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氮作为植物必需的大量营养元素,是叶绿素、氨基酸和核酸等关键生命物质的重要组成成分。硝酸盐作为农业土壤及众多自然生态系统中植物可利用的主要氮源,其高效利用对保障粮食安全至关重要。然而,在全球人口持续增长和气候变化加剧的背景下,当前作物生产仍高度依赖大量氮肥投入。氮肥利用率低下不仅造成生产成本增加和资源浪费,更引发了一系列环境问题。因此,深入揭示植物硝酸盐转运的分子调控机制,并探究其在豆科植物根瘤共生固氮中的作用,对培育氮高效作物品种,及优化氮素营养管理策略具有重要的科学价值与实践意义。本文系统综述了植物硝酸盐转运系统的组成与功能,及其与钙信号、激素信号和磷营养等途径的互作调控网络;同时,总结了硝酸盐调控豆科植物结瘤与固氮的关键过程与分子基础,以期为深入解析作物氮高效利用机制、推动农业可持续发展提供理论依据。
Abstract:Nitrogen is an essential element for plants,and it acts as a key component of critical biomolecules such as chlorophyll,amino acids,and nucleic acids.As the primary source of available nitrogen in agricultural soils and many natural ecosystems,the efficient utilization of nitrate is crucial for ensuring global food security.However,driven by continuous global population growth and intensifying climate change,crop production still heavily relies on substantial nitrogen fertilizer inputs.The low efficiency of nitrogen fertilizer use not only increases production costs and leads to resource wastage but also causes a series of environmental problems.Therefore,elucidating the molecular regulatory mechanisms of nitrate transport in plants and exploring its role in symbiotic nitrogen fixation in legume hold significant scientific value and practical importance for developing nitrogen-efficient crop varieties and optimizing nitrogen nutrient management strategies.This review systematically summarizes the composition and function of plant nitrate transporters,as well as their interactive regulatory networks with calcium signaling,hormonal signaling,and phosphorus nutrition pathways.Furthermore,it outlines the key processes and molecular basis of nitrate-mediated regulation of nodulation and nitrogen fixation in legumes,aiming to provide a theoretical foundation for further deciphering the mechanisms of nitrogen use efficiency in crops and promoting sustainable agricultural development.
ALBORESI A,GESTIN C,LEYDECKER M T,et al.,2005.Nitrate,a signal relieving seed dormancy in Arabidopsis.Plant Cell Environ.,28(4):500-512.
ALMAGRO A,LIN S H,TSAY Y F,2008.Characterization of the Arabidopsis nitrate transporter NRT1.6 reveals a role of nitrate in early embryo development.Plant Cell,20(12):3289-3299.
ALVAREZ J M,RIVERAS E,VIDAL E A,et al.,2014.Systems approach identifies TGA1 and TGA4 transcription factors as important regulatory components of the nitrate response of Arabidopsis thaliana roots.Plant J.,80(1):1-13.
ALVAREZ J M,VIDAL E A,GUTIÉRREZ R A,2012.Integration of local and systemic signaling pathways for plant N responses.Curr.Opin.Plant Biol.,15(2):185-191.
ASIM M,ULLAH Z,OLUWASEUN A,et al.,2020.Signalling overlaps between nitrate and auxin in regulation of the root system architecture:insights from the Arabidopsis thaliana.Int.J.Mol.Sci.,21(8):2880.
BARBULOVA A,ROGATO A,D’APUZZO E,et al.,2007.Differential effects of combined N sources on early steps of the nod factor-dependent transduction pathway in Lotus japonicus.Mol.Plant Microbe Interact.,20(8):994-1003.
BELLEGARDE F,TJAHJONO O,YOSHINO-KIDA M,et al.,2025.Fluctuations in nitrate availability impact cytokinin biosynthesis through histone modifications of IPT3 in Arabidopsis roots for growth acclimation.Plant Commun.,6(11):101531.
BREAKSPEAR A,LIU C W,ROY S,et al.,2014.The root hair “infectome” of Medicago truncatula uncovers changes in cell cycle genes and reveals a requirement for auxin signaling in rhizobial infection.Plant Cell,26(12):4680-4701.
BUCHNER P,HAWKESFORD M J,2014.Complex phylogeny and gene expression patterns of members of the nitrate transporter 1/peptide transporter family (NPF) in wheat.J.Exp.Bot.,65(19):5697-5710.
CABA J M,CENTENO M L,FERNÁNDEZ B,et al.,2000.Inoculation and nitrate alter phytohormone levels in soybean roots:differences between a supernodulating mutant and the wild type.Planta,211(1):98-104.
