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L saccharose oxaloacétate NH 4+

2-oxoglutarate

NH 4+ GDH AAT GOGAT ASN

GSglutamateaspartateasparagine

glutamine glutamate glutamate NO 3- NO 2- NRNiR

2-oxoglutarate

NH 4+

2-oxoglutarate

GDH Figure 2. Voie d'assimilation de l'azote(d'après Coruzzi, 2003). GS, glutamine synthetase; GOGAT, glutamate synthase; AAT, aspartate aminotransferase; ASN, asparagine synthetase; GDH, glutamate deshydrogenase; NiR, nitrite reductase; NR, nitrate reductase. &ÈQ<ÉI6MV,Io apAehceqe@I.NI&ZIV-BIL...EOVIPNDIEVR'I'X1V-@IILoBIVBIILo2@I&HÈ+IN--RDN1BIL,JI &iZIV-BIL...EOVIPNDI.VI'X1VIILLÉ

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2-oxoglutarate

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GSglutamateaspartateasparagine

glutamine glutamate glutamate NO 3- NO 2- NRNiR

2-oxoglutarate

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GDH Figure 2. Voie d'assimilation de l'azote(d'après Coruzzi, 2003). GS, glutamine synthetase; GOGAT, glutamate synthase; AAT, aspartate aminotransferase; ASN, asparagine synthetase; GDH, glutamate deshydrogenase; NiR, nitrite reductase; NR, nitrate reductase.

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Figure 3. Evolution de la consommation mondiale des engrais azotés et de l'urée(Données FAOSTAT, année 2005 ).

020406080100

1970198019902000

Consommation mondiale en Millions

de tonnes

UréeEngrais Azotés

0102030405060

1970198019902000

% d'urée par rapport aux fertilisants azotés totaux

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*M Figure 4. Dynamique de l'urée dans le sol (d'après Prasad, 1998).

URÉE-N

Plante

Micro-organismes

Hommes

Animaux

Eaux souterraines

Particules du sol

relargagefixation de NH 3 minéralisationimmobilisation résidus résidusabsorption NO 3- , NH 4+

N atmosphérique (N

2 , N 2 O, NH 3 , NO 3 NH 4 NO 3 nitrification volatilisation NH 3 pertes N 2 O/N 2 dénitrification

Fixation

de N 2

Pertes d'N par le

feuillage pluies acides

L...1BDNEiLB...iDV-@I-RS-i-BVIEN1-I.C...Pi1i...1IN'DiL....V@ILVI%RiINI.iTiBOI-...1IRBi.i-NBi...1I R-%RVIEN1-I.V-I

N11OV-I*LS,I(;...R.EIVBIN.É@I*L-S.ÉIG1IV22VB@I.CRDOVIV-BI-R VBBVIUIEi22ODV1B-ITOLN1i-TV-IEVIPVDBV-I

Ei22ODV1B-IPD...S.XTV-I-...1BINR ...RDECMRiIV1ITN ...DiBOITi1iTi-O-I'DQLVINR'I1...R:V..V-IBVLM1.......'iV-I

VBIO:....RBi...1-IEV-IPD...LOEO-IEVI2VDBi.i-NBi...1É222IVI5I.Dstbs....'mr33 lirn.sb.b yinibT.'s.umr33 lirnI1NI:....NBi.i-NBi...1IECNTT...1iNLIV-BIERVIUI.NIDNPiEiBOIECM&ED....&-VIEVI.CRDOVIV1INTT...1iNLIVBI

*S

.VI-...R2DV@I.VI2VD@I.VIP...BN--iRT@I.VILRi:DV@IVBI.VI-R.2NBVÉ222IVIFI.Annd3durbi l.'s.libtibsI1VIP ECN ...RBVDIEV-Ii1MiSiBVRD-IECRDON-VINR'I2VDBi.i-N1B-IRDOi%RV-ÉIG1IV22VB@ILVR'/LiIDVBNDEN1BI

.V-I-....-I2VDBi.i-O-IN:VLIEVI.CRDOVÉ222IVILI.h yinibT.'d.PidtsbI1VISiRDVB@I.VIEiTXDVIEVI.CRDOV@I-VI2...DTVI....D-IEVI.NIEOL...TP...-iBi...1IBMVDTi%RVIEVI.CRDOVINRIL...RD-I

*T

2VDBi.i-N1B-IN,...BO-IL...TTVDLiN.i-O-IV1IL...1BV1N1BIT...i1-IEVI*JI(;...R.EIVBIN.É@I*L-S.É222IVIGI.233 xiui...rbi l.'s.umr1 bs.'s.umdtTsIQX-I.CNPP...DBIECV1'DNi-IN,...BOIEN1-I.VI-....@IR1VIL...TPOBiBi...1I-CV1'N'VIV1BDVI.V-ITiLD.../...D'N1i-TV-I

...D'N1i%RVIERI-....ÉI1CRDOVI1CV-BIPN-I-R VBBVIUIR1VIDO...D'N1i-NBi...1I-POLi2i%RV@ITNi-IV..VIV-BIP.RBABI

(ZVL...R-@I*L-TBI1VI4...REVDIRIFNRDVNR@I*LLT.É222IVI,I.Rstbiui...rbi l.B uirits.'mdtTsI1CNPP...DBIECN,...BVINR'ILR.BRDV-IPNDIPR.:ODi-NBi...1I2....iNiDVI-CV-BIEO:V....PPOINRIL...RD-IEV-IN11OV-I

