Abstract:Nitrogen is the main component of proteins, nucleic acids and enzymes, and plays an important role in plant growth and development. NH4+ uptake and assimilation consumes low energy compared to NO3- assimilation, NH4+ absorption and utilization efficiency directly affects plant growth and development. Plants can activate complex regulatory networks to optimize NH4+ uptake and utilization due to external fluctuations in NH4+ supply. This review summarized the ways in which ammonium transporter participates in NH4+ perception and the regulatory mechanism at the transcription and protein levels. In addition, it was summarized that the stress responses of AMTs and NH4+ assimilation-related genes under abiotic stresses such as low nitrogen, drought, and salt stress. This paper provides a theoretical basis for high nitrogen use efficiency of plant under abiotic stress.
[1] 丁庆倩,王小婷,胡利琴,等.2018.谷子MYB类转录因子SiMYB42提高转基因拟南芥低氮胁迫耐性[J].遗传,40(04):327-338.
(Ding Q Q,Wang X T,Hu L Q,et al.2018.MYB-like transcription factor SiMYB42 from foxtail millet (Setariaitalica L.) enhances Arabidopsis tolerance to low-nitrogen stress[J].Hereditas,40(04):327-338.
[2] 董越梅,李久蒂,朱至清,等.2000.铵载体(AMT)研究进展[J].植物学通报,17(1):39-45.
(Dong Y M,Li J D,Zhu Z Q,et al.2000.Molecular research progress of ammonium transporter[J].Chinese Bulletin of Botany,17(1):39-45.
[3] 李静,张冰玉,苏晓华,等.2012.植物中的铵根及硝酸根转运蛋白研究进展.南京林业大学学报(自然科学版),36(04):137-143.
(Li J,Zhang B Y,Su X H,et al.2012.Research progress of ammonium and nitrate transporters in plants[J].Nanjing Forestry University (Natural Science Edition),36(04):137-143.)
[4] 李磊,罗杰,李红,等.2011.毛果杨全基因组铵转运蛋白家族成员及其序列分析[J].西北农林科技大学学报(自然科学版),39(02):133-142.
(Li L,Luo J,Li H,et al.2011.Genome-wide analysis of the ammonium transporter gene family in Populus trichocarpa[J].Journal of Northwest A&F University(Nat.Sci.Ed.),39(02):133-142.)
[5] 李园枚,傅明辉,蒋丽花.2012.植物铵转运蛋白研究进展[J].广东农业科学,19:148-151.
(Li Y M,Fu M H,Jiang L H.2012,Research progress on plant AMT[J].Guangdong Agricultural Science,19:148-151.)
[6] 刘婷,尚忠林.2016.植物对铵态氮的吸收转运调控机制研究进展[J].植物生理学报,52(6):799-809.
(Liu T,Shang Z L.2016.Research progress on the regulation mechanism of ammonium nitrogen absorption and transport in plants[J].Plant Physiology,52(6):799-809.)
[7] 骆媛媛,柳参奎.2009.植物中铵转运蛋白的研究进展[J].基因组学与应用生物学,28(2):373-379.
(Luo Y Y,Liu C K.2009.Research progress of ammonium transporter in plants[J].Genomics and Applied Biology,28(2):373-379.)
[8] 王英,吕德国,秦嗣军,等.2010.低温对山定子和平邑甜茶幼苗根系氮代谢酶及游离氨基酸的影响[J].园艺学报,37(02):179-184.
(Wang Y,Lv D G,Qing S Jet al.2010.Effects of low-temperature on enzymes activities of nitrogen metabolism and free amino acids contents in root of malus baccata borkh.and malus hupehensis rehd.seedlings[J].Acta Horticulturae Sinica,37(02):179-184.)
[9] 杨寒.2016.毛果杨铵转运蛋白PtrAMT1;6功能鉴定和遗传转化[D].硕士学位论文,东北林业大学,导师:刘关君,pp.35-36.