CAMARGO A,LLAMAS A,SCHNELL R A,et al.,2007.Nitrate signaling by the regulatory gene NIT2 in chlamydomonas.Plant Cell,19(11):3491-3503.
CASTAINGS L,MARCHIVE C,MEYER C,et al.,2011.Nitrogen signalling in Arabidopsis:how to obtain insights into a complex signalling network.J.Exp.Bot.,62(4):1391-1397.
CEREZO M,TILLARD P,FILLEUR S,et al.,2001.Major alterations of the regulation of root NO3-uptake are associated with the mutation of NRT2.1 and NRT2.2 genes in Arabidopsis.Plant Physiol.,127(1):262-271.
CHIU C C,LIN C S,HSIA A P,et al.,2004.Mutation of a nitrate transporter,AtNRT1:4,results in a reduced petiole nitrate content and altered leaf development.Plant Cell Phy-siol.,45(9):1139-1148.
CHOPIN F,ORSEL M,DORBE M F,et al.,2007.The Arabidopsis AtNRT2.7 nitrate transporter controls nitrate content in seeds.Plant Cell,19(5):1590-1602.
CHRISTENSEN S,ROUSK K,2024.Global N2O emissions from our planet:which fluxes are affected by man,and can we reduce these iScience,27(2):109042.
CHU X Q,LI M Z,ZHANG S J,et al.,2021.HBI1-TCP20 interaction positively regulates the CEPs-mediated systemic nitrate acquisition.J.Integr.Plant Biol.,63(5):902-912.
CRAWFORD N M,FORDE B G,2002.Molecular and developmental biology of inorganic nitrogen nutrition.Arabidopsis Book,1:e0011.
DANISMAN S,VAN DER WAL F,DHONDT S,et al.,2012.Arabidopsis class Ⅰ and class Ⅱ TCP transcription factors regulate jasmonic acid metabolism and leaf development antagonistically.Plant Physiol.,159(4):1511-1523.
DECHORGNAT J,NGUYEN C T,ARMENGAUD P,et al.,2011.From the soil to the seeds:the long journey of nitrate in plants.J.Exp.Bot.,62(4):1349-1359.
DENYER T,MA X L,KLESEN S,et al.,2019.Spatiotemporal developmental trajectories in the Arabidopsis root revealed using high-throughput single-cell RNA sequencing.Dev.Cell,48(6):840-852.e5.
DODD A N,KUDLA J,SANDERS D,2010.The language of calcium signaling.Annu.Rev.Plant Biol.,61:593-620.
DONG Q Y,BAI B W,ALMUTAIRI B O,et al.,2021.Emerging roles of the CBL-CIPK calcium signaling network as key regulatory hub in plant nutrition.J.Plant Physiol.,257:153335.
FAN S C,LIN C S,HSU P K,et al.,2009.The Arabidopsis nitrate transporter NRT1.7,expressed in phloem,is responsible for source-to-sink remobilization of nitrate.Plant Cell,21(9):2750-2761.
FAN X R,TANG Z,TAN Y W,et al.,2016.Overexpression of a pH-sensitive nitrate transporter in rice increases crop yields.Proc.Natl.Acad.Sci.USA,113(26):7118-7123.
FENG H M,FAN X R,FAN X R,et al.,2011.Multiple roles of nitrate transport accessory protein NAR2 in plants.Plant Signal.Behav.,6(9):1286-1289.
FORDE B G,2000.Nitrate transporters in plants:structure,function and regulation.Biochim.Biophys.Acta (BBA) Biomembr.,1465(1-2):219-235.
FUJIKAKE H,YAMAZAKI A,OHTAKE N,et al.,2003.Quick and reversible inhibition of soybean root nodule growth by nitrate involves a decrease in sucrose supply to nodules.J.Exp.Bot.,54(386):1379-1388.
GAO Y Y,QI S D,WANG Y,2022.Nitrate signaling and use efficiency in crops.Plant Commun.,3(5):100353.
GRUBER N,GALLOWAY J N,2008.An earth-system perspective of the global nitrogen cycle.Nature,451(7176):293-296.
GUAN P Z,WANG R C,NACRY P,et al.,2014.Nitrate foraging by Arabidopsis roots is mediated by the transcription factor TCP20 through the systemic signaling pathway.Proc.Natl.Acad.Sci.USA,111(42):15267-15272.
HERVÉ C,DABOS P,BARDET C,et al.,2009.In vivo interference with AtTCP20 function induces severe plant growth alterations and deregulates the expression of many genes important for development.Plant Physiol.,149(3):1462-1477.
