeAMT5 | Mycorrhizal roots | No | Potentially transferring ammonium to host plant | [72] |
PttAMT1;2 | Mycorrhizal roots | No | Potentially transferring ammonium to host plant | [73] |
PoptrAMT1.2b, PoptrAMT1.3, PoptrAMT1.4a, and PtrAMT1;2 | Mycorrhizal roots | No | Potentially transferring ammonium to host plant | [40,73] |
Required for root development | ||||
AtAMT1;3 | Root | Yes | Required for high-order lateral root branching upon ammonium exposure | [74] |
LjAMT1;3 | Root | Yes | Required for short root phenotype upon high concentration of ammonium exposure | [75,76] |
LjAMT2;3 | Mycorrhizal roots | Yes | Required for root premature arbuscule degeneration suppression | [77] |
Roles in plant disease defense | ||||
AtAMT1;1 | Root | Yes | Enhances resistance to necrotrophic fungus Plectosphaerella cucumerina and reduces sensitivity to hemibiotrophic bacterium Pseudomonas syringae | [81] |
TaAMT2;3a | Leaf | Yes | Retards the growth of P. striiformis | [82] |
TaAMT1;1a, TaAMT1;1b, and TaAMT1;3 | Leaf | No | Participates in plant–pathogen interaction by transport of ammonium | [83] |
Protein Name | Transport Mechanisms | Supporting Evidence | References |
---|---|---|---|
NH4+ uniport | |||
LeAMT1;1 | NH4+ uniport | (i) Electrogenic transport of ammonium. (ii) The more negative the membrane potential, the smaller the Km value, suggesting the cationic transport. (iii) Reversal potential moves towards positive direction only by ammonium introduction. (iv) pH-independent. (v) Each methylammonium ion transported carries a positive elementary charge. | [46,49] |
AtAMT1;1 | NH4+ uniport | (i) Electrogenic transport of ammonium.(ii) pH-independent. | [14] |
OsAMT1;1 | NH4+ uniport | (i) Electrogenic transport of ammonium. (ii) pH-independent. | [32] |
NH3/H+ cotransport | |||
TaAMT1;1 | NH3/H+ cotransport | (i) Electrogenic transport of ammonium. (ii) Each methylammonium ion transported carries a positive elementary charge. (iii) Stimulated by acidic pH. | [50] |
AtAMT1;2 | NH3/H+ cotransport | (i) Electrogenic transport of ammonium. (ii) Mutation of Q67H and W145S results in uncoupling transport of NH3 and H+. | [84] |
NH3 cotransport | |||
AtAMT2 | NH3 cotransport | (i) Electroneutral transport of ammonium. (ii) pH-independent. | [28] |
LjAMT2;2 | NH3 cotransport | (i) Electroneutral transport of ammonium. (ii) Stimulated by acidic pH. | [70] |
NH4+/H+ cotransport | |||
PvAMT1;1 | NH4+/H+ cotransport | (i) Electrogenic transport of ammonium. (ii) NH4+ and H+ ions together describe the ideal slope of reversal potential changes against 10-fold substrate concentration changes, and H199E mutation causes only NH4+ to describe the ideal slope. (iii) Cytosol acidification by PvAMT1;1 upon ammonium exposure, but this is not the case by H199E mutation. | [51] |
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Hao, D.-L.; Zhou, J.-Y.; Yang, S.-Y.; Qi, W.; Yang, K.-J.; Su, Y.-H. Function and Regulation of Ammonium Transporters in Plants. Int. J. Mol. Sci. 2020, 21, 3557. https://doi.org/10.3390/ijms21103557
Hao D-L, Zhou J-Y, Yang S-Y, Qi W, Yang K-J, Su Y-H. Function and Regulation of Ammonium Transporters in Plants. International Journal of Molecular Sciences. 2020; 21(10):3557. https://doi.org/10.3390/ijms21103557
Chicago/Turabian StyleHao, Dong-Li, Jin-Yan Zhou, Shun-Ying Yang, Wei Qi, Ke-Jun Yang, and Yan-Hua Su. 2020. "Function and Regulation of Ammonium Transporters in Plants" International Journal of Molecular Sciences 21, no. 10: 3557. https://doi.org/10.3390/ijms21103557