(Yang H.2016.Function identification and genetic transformation of PrtAMT1;6,an ammonium tansporter of Populus trichocarpa[D].Thesis for M.S.,Northeast Forestry University,Supervisor:Liu G J,pp.35-36.)
[10] 张体付,葛敏,张晓林,等.2012.玉米Dof转录因子MZ1g40在氮利用途径中的功能分析[C].//2012年全国玉米遗传育种学术研讨会暨新品种展示观摩会论文及摘要集,pp.9.
(Zhang T F,Ge M,Zhang X L,et al.2012.Functional analysis of maize dof transcription factor MZ1g40 in nitrogen utilization pathway[C].//2012 National Maize Genetics and Breeding Academic Symposium and New Variety Exhibition and Observation Papers and Abstracts,pp.9.)
[11] Abouelsaad I,Weihrauch D,Renault S.2016.Effects of salt stress on the expression of key genes related to nitrogen assimilation and transport in the roots of the cultivated tomato (Solanum lycopersicon) and its wild salt-tolerant relative (Solanum pennellii)[J].Scientia Horticulturae,211:70-78.
[12] Britto D T,Kronzucker H J.2002.NH4+ toxicity in higher plants:A critical review[J].Journal of Plant Physiology,159(6):567-584.
[13] Britto D T,Siddiqi M Y,Glass A D M,et al.2001.Futile transmembrane NH4+ cycling:A cellular hypothesis to explain ammonium toxicity in plants[J].Proceedings of the National Academy of Sciences of the USA,98(7):4255-4258.
[14] Brix H,Dyhr-Jensen K,Lorenzen B.2002.Root-zone acidity and nitrogen source affects Typha latifolia L.growth and uptake kinetics of ammonium and nitrate[J].Journal of Experimental Botany,53(379):2441-2450.
[15] Bu Y Y,Takano T,Liu S K.2019.The role of ammonium transporter (AMT) against salt stress in plants[J].Plant Signaling & Behavior,14(8):1-3.
[16] Camanes G,Cerezo M,Primo-Millo E,et al.2009.Ammonium transport and CitAMT1 expression are regulated by N in Citrus plants[J].Planta,229(2):331-342.
[17] Castro-Rodriguez V,Assaf-Casals I,Perez-Tienda J,et al.2016.Deciphering the molecular basis of ammonium uptake and transport in maritime pine[J].Plant Cell and Environment,39(8):1669-1682.
[18] Chaves M M,Flexas J,Pinheiro C.2008.Photosynthesis under drought and salt stress:regulation mechanisms from whole plant to cell[J].Annals of Botany,103(4):551-560.
[19] Cheng Y R,Wang C,Chai S Y,et al.2017.Ammonium N influences the uptakes,translocations,subcellular distributions and chemical forms of Cd and Zn to mediate the Cd/Zn interactions in dwarf polish wheat (Triticum polonicum L.) seedlings[J].Chemosphere,193:1164-1171.
[20] Coskun D,Britto D T,Li M Y,et al.2013.Rapid ammonia gas transport accounts for futile transmembrane cycling under NH3/NH4+ toxicity in plant roots[J].Plant Physiology,163(4):1859-1867.
[21] Couturier J,Montanini B,Martin F,et al.2007.The expanded family of ammonium transporters in the perennial poplar plant[J].New Phytologist,174(1):137-150.
[22] Cui L J,Cao R,Li J L,et al.2006.High temperature effects on ammonium assimilation in leaves of two Festuca arundinacea cultivars with different heat susceptibility[J].Plant Growth Regulation,49(2-3):127-136.
[23] Ding L,Gao C M,Li Y R,et al.2015.The enhanced drought tolerance of rice plants under ammonium is related to aquaporin (AQP)[J].Plant Science,23(4):14-21.
[24] Dluzniewska P,Gessler A,Dietrich H,et al.2007.Nitrogen uptake and metabolism in Populus x canescens as affected by salinity[J].New Phytologist,173(2):279-293.