HIREL B,LE GOUIS J,NEY B,et al.,2007.The challenge of improving nitrogen use efficiency in crop plants:towards a more central role for genetic variability and quantitative genetics within integrated approaches.J.Exp.Bot.,58(9):2369-2387.
HIREL B,TÉTU T,LEA P J,et al.,2011.Improving nitrogen use efficiency in crops for sustainable agriculture.Sustainability,3(9):1452-1485.
HSU P K,TSAY Y F,2013.Two phloem nitrate transporters,NRT1.11 and NRT1.12,are important for redistributing xylem-borne nitrate to enhance plant growth.Plant Physiol.,163(2):844-856.
HU B,JIANG Z M,WANG W,et al.,2019.Nitrate-NRT1.1B-SPX4 cascade integrates nitrogen and phosphorus signalling networks in plants.Nat.Plants,5(4):401-413.
HU B,WANG W,OU S J,et al.,2015.Variation in NRT1.1B contributes to nitrate-use divergence between rice subspecies.Nat.Genet.,47(7):834-838.
HU H C,WANG Y Y,TSAY Y F,2009.AtCIPK8,a CBL-interacting protein kinase,regulates the low-affinity phase of the primary nitrate response.Plant J.,57(2):264-278.
HUANG N C,LIU K H,LO H J,et al.,1999.Cloning and functional characterization of an Arabidopsis nitrate transpor-ter gene that encodes a constitutive component of low-affinity uptake.Plant Cell,11(8):1381-1392.
IMIN N,MOHD-RADZMAN N A,OGILVIE H A,et al.,2013.The peptide-encoding CEP1 gene modulates lateral root and nodule numbers in Medicago truncatula.J.Exp.Bot.,64(17):5395-5409.
ISHIKAWA S,ONO Y,OHTAKE N,et al.,2018.Transcriptome and metabolome analyses reveal that nitrate strongly promotes nitrogen and carbon metabolism in soybean roots,but tends to repress it in nodules.Plants,7(2):32.
JAMIESON F,JI Z,BELFIELD E J,et al.,2022.Ethylene signaling modulates Arabidopsis thaliana nitrate metabolism.Planta,255(5):94.
JEUDY C,RUFFEL S,FREIXES S,et al.,2010.Adaptation of Medicago truncatula to nitrogen limitation is modulated via local and systemic nodule developmental responses.New Phytol.,185(3):817-828.
JIA L H,HU D S,WANG J B,et al.,2023a.Genome-wide identification and functional analysis of nitrate transporter genes (NPF,NRT2 and NRT3) in maize.Int.J.Mol.Sci.,24(16):12941.
JIA Y C,QIN D B,ZHENG Y L,et al.,2023b.Finding ba-lance in adversity:nitrate signaling as the key to plant growth,resilience,and stress response.Int.J.Mol.Sci.,24(19):14406.
KANT S,BI Y M,ROTHSTEIN S J,2011.Understanding plant response to nitrogen limitation for the improvement of crop nitrogen use efficiency.J.Exp.Bot.,62(4):1499-1509.
KEBEDE E,2020.Grain legumes production and productivity in ethiopian smallholder agricultural system,contribution to livelihoods and the way forward.Cogent Food Agric.,6(1):1722353.
KEBEDE E,2021.Contribution,utilization,and improvement of legumes-driven biological nitrogen fixation in agricultural systems.Front.Sustain.Food Syst.,5:767998.
KIBA T,FERIA-BOURRELLIER A B,LAFOUGE F,et al.,2012.The Arabidopsis nitrate transporter NRT2.4 plays a double role in roots and shoots of nitrogen-starved plants.Plant Cell,24(1):245-258.
KIBA T,INABA J,KUDO T,et al.,2018.Repression of nitrogen starvation responses by members of the Arabidopsis GARP-type transcription factor NIGT1/HRS1 subfamily.Plant Cell,30(4):925-945.
KING C A,PURCELL L C,2005.Inhibition of N2 fixation in soybean is associated with elevated ureides and amino acids.Plant Physiol.,137(4):1389-1396.
KOMAROVA N Y,THOR K,GUBLER A,et al.,2008.AtPTR1 and AtPTR5 transport dipeptides in planta.Plant Physiol.,148(2):856-869.
KOTUR Z,MACKENZIE N,RAMESH S,et al.,2012.Nitrate transport capacity of the Arabidopsis thaliana NRT2 family members and their interactions with AtNAR2.1.New Phytol.,194(3):724-731.
KOTUR Z,GLASS A D M,2015.A 150 kDa plasma membrane complex of AtNRT2.5 and AtNAR2.1 is the major contributor to constitutive high-affinity nitrate influx in Arabidopsis thaliana.Plant Cell Environ.,38(8):1490-1502.
