Rice Science ›› 2019, Vol. 26 ›› Issue (6): 343-355.DOI: 10.1016/j.rsci.2018.12.007
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Received:
2018-10-12
Accepted:
2018-12-19
Online:
2019-11-28
Published:
2019-08-19
Saiful Islam Md. Sensing and Uptake of Nitrogen in Rice Plant: A Molecular View[J]. Rice Science, 2019, 26(6): 343-355.
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Fig. 1. Candidate NO3- transporters for improving nitrogen use efficiency (NUE) and yield of rice.The orange box indicates genes that could give positive effects on rice for uptake, transport and remobilization of NO3- in normal growth conditions. As examples, NRT1.1 (and another allele, NRT1.1B), NRT1.1b, NRT1.2, NRT2.1 and NRT2.2 (cooperating with NAR2.1) are responsible for NO3- uptake from soil. NRT2.3a (cooperating with NAR2.1) is a root-to-shoot NO3- transporter, and NRT2.3b is responsible for transporting of NO3- to the shoot and remobilizing N into the grain. The blue box indicates genes that can increase rice NUE and yield under salinity, such as NPF2.4 and NPF2.1, by regulating NO3- and K+ distributions in rice plants.
Gene | Gene locus | Function | Spatial expression | Transcriptional regulation by N | Effect on plant development | Reference |
---|---|---|---|---|---|---|
OsAMT1.1 | LOC_Os04g43070 | Dual-affinity NH4+ transporter, root-to-shoot transporter | Root epidermis and vascular tissues, root-shoot junction, vascular bundle, and leaf mesophyll cells | Induced by NH4+ and repressed by N starvation | Promote root growth and grain yield | |
OsAMT1.2 | LOC_Os02g40710 | NH4+ transporter | Vascular tissues, endodermis and pericycle in primary root tip | Induced by NH4+ and repressed by N starvation | Unknown | |
OsAMT1.3 | LOC_Os02g40730 | NH4+ transporter | Root vascular tissue and lateral root primordia | Repressed by NH4+ and induced by N starvation | Promote root elongation | |
OsAMT2.1 | LOC_Os05g39240 | Low-affinity NH4+ transporter | Root, shoot, leaf blade and sheath | Irresponsive to N supply | Unknown | |
OsAMT2.2 | LOC_Os01g61510 | NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT2.3 | LOC_Os01g61550 | NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT3.1 | LOC_Os01g65000 | Putative NH4+ transporter | Unknown | Induced by N starvation | Unknown | |
OsAMT3.2 | LOC_Os03g62200 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT3.3 | LOC_Os02g34580 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT4 | LOC_Os03g53780 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsNRT1.1b | LOC_Os10g40600 | Dual-affinity NO3- transporter | Root epidermis, hairs and vascular tissues, and leaf sheath, leaf blade and culm | Induced by NO3- | Grain yield increase | |
OsNPF2.4 | LOC_Os03g48180 | Low-affinity NO3- transporter, root-to-shoot transporter | Root epidermis, xylem parenchyma, and phloem companion cells and leaf phloem | Induced by high NO3- supply | Promote root growth and biomass, and grain yield | |
OsNRT2.1 | LOC_Os02g02170 | High-affinity NO3- transporter | Abundantly expressed in most cell types in root, but weakly expressed in shoot | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNRT2.2 | LOC_Os02g02190 | High-affinity NO3 transporter | Abundantly expressed in most cell types in roots, but weakly expressed in shoot | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNRT2.3a | LOC_Os01g50820 | High-affinity NO3- transporter, transport nitrate from root to shoot, not root nitrate uptake | Vascular tissues at root and root-shoot junction | Induced by NO3- and repressed by NH4+ | Promote root growth and biomass | |
OsNRT2.3b | LOC_Os01g50820 | High-affinity NO3- transporter, sensing cytosolic pH changes | Vascular tissue, mainly in phloem in both root and shoot | Irresponsive to N supply | Improved grain yield and nitrogen use efficiency | |
OsNRT2.4 | LOC_Os01g36720 | Putative high-affinity NO3- transporter | Lateral root primordia and shoot vascular tissues | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNAR2.1 | LOC_Os02g38230 | Partner proteins of OsNRT2.1, OsNRT2.2 and OsNRT2.3a | Epidermis and vascular tissue in root and shoot | Induced by NO3- and repressed by NH4+ | Promote root and shoot growth | |
OsNAR2.2 | LOC_Os04g40410 | Putative partner protein of OsNRT2s | Unknown | Induced by high NO3- and NH4+ | Unknown |
Table 1 Different genes involved in N transport in rice plant.