[25] Dominique L,Yuan L,Kojima S,et al.2006.Additive contribution of AMT1;1 and AMT1;3 to high-affinity ammonium uptake across the plasma membrane of nitrogen-deficient Arabidopsis roots[J].Plant Journal,48(4):522-534.
[26] Ehlting B,Dluzniewska P,Dietrich H,et al.2007.Interaction of nitrogen nutrition and salinity in Grey poplar (Populus tremula x alba)[J].Plant Cell and Environment,30(7):796-811.
[27] Espen L,Nocito F F,Cocucci M.2004.Effect of NO3- transport and reduction on intracellular pH:An in vivo NMR study in maize roots[J].Journal of Experimental Botany,55(405):2053-2061.
[28] Fan T F,chen X Y,Shi D X,et al.2017.Molecular identification of tobacco NtAMT1.3 that mediated ammonium root-influx with high affinity and improved plant growth on ammonium when overexpressed in Arabidopsis and tobacco[J].Plant Science,264:102-111.
[29] Fernandez-Crespo E,Camanes G,Garcia-Agustin P.2012.Ammonium enhances resistance to salinity stress in citrus plants[J].Journal of Plant Physiology,169(12):1183-1191.
[30] Fernandez-Crespo E,Gomez-Pastor R,Scalschi L,et al.2014.NH4+ induces antioxidant cellular machinery and provides resistance to salt stress in citrus plants[J].Trees-Structure and Function,28(6):1693-1704.
[31] Ferreira L M,de Souza V M,Tavares O C H,et al.2015.OsAMT1.3 expression alters rice ammonium uptake kinetics and root morphology[J].Plant Biotechnology Reports,9(4):221-229.
[32] Gazzarrini S,Lejay L,Gojon A,et al.1999.Three functional transporters for constitutive,diurnally regulated,and starvation-induced uptake of ammonium into arabidopsis roots[J].Plant Cell,11(5):937-947.
[33] Guo S,Zhou Y,Li Y,et al.2008.Effects of different nitrogen forms and osmotic stress on water use efficiency of rice (Oryza sativa)[J].Annals of Applied Biology,153(1):127-134.
[34] Hessini K,Ben Hamed K,Gandour M,et al.2013.Ammonium nutrition in the halophyte Spartina alterniflora under salt stress:Evidence for a priming effect of ammonium[J].Plant and Soil,370(1-2):163-173.
[35] Huang L L,Li M J,Shao Y,et al.2018.Ammonium uptake increases in response to PEG-induced drought stress in Malus hupehensis Rehd.[J].Environmental and Experimental Botany,151:32-42.
[36] Iwamoto M,Tagiri A.2016.MicroRNA-targeted transcription factor gene RDD1 promotes nutrient ion uptake and accumulation in rice[J].Plant Journal,85(4):466-477.
[37] Jia J B,Zhou J,Shi W G,et al.2017.Comparative transcriptomic analysis reveals the roles of overlapping heat-/drought-responsive genes in Poplars exposed to high temperature and drought[J].Scientific Reports,7:43215.
[38] Kant S,Kant P,Lips H,et al.2007.Partial substitution of NO3- by NH4+ fertilization increases ammonium assimilating enzyme activities and reduces the deleterious effects of salinity on the growth of barley[J].Journal of Plant Physiology,164(3):303-311.
[39] Katz A,Waridel P,Shevchenko A,et al.2007.Salt-induced changes in the plasma membrane proteome of the halotolerant alga dunaliella salina as revealed by blue native gel electrophoresis and nano-LC-MS/MS analysis[J].Molecular & Cellular Proteomics,6(9):1459-1472.
[40] Kuznetsov V V,Rakitin V Y,Zholkevich V N.1999.Effects of preliminary heat-shock treatment on accumulation of osmolytes and drought resistance in cotton plants during water deficiency[J].Physiologia Plantarum,107(4):399-406.
[41] Li C,Tang Z,Wei J,et al.2016.The OsAMT1.1 gene functions in ammonium uptake and ammonium-potassium homeostasis over low and high ammonium concentration ranges[J].Journal of Genetics and Genomics,43(11):639-649.