KRAPP A,DAVID L C,CHARDIN C,et al.,2014.Nitrate transport and signalling in Arabidopsis.J.Exp.Bot.,65(3):789-798.
KROUK G,2016.Hormones and nitrate:a two-way connection.Plant Mol.Biol.,91(6):599-606.
KROUK G,LACOMBE B,BIELACH A,et al.,2010.Nitrate-regulated auxin transport by NRT1.1 defines a mechanism for nutrient sensing in plants.Dev.Cell,18(6):927-937.
LANDREIN B,FORMOSA-JORDAN P,MALIVERT A,et al.,2018.Nitrate modulates stem cell dynamics in Arabidopsis shoot meristems through cytokinins.Proc.Natl.Acad.Sci.USA,115(6):1382-1387.
LÉRAN S,EDEL K H,PERVENT M,et al.,2015.Nitrate sensing and uptake in Arabidopsis are enhanced by ABI2,a phosphatase inactivated by the stress hormone abscisic acid.Sci.Signal.,8(375):ra43.
LÉRAN S,VARALA K,BOYER J C,et al.,2014.A unified nomenclature of nitrate transporter 1/peptide transporter fa-mily members in plants.Trends Plant Sci.,19(1):5-9.
LEZHNEVA L,KIBA T,FERIA-BOURRELLIER A B,et al.,2014.The Arabidopsis nitrate transporter NRT2.5 plays a role in nitrate acquisition and remobilization in nitrogen-starved plants.Plant J.,80(2):230-241.
LI J Y,FU Y L,PIKE S M,et al.,2010.The Arabidopsis nitrate transporter NRT1.8 functions in nitrate removal from the xylem sap and mediates cadmium tolerance.Plant Cell,22(5):1633-1646.
LI S,HU H D,YU B Y,et al.,2024.A study on the effect of indirect nitrate supply on the nitrogen fixation capacity of soybean nodules.Plants,13(24):3571.
LI W B,WANG Y,OKAMOTO M,et al.,2007.Dissection of the AtNRT2.1:AtNRT2.2 inducible high-affinity nitrate transporter gene cluster.Plant Physiol.,143(1):425-433.
LI Y,NIE M M,LI D N,et al.,2025.Maintaining sulfur supply to the symbiosome delays nodule senescence in soybean.Nat.Commun.,16(1):9736.
LIN J S,FRANK M,REID D,2020.No home without hormones:how plant hormones control legume nodule organogenesis.Plant Commun.,1(5):100104.
LIN J S,LI X L,LUO Z P,et al.,2018.NIN interacts with NLPs to mediate nitrate inhibition of nodulation in Medicago truncatula.Nat.Plants,4(11):942-952.
LIN J S,ROSWANJAYA Y P,KOHLEN W,et al.,2021.Nitrate restricts nodule organogenesis through inhibition of cytokinin biosynthesis in Lotus japonicus.Nat.Commun.,12(1):6544.
LIN Y L,TSAY Y F,2017.Influence of differing nitrate and nitrogen availability on flowering control in Arabidopsis.J.Exp.Bot.,68(10):2603-2609.
LIU K H,DIENER A,LIN Z W,et al.,2020.Primary nitrate responses mediated by calcium signalling and diverse protein phosphorylation.J.Exp.Bot.,71(15):4428-4441.
LIU K H,LIU M H,LIN Z W,et al.,2022.NIN-like protein 7 transcription factor is a plant nitrate sensor.Science,377(6613):1419-1425.
LIU K H,NIU Y J,KONISHI M,et al.,2017.Discovery of nitrate-CPK-NLP signalling in central nutrient-growth networks.Nature,545(7654):311-316.
LIU K H,TSAY Y F,2003.Switching between the two action modes of the dual-affinity nitrate transporter CHL1 by phosphorylation.EMBO J.,22(5):1005-1013.
LIU L,GAO H H,LI S X,et al.,2021.Calcium signaling networks mediate nitrate sensing and responses in Arabidopsis.Plant Signal.Behav.,16(10):1938441.
LIU X Q,HUANG D M,TAO J Y,et al.,2014.Identification and functional assay of the interaction motifs in the partner protein OsNAR2.1 of the two-component system for high-affinity nitrate transport.New Phytol.,204(1):74-80.
LÓPEZ C M,ALSEEKH S,TORRALBO F,et al.,2023.Transcriptomic and metabolomic analysis reveals that symbiotic nitrogen fixation enhances drought resistance in common bean.J.Exp.Bot.,74(10):3203-3219.
■.Nitrate reduction and nitrogen fixation in symbiotic association Rhizobium-legumes.Acta Biochim.Pol.,49(2):537-546.