Gene | Gene locus | Function | Spatial expression | Transcriptional regulation by N | Effect on plant development | Reference |
---|---|---|---|---|---|---|
OsAMT1.1 | LOC_Os04g43070 | Dual-affinity NH4+ transporter, root-to-shoot transporter | Root epidermis and vascular tissues, root-shoot junction, vascular bundle, and leaf mesophyll cells | Induced by NH4+ and repressed by N starvation | Promote root growth and grain yield | |
OsAMT1.2 | LOC_Os02g40710 | NH4+ transporter | Vascular tissues, endodermis and pericycle in primary root tip | Induced by NH4+ and repressed by N starvation | Unknown | |
OsAMT1.3 | LOC_Os02g40730 | NH4+ transporter | Root vascular tissue and lateral root primordia | Repressed by NH4+ and induced by N starvation | Promote root elongation | |
OsAMT2.1 | LOC_Os05g39240 | Low-affinity NH4+ transporter | Root, shoot, leaf blade and sheath | Irresponsive to N supply | Unknown | |
OsAMT2.2 | LOC_Os01g61510 | NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT2.3 | LOC_Os01g61550 | NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT3.1 | LOC_Os01g65000 | Putative NH4+ transporter | Unknown | Induced by N starvation | Unknown | |
OsAMT3.2 | LOC_Os03g62200 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT3.3 | LOC_Os02g34580 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsAMT4 | LOC_Os03g53780 | Putative NH4+ transporter | Unknown | Unknown | Unknown | |
OsNRT1.1b | LOC_Os10g40600 | Dual-affinity NO3- transporter | Root epidermis, hairs and vascular tissues, and leaf sheath, leaf blade and culm | Induced by NO3- | Grain yield increase | |
OsNPF2.4 | LOC_Os03g48180 | Low-affinity NO3- transporter, root-to-shoot transporter | Root epidermis, xylem parenchyma, and phloem companion cells and leaf phloem | Induced by high NO3- supply | Promote root growth and biomass, and grain yield | |
OsNRT2.1 | LOC_Os02g02170 | High-affinity NO3- transporter | Abundantly expressed in most cell types in root, but weakly expressed in shoot | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNRT2.2 | LOC_Os02g02190 | High-affinity NO3 transporter | Abundantly expressed in most cell types in roots, but weakly expressed in shoot | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNRT2.3a | LOC_Os01g50820 | High-affinity NO3- transporter, transport nitrate from root to shoot, not root nitrate uptake | Vascular tissues at root and root-shoot junction | Induced by NO3- and repressed by NH4+ | Promote root growth and biomass | |
OsNRT2.3b | LOC_Os01g50820 | High-affinity NO3- transporter, sensing cytosolic pH changes | Vascular tissue, mainly in phloem in both root and shoot | Irresponsive to N supply | Improved grain yield and nitrogen use efficiency | |
OsNRT2.4 | LOC_Os01g36720 | Putative high-affinity NO3- transporter | Lateral root primordia and shoot vascular tissues | Induced by NO3- and repressed by NH4+ | Unknown | |
OsNAR2.1 | LOC_Os02g38230 | Partner proteins of OsNRT2.1, OsNRT2.2 and OsNRT2.3a | Epidermis and vascular tissue in root and shoot | Induced by NO3- and repressed by NH4+ | Promote root and shoot growth | |
OsNAR2.2 | LOC_Os04g40410 | Putative partner protein of OsNRT2s | Unknown | Induced by high NO3- and NH4+ | Unknown |
Fig. 2. Schematic representation of proposed evolution and predicted functions for nitrate transporters (NAR2/NRT2) in rice.NAR2.1, NAR2.2, NRT2.1, NRT2.2 and NRT2.3a are expressed mainly in roots; NRT2.3b and NRT2.4 are expressed mainly in shoots (Feng et al, 2011; Yan et al, 2011). Both NRT2.1 and NRT2.2 associated with the nitrate transport (NAR2.1) in the high-affinity concentration range. NRT2.3a requires NAR2.1 for the nitrate transport function, and the protein has a 10-fold lower affinity for nitrate than NRT2.1 and NRT2.2. NAR2.1 can provide a switch, depending on the partner transporter, to enable a rapid response in nitrate uptake over the dynamic ranges of external concentrations of nitrate (Feng et al, 2011; Yan et al, 2011). In contrast, NRT2.3b can function in nitrate transport independently, mainly in the shoot, and its overexpression can greatly improve nitrogen use efficiency and grain yield in rice (Feng et al, 2011; Yan et al, 2011). The solid red arrows represent defined direct functions of the transporters in nitrate uptake and translocation; the dashed arrows represent presumed relationships based on the tissue localization of the genes in rice plant and functional expression in oocytes. The blue arrows indicate the proposed evolution of individual members of the NAR2 and NRT2 nitrate transporter families. Black arrows indicate the possible relationships between NAR2.1 and root growth and between the functions of NRT2 members and plant growth and development.
Fig. 3. Regulation of root-to-shoot transport of nitrate.Nitrate transporter 1.5 (NRT1.5), expressed in the xylem pole pericycle, mediates xylem-loading of nitrate (red arrow) and facilitates root-to-shoot xylem transport of nitrate (blue arrow). NRT1.9, expressed in companion cells of phloem, mediates phloem loading (dark-purple arrow) for downward phloem transport of nitrate (dark blue arrow), which may facilitate xylem-to-phloem transfer of nitrate (broken purple arrow), and then reduce root-to-shoot nitrate transport. NRT1.8, expressed in xylem parenchyma, mediates nitrate removal from xylem (green arrow), and could also reduce root-to-shoot nitrate transport.Co, Cortex; En, Endodermis; Pe, Pericycle; Ph, Phloem; Xy, Xylem.
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