[42] Li G J,Zhang L,Wang M,et al.2019.The Arabidopsis AMOT1/EIN3 gene plays an important role in the amelioration of ammonium toxicity[J].Journal of Experimental Botany,70(4):1375-1388.
[43] Lima J E,Kojima S,Takahashi H,et al.2010.Ammonium triggers lateral root branching in Arabidopsis in an AMMONIUM TRANSPORTER1;3-dependent manner[J].Plant Cell,22(11):3621-3633.
[44] Liu F,Xu Y R,Chang K X,et al.2019.The long noncoding RNA T5120 regulates nitrate response and assimilation in Arabidopsis[J].New Phytologist,224(1):117-131.
[45] Liu Y,Wiren N.2017.Ammonium as a signal for physiological and morphological responses in plants[J].Journal of Experimental Botany,68(10):2581-2592.
[46] Loque D,Lalonde S,Looger L L,et al.2007.A cytosolic trans-activation domain essential for ammonium uptake[J].Nature,446(7132):195-198.
[47] Meng S,Su L,Li Y M,et al.2016a.Nitrate and ammonium contribute to the distinct nitrogen metabolism of Populus simonii during moderate salt stress[J].PLOS ONE,11(3):e0150354
[48] Meng S,Zhang C X,Su L,et al.2016b.Distinct effect of pH on N uptake and assimilation in two conifer species[J].Trees-Structure and Function,30(5):1607-1618.
[49] Meng S.,Zhang C.,Su L.,Li Y.,andZhao Z.2016c.Nitrogen uptake and metabolism of Populus simonii in response to PEG-induced drought stress[J].Environmental and Experimental Botany,123:78-87.
[50] Miranda R D,Gomes E,Prisco J T,et al.2016.Ammonium improves tolerance to salinity stress in Sorghum bicolor plants[J].Plant Growth Regulation,78(1):121-131.
[51] Neuhauser B,Dynowski M,Mayer M,et al.2007.Regulation of NH4+ transport by essential cross talk between AMT monomers through the carboxyl tails[J].Plant Physiology,143(4):1651-1659.
[52] Patterson K,Cakmak T,Cooper A,et al.2010.Distinct signalling pathways and transcriptome response signatures differentiate ammonium and nitrate-supplied plants[J].Plant Cell and Environment,33(9):1486-1501.
[53] Qasim B,Motelica-Heino M,Bourgerie S,et al.2015.Effect of nitrate and ammonium fertilization on Zn,Pb,and Cd phytostabilization by Populus euramericana Dorskamp in contaminated technosol[J].Environmental Science and Pollution Research,22(23):18759-18771.
[54] Ranathunge K,El-kereamy A,Gidda S,et al.2014.AMT1;1 transgenic rice plants with enhanced NH4+ permeability show superior growth and higher yield under optimal and suboptimal NH4+ conditions[J].Journal of Experimental Botany,65(4):965-979.
[55] Reid R,Hayes J.2003.Mechanisms and control of nutrient uptake in plants[J].International Review of Cytology,29:73-114.
[56] Robertson G P,Vitousek P M.2009.Nitrogen in agriculture:Balancing the cost of an essential resource[J].Annual Review of Environment and Resources,34(1):97-125.
[57] Rymarquis L A,Kastenmayer J P,Huttenhofer A G,et al.2008.Diamonds in the rough:mRNA-like non-coding RNAs[J].Trends in Plant Science,13(7):329-334.
[58] Shinozaki K,Yamaguchi-Shinozaki K.2000.Molecular responses to dehydration and low temperature:Differences and cross-talk between two stress signaling pathways[J].Current Opinion in Plant Biology,3(3):217-223
[59] Straub T,Ludewig U,Neuhauser B.2017.The kinase CIPK23 inhibits ammonium transport in Arabidopsis thaliana[J].Plant Cell,29(2):409-422.
[60] Suarez M F,Avila C,Gallardo F,et al.2002.Molecular and enzymatic analysis of ammonium assimilation in woody plants[J].Journal of Experimental Botany,53(370):891-904.