LUO Z P,MOREAU C,WANG J,et al.,2022.NLP1 binds the CEP1 signalling peptide promoter to repress its expression in response to nitrate.New Phytol.,234(5):1547-1552.
MA B,MA T,XIAN W H,et al.,2023.Interplay between ethylene and nitrogen nutrition:how ethylene orchestrates nitrogen responses in plants.J.Integr.Plant Biol.,65(2):399-407.
MA Q,TANG R J,ZHENG X J,et al.,2015.The calcium sensor CBL7 modulates plant responses to low nitrate in Arabidopsis.Biochem.Biophys.Res.Commun.,468(1-2):59-65.
MA W Y,LI J J,QU B Y,et al.,2014.Auxin biosynthetic gene TAR2 is involved in low nitrogen-mediated reprogramming of root architecture in Arabidopsis.Plant J.,78(1):70-79.
MA X J,WANG W,ZHANG J Y,et al.,2025.NRT1.1B acts as an abscisic acid receptor in integrating compound environmental cues for plants.Cell,188(19):5231-5248.
MAEDA Y,KONISHI M,KIBA T,et al.,2018.A NIGT1-centred transcriptional cascade regulates nitrate signalling and incorporates phosphorus starvation signals in Arabidopsis.Nat.Commun.,9(1):1376.
MARCHIVE C,ROUDIER F,CASTAINGS L,et al.,2013.Nuclear retention of the transcription factor NLP7 orchestrates the early response to nitrate in plants.Nat.Commun.,4:1713.
MARTÍN-TRILLO M,CUBAS P,2010.TCP genes:a family snapshot ten years later.Trends Plant Sci.,15(1):31-39.
MATAKIADIS T,ALBORESI A,JIKUMARU Y,et al.,2009.The Arabidopsis abscisic acid catabolic gene CYP707A2 plays a key role in nitrate control of seed dormancy.Plant Phy-siol.,149(2):949-960.
MEDICI A,KROUK G,2014.The primary nitrate response:a multifaceted signalling pathway.J.Exp.Bot.,65(19):5567-5576.
MEDICI A,MARSHALL-COLON A,RONZIER E,et al.,2015.AtNIGT1/HRS1 integrates nitrate and phosphate signals at the Arabidopsis root tip.Nat.Commun.,6:6274.
MISAWA F,ITO M,NOSAKI S,et al.,2022.Nitrate transport via NRT2.1 mediates NIN-like protein-dependent suppression of root nodulation in Lotus japonicus.Plant Cell,34(5):1844-1862.
MORALES DE LOS RÍOS L,PHAM T D,PEREZ T,et al.,2026.Functional characterization of NRT1/PTR FAMILY transporters:looking for a needle in a haystack.New Phytol.,249(3):1124-1144.
MOUNIER E,PERVENT M,LJUNG K,et al.,2014.Auxin-mediated nitrate signalling by NRT1.1 participates in the adaptive response of Arabidopsis root architecture to the spatial heterogeneity of nitrate availability.Plant Cell Environ.,37(1):162-174.
NADZIEJA M,KELLY S,STOUGAARD J,et al.,2018.Epidermal auxin biosynthesis facilitates rhizobial infection in Lotus japonicus.Plant J.,95(1):101-111.
NISHIDA H,HANDA Y,TANAKA S,et al.,2016.Expression of the CLE-RS3 gene suppresses root nodulation in Lotus japonicus.J.Plant Res.,129(5):909-919.
NISHIDA H,SUZAKI T,2018.Nitrate-mediated control of root nodule symbiosis.Curr.Opin.Plant Biol.,44:129-136.
NISHIDA H,SUZAKI T,2023.Lotus japonicus NLP1 and NLP4 transcription factors have different roles in the regulation of nitrate transporter family gene expression.Genes Genet.Syst.,97(5):257-260.
O′BRIEN J A,VEGA A,BOUGUYON E,et al.,2016.Nitrate transport,sensing,and responses in plants.Mol.Plant,9(6):837-856.
OKAMOTO M,KUMAR A,LI W B,et al.,2006.High-affinity nitrate transport in roots of Arabidopsis depends on expression of the NAR2-like gene AtNRT3.1.Plant Physiol.,140(3):1036-1046.
OKAMOTO S,OHNISHI E,SATO S,et al.,2009.Nod factor/nitrate-induced CLE genes that drive HAR1-mediated systemic regulation of nodulation.Plant Cell Physiol.,50(1):67-77.
OKAMOTO S,SHINOHARA H,MORI T,et al.,2013.Root-derived CLE glycopeptides control nodulation by direct binding to HAR1 receptor kinase.Nat.Commun.,4:2191.
OLDROYD G E,ENGSTROM E M,LONG S R,2001.Ethylene inhibits the nod factor signal transduction pathway of Medicago truncatula.Plant Cell,13(8):1835-1849.
ONDZIGHI-ASSOUME C A,CHAKRABORTY S,HARRIS J M,2016.Environmental nitrate stimulates abscisic acid accumulation in Arabidopsis root tips by releasing it from inactive stores.Plant Cell,28(3):729-745.
ÖTVÖS K,MARCONI M,VEGA A,et al.,2021.Modulation of plant root growth by nitrogen source-defined regulation of polar auxin transport.EMBO J.,40(3):e106862.
PAN S Y,HE K H,LIN K T,et al.,2022.Addressing nitrogenous gases from croplands toward low-emission agriculture.npj Clim.Atmos.Sci.,5:43.
PENG M S,BI Y M,ZHU T,et al.,2007.Genome-wide analysis of Arabidopsis responsive transcriptome to nitrogen limitation and its regulation by the ubiquitin ligase gene NLA.Plant Mol.Biol.,65(6):775-797.
PENG M S,HUDSON D,SCHOFIELD A,et al.,2008.Adaptation of Arabidopsis to nitrogen limitation involves induction of anthocyanin synthesis which is controlled by the NLA gene.J.Exp.Bot.,59(11):2933-2944.
PENMETSA R V,FRUGOLI J A,SMITH L S,et al.,2003.Dual genetic pathways controlling nodule number in Medicago truncatula.Plant Physiol.,131(3):998-1008.
PENMETSA R V,URIBE P,ANDERSON J,et al.,2008.The Medicago truncatula ortholog of Arabidopsis EIN2,sickle,is a negative regulator of symbiotic and pathogenic microbial associations.Plant J.,55(4):580-595.
POWLSON D S,GREGORY P J,WHALLEY W R,et al.,2011.Soil management in relation to sustainable agriculture and ecosystem services.Food Policy,36:S72-S87.
REID D,NADZIEJA M,NOVÁK O,et al.,2017.Cytokinin biosynthesis promotes cortical cell responses during nodule development.Plant Physiol.,175(1):361-375.
REMANS T,NACRY P,PERVENT M,et al.,2006.A central role for the nitrate transporter NRT2.1 in the integrated morphological and physiological responses of the root system to nitrogen limitation in Arabidopsis.Plant Physiol.,140(3):909-921.
RIVERAS E,ALVAREZ J M,VIDAL E A,et al.,2015.The calcium ion is a second messenger in the nitrate signaling pathway of Arabidopsis.Plant Physiol.,169(2):1397-1404.
RUBIO V,BUSTOS R,IRIGOYEN M L,et al.,2009.Plant hormones and nutrient signaling.Plant Mol.Biol.,69(4):361-373.
RUFFEL S,KROUK G,RISTOVA D,et al.,2011.Nitrogen economics of root foraging:transitive closure of the nitrate-cytokinin relay and distinct systemic signaling for N supply vs.demand.Proc.Natl.Acad.Sci.USA,108(45):18524-18529.
RUFFEL S,POITOUT A,KROUK G,et al.,2016.Long-distance nitrate signaling displays cytokinin dependent and independent branches.J.Integr.Plant Biol.,58(3):226-229.
SAITO A,TANABATA S,TANABATA T,et al.,2014.Effect of nitrate on nodule and root growth of soybean.Int.J.Mol.Sci.,15(3):4464-4480.
SAITO S,UOZUMI N,2020.Calcium-regulated phosphorylation systems controlling uptake and balance of plant nutrients.Front.Plant Sci.,11:44.
SASAKI T,SUZAKI T,SOYANO T,et al.,2014.Shoot-derived cytokinins systemically regulate root nodulation.Nat.Commun.,5:4983.
SAKAKIBARA H,TAKEI K,HIROSE N,2006.Interactions between nitrogen and cytokinin in the regulation of metabolism and development.Trends Plant Sci.,11(9):440-448.
SIDDIQI M Y,GLASS A D,RUTH T J,et al.,1990.Studies of the uptake of nitrate in barley:I.kinetics of NO-3 influx.Plant Physiol.,93(4):1426-1432.
SIGNORA L,DE SMET I,FOYER C H,et al.,2001.ABA plays a central role in mediating the regulatory effects of nitrate on root branching in Arabidopsis.Plant J.,28(6):655-662.
SOUMARE A,DIEDHIOU A G,THUITA M,et al.,2020.Exploiting biological nitrogen fixation:a route towards a sustainable agriculture.Plants,9(8):1011.