[61] Suenaga A.2003.Constitutive expression of a novel-type ammonium transporter OsAMT2 in rice plants[J].Plant and Cell Physiology,44(2):206-211.
[62] Sun Y C,Sheng S,Fan T F,et al.2019.Molecular identification and functional characterization of GhAMT1.3 in ammonium transport with a high affinity from cotton (Gossypium hirsutum L.)[J].Physiologia Plantarum,167(2):217-231.
[63] Tang Z H,Liu Y J,Guo X R,et al.2011.The combined effects of salinity and nitrogen forms on Catharanthus roseus:The role of internal ammonium and free amino acids during salt stress[J].Journal of Plant Nutrition and Soil Science,174(1):135-144.
[64] Vessey J K,Henry L T,Chaillou S,et al.1990.Root-zone acidity affects relative uptake of nitrate and ammonium from mixed nitrogen-sources[J].Journal of Plant Nutrition,13(1):95-116.
[65] Xiong L M,Wang R G,Mao G H,et al.2006.Identification of drought tolerance determinants by genetic analysis of root response to drought stress and abscisic acid[J].Plant Physiology,142(3):1065-1074.
[66] Xu C P,Jiang Z C,Huang B R.2011.Nitrogen deficiency-induced protein changes in immature and mature leaves of creeping bentgrass[J].Journal of the American Society for Horticultural Science,136(6):399-407.
[67] Xuan W,Beeckman T,Xu G H.2017.Plant nitrogen nutrition:sensing and signaling[J].Current Opinion in Plant Biology,39:57-65.
[68] Yang S Y,Hao D L,Cong Y,et al.2015.The rice OsAMT1;1 is a proton-independent feedback regulated ammonium transporter[J].Plant Cell Reports,34(2):321-330.
[69] Yu C,Lv D G,Qin S J,et al.2010.Changes in photosynthesis,fluorescence,and nitrogen metabolism of hawthorn (Crataegus pinnatifida) in response to exogenous glutamic acid[J].Photosynthetica,48(3):339-347.
[70] Yuan L X,Graff L,Loque D,et al.2009.AtAMT1;4,a pollen-specific high-affinity ammonium transporter of the plasma membrane in Arabidopsis[J].Plant and Cell Physiology,50(1):13-25.
[71] Yuan L X,Loque D,Kojima S,et al.2007.The organization of high-affinity ammonium uptake in Arabidopsis roots depends on the spatial arrangement and biochemical properties of AMT1-type transporters[J].Plant Cell,19(8):2636-2652.
[72] Yutaka S,Akira I,Satomi S,et al.2003.Distinct expression and function of three ammonium transporter genes (OsAMT1;1 - 1;3) in rice[J].Plant and Cell Physiology,44(7):726-734.
[73] Zhang C X,Meng S,Li M J,et al.2018.Transcriptomic insight into nitrogen uptake and metabolism of Populus simonii in response to drought and low nitrogen stresses[J].Tree Physiology,38(11):1672-1684.
[74] Zhang C X,Meng S,Li Y M,et al.2014.Net NH4+ and NO3- fluxes,and expression of NH4+ and NO3- transporter genes in roots of Populus simonii after acclimation to moderate salinity[J].Trees-Structure and Function,28(6):1813-1821.
[75] Zhang J S,Zhou Z Y,Bai J J,et al.2020.Disruption of MIR396e and MIR396f improves rice yield under nitrogen-deficient conditions[J].National Science Review,7(1):102-112.
[76] Zhang S M,Yang C,Chen M M,et al.2019.Influence of nitrogen availability on Cd accumulation and acclimation strategy of Populus leaves under Cd exposure[J].Ecotoxicology and Environmental Safety,180:439-448.
[77] Zhao B T,Zhu X F,Jung J H,et al.2016.Effect of brassinosteroids on ammonium uptake via regulation of ammonium transporter and N-metabolism genes in Arabidopsis[J].Biologia Plantarum,60(3):563-571.