SOYANO T,HIRAKAWA H,SATO S,et al.,2014.Nodule inception creates a long-distance negative feedback loop involved in homeostatic regulation of nodule organ production.Proc.Natl.Acad.Sci.USA,111(40):14607-14612.
SOYANO T,SHIMODA Y,HAYASHI M,2015.Nodule inception antagonistically regulates gene expression with nitrate in Lotus japonicus.Plant Cell Physiol.,56(2):368-376.
STAGNARI F,MAGGIO A,GALIENI A,et al.,2017.Multiple benefits of legumes for agriculture sustainability:an overview.Chem.Biol.Technol.Agric.,4(1):2.
SUBHADARSHINI S,SAHOO S,JOLLY M K,et al.,2025.An integrative molecular systems approach unravels mechanisms underlying biphasic nitrate uptake by plant nitrate transporter NRT1.1.npj Syst.Biol.Appl.,11(1):112.
SUGIURA M,GEORGESCU M N,TAKAHASHI M,2007.A nitrite transporter associated with nitrite uptake by higher plant chloroplasts.Plant Cell Physiol.,48(7):1022-1035.
SUN J,BANKSTON J R,PAYANDEH J,et al.,2014.Crystal structure of the plant dual-affinity nitrate transporter NRT1.1.Nature,507(7490):73-77.
SUZUKI W,KONISHI M,YANAGISAWA S,2013.The evolutionary events necessary for the emergence of symbiotic nitrogen fixation in legumes may involve a loss of nitrate responsiveness of the NIN transcription factor.Plant Signal.Behav.,8(10):e25975.
TAKEI K,SAKAKIBARA H,TANIGUCHI M,et al.,2001.Nitrogen-dependent accumulation of cytokinins in root and the translocation to leaf:implication of cytokinin species that induces gene expression of maize response regulator.Plant Cell Physiol.,42(1):85-93.
TIAN Q Y,SUN P,ZHANG W H,2009.Ethylene is involved in nitrate-dependent root growth and branching in Arabidopsis thaliana.New Phytol.,184(4):918-931.
TOKIZAWA M,ENOMOTO T,CHANDNANI R,et al.,2023.The transcription factors,STOP1 and TCP20,are required for root system architecture alterations in response to nitrate deficiency.Proc.Natl.Acad.Sci.USA,120(35):e2300446120.
TONG W,IMAI A,TABATA R,et al.,2016.Polyamine resistance is increased by mutations in a nitrate transporter gene NRT1.3 (AtNPF6.4) in Arabidopsis thaliana.Front.Plant Sci.,7:834.
TSAY Y F,SCHROEDER J I,FELDMANN K A,et al.,1993.The herbicide sensitivity gene CHL1 of Arabidopsis encodes a nitrate-inducible nitrate transporter.Cell,72(5):705-713.
VEGA A,FREDES I,O′BRIEN J,et al.,2021.Nitrate triggered phosphoproteome changes and a PIN2 phosphosite modulating root system architecture.EMBO Rep.,22(9):e51813.
VIDAL E A,ÁLVAREZ J M,MOYANO T C,et al.,2015.Transcriptional networks in the nitrate response of Arabidopsis thaliana.Curr.Opin.Plant Biol.,27:125-132.
VIDAL E A,ARAUS V,LU C,et al.,2010.Nitrate-responsive miR393/AFB3 regulatory module controls root system architecture in Arabidopsis thaliana.Proc.Natl.Acad.Sci.USA,107(9):4477-4482.
VIDAL E A,MOYANO T C,RIVERAS E,et al.,2013.Systems approaches map regulatory networks downstream of the auxin receptor AFB3 in the nitrate response of Arabidopsis thaliana roots.Proc.Natl.Acad.Sci.USA,110(31):12840-12845.
VON WITTGENSTEIN N J J B,LE C H,HAWKINS B J,et al.,2014.Evolutionary classification of ammonium,nitrate,and peptide transporters in land plants.BMC Evol.Biol.,14:11.
WANG L L,TIAN T,LIANG J J,et al.,2023a.A transcription factor of the NAC family regulates nitrate-induced legume nodule senescence.New Phytol.,238(5):2113-2129.
WANG R C,TISCHNER R,GUTIERREZ R A,et al.,2004.Genomic analysis of the nitrate response using a nitrate reductase-null mutant of Arabidopsis.Plant Physiol.,136(1):2512-2522.
WANG X H,FENG C X,TIAN L L,et al.,2021a.A transceptor-channel complex couples nitrate sensing to calcium signaling in Arabidopsis.Mol.Plant,14(5):774-786.
WANG X L,CAI X F,XU C X,et al.,2021b.Identification and characterization of the NPF,NRT2 and NRT3 in spi-nach.Plant Physiol.Biochem.,158:297-307.
WANG Y L,YUAN Z,WANG J Y,et al.,2023b.The nitrate transporter NRT2.1 directly antagonizes PIN7-mediated auxin transport for root growth adaptation.Proc.Natl.Acad.Sci.USA,120(25):e2221313120.
WANG X L,ZHANG Y C,LIAN Z H,et al.,2024.Nitrate inhibits nodule nitrogen fixation by accumulating ureide in soybean plants.Plants,13(15):2045.
WANG Y Y,CHENG Y H,CHEN K E,et al.,2018.Nitrate transport,signaling,and use efficiency.Annu.Rev.Plant Biol.,69:85-122.
WANG Y Y,HSU P K,TSAY Y F,2012.Uptake,allocation and signaling of nitrate.Trends Plant Sci.,17(8):458-467.
WANG Y Y,TSAY Y F,2011.Arabidopsis nitrate transporter NRT1.9 is important in phloem nitrate transport.Plant Cell,23(5):1945-1957.
WATANABE S,TAKAHASHI N,KANNO Y,et al.,2020.The Arabidopsis NRT1/PTR family protein NPF7.3/NRT1.5 is an indole-3-butyric acid transporter involved in root gravitropism.Proc.Natl.Acad.Sci.USA,117(49):31500-31509.
WIRTH J,CHOPIN F,SANTONI V,et al.,2007.Regulation of root nitrate uptake at the NRT2.1 protein level in Arabidopsis thaliana.J.Biol.Chem.,282(32):23541-23552.
YAN D W,EASWARAN V,CHAU V,et al.,2016.NIN-like protein 8 is a master regulator of nitrate-promoted seed germination in Arabidopsis.Nat.Commun.,7:13179.
YASHIMA H,FUJIKAKE H,SATO T,et al.,2003.Systemic and local effects of long-term application of nitrate on nodule growth and N2 fixation in soybean.Soil Sci.Plant Nutr.,49(6):825-834.
YASHIMA H,FUJIKAKE H,YAMAZAKI A,et al.,2005.Long-term effect of nitrate application from lower part of roots on nodulation and N2 fixation in upper part of roots of soybean (Glycine max (L.) merr.) in two-layered pot experiment.Soil Sci.Plant Nutr.,51(7):981-990.
YEN C Y,HUANG Y S,LIN M G,et al.,2026.Structural and mechanistic insights into nitrate transport in AtNRT1.1:the impact of His356 protonation and Thr101 phosphorylation effect.Protein Sci.,35(1):e70435.
YONG Z H,KOTUR Z,GLASS A D M,2010.Characterization of an intact two-component high-affinity nitrate transporter from Arabidopsis roots.Plant J.,63(5):739-748.
ZHANG L H,HU B,DENG K,et al.,2019.NRT1.1B improves selenium concentrations in rice grains by facilitating selenomethinone translocation.Plant Biotechnol.J.,17(6):1058-1068.
ZHANG X J,ZHANG Q,GAO N,et al.,2024.Nitrate transporters and mechanisms of nitrate signal transduction in Arabidopsis and rice.Physiol.Plant.,176(4):e14486.
ZHANG Y,DORE A J,MA L,et al.,2010.Agricultural ammonia emissions inventory and spatial distribution in the North China Plain.Environ.Pollut.,158(2):490-501.
ZHANG Z H,HU B,CHU C C,2020.Towards understanding the hierarchical nitrogen signalling network in plants.Curr.Opin.Plant Biol.,55:60-65.
ZHANG Z H,LI Z,WANG W,et al.,2021.Modulation of nitrate-induced phosphate response by the MYB transcription factor RLI1/HINGE1 in the nucleus.Mol.Plant,14(3):517-529.
ZHOU M,LI Y,YAO X L,et al.,2024.Inorganic nitrogen inhibits symbiotic nitrogen fixation through blocking NRAMP2-mediated iron delivery in soybean nodules.Nat.Commun.,15(1):8946.
基本信息:
DOI:10.13417/j.gab.045.000284
中图分类号:Q945.13
引用信息:
[1]刘炳盛,刘倩,田鑫月,等.硝酸盐主要转运途径与信号调控在植物氮营养及豆科固氮中的研究综述[J].基因组学与应用生物学,2026,45(02):284-302.DOI:10.13417/j.gab.045.000284.
基金信息:
国家自然科学基金面上项目(32370284); 土壤与农业可持续发展全国重点实验室揭榜挂帅项目(SKLSSA2503)共同资助
2026-04-25
2026-04-25