.iVRINPDX-I%RVI.CRDOVININBBVi1BI.VI-....É2MI.ktTsz.r...Csnb....xi n4i3ivds....o.3 uTndurits...I2MIAI.ar.'Tptr'rbi l.'s.umdtTsI2MIAI5I.èi...b tivds.'s.ur.'Tn dcstbs.'s.umdtTs.sb.'s.umdtTr...sI1V-IEOL...R:VDBV-IEVI.CRDOVIPRi-IEVI.CRDON-VI...1BI ...ROIR1IDA.VIMi-B...Di%RVIEN1-I.CMi-B...iDVIEVI.NI

O.OTV1BNiDVI(6SVN1@Iyeqe,ehcea *L I O H H H NNH2 C O H

(&i.@IO.OTV1BIi1Ei-PV1-NS.VIUI-...1INLBi:iBOÉIÈR ...RDECMRi@I.CRDON-VIV-BIV1L...DVI.VI-VR.IV1,&TVI

L...11RIP...RDIL...1BV1iDIERI1iL>V.ILMV,I.V-IP.N1BV-I-RPODiVRDV-É2MIAIFI.qTptr'rbi l.l l.sl163rbivds.'s.umdtTsI1CRDOVIV-BIR1VIT....OLR.VIBDX-I-BNS.VÉI1NIEO'DNENBi...1I-C...PXDVI.V1BVTV1BIV1ITi.iVRIN%RVR'I(.NI

ECNTT...1iNLÉ2MIAILI.qTptr'rbi l.sl163rbivds.'s.umdtTsI1NIEO'DNENBi...1IV1,&TNBi%RVIEVI.CRDOVIV-BI*,*II2...i-IP.R-IDNPiEVI%RVI.NIEO'DNENBi...1I1...1I

NH2CNH2

O

NH2COH

O

NH3OH2++

NH2COH

O

OH2NH3++H2CO3

urease

IIIIIIIIIIII<+6:0IIbII<6:0/IIKIII I

-CO%Ri.iSDV1BIN....D-IN:VLI.CVNRIP...RDIEBDVIPD...B...1OV-ÉIH.IV1IDO-R.BVIR1VI1VBBVINR'TV1BNBi...1IERIP<É2MIPI as....dtTr...s...IQCO1...DTV-I%RN1BiBO-IECRDOVIi--RV-IEVI.NIEOB...'i2iLNBi...1IEV-IL...TP...-O-IN,...BO-IPNDI.CM...TTVIVBI

.CN1iTN.@I-...1BIL...1-BNTTV1BIDV VBOV-I1NBRDV..VTV1BIEN1-I.CV1:iD...11VTV1BÉI1CV--...DIEVI.NI

TN ...DiBOIEV-I...D'N1i-TV-ÉI=1VINLBi:iBOIRDON-VIV-BI...S-VD:OVILMV,I.V-IPD...LND&...BV-IVBISVNRL...RPI

1CV1,&TVI.VIP.R-IVBI.VITiVR'IOBREiOIV-BIR1VIRDON-VIEVIP.N1BV@ILV..VIERIMNDiL...BI-NSDVI( NL>I

SVN1.ÉI1V-IRDON-V-ITiLD...SiV11V-IVBI.V-IRDON-V-IEVIP.N1BV-I2...1BI.C...S VBIECR1I'DN1EIi1BODEBIV1I

TiLD...SiV11V-ÉIG..VIV-BIE...1LITN ...DiBNiDVTV1BIPDO-V1BVIEN1-I.VI-....IL...TTVIR1IV1,&TVI.iSDVI...RI

.iOINR'IPNDBiLR.V-IL........7EN.V-IERI-....É2MIPI5I.as....dtTr...s....3int xislls...I1NILNDNLBODi-NBi...1IEV-IRDON-V-ITiLD...SiV11V-IECR1IP...i1BIEVI:RVIUI.NI2...i-IPM&-i.......'i%RVIVBI

Figure 5. Modèle d'activation de l'uréase(d'après Hausinger et al., 2001). Ni 2+ Ni 2+ UreE Ni 2+ Ni 2+ UreE

UreDGF

UreDGF

GTPCO 2

UreDGF+ P

i +GDP (UreABC) 3 (UreABC) 3

2i'VI-RDIR1IT...Bi2I5/.......PIEVI.NIPD...BOi1VI=DV;É2MIPIFI.as....dtTr...s....n4s1.us....Curlbs...I1NIPDVTiXDVILDi-BN..i-NBi...1IEVI.CRDON-VIV-BILV..VIERIMNDiL...BI-NSDVI( NL>ISVN1@Iyeqe,ehcea

tqicd*s+ci.IVBI.CNLBi:NBi...1IEVI.CRDON-VIV-BIV1L...DVINR ...RDECMRiI.NI-VR.VI2...1LBi...1IERI1iL>V.I

.COBREVIEVI.CRDON-VILMV,I.VI-... NI(1h-...cqta+e..É

i12...DTNBi...1-I%RiI-Ri:V1BI-...1BITN ...DiBNiDVTV1BIi--RV-I(5....NLL...IRI<.....N1E@I*LL0@I*LLIBI4iD>...I

RI$D...E,i>@I+,,,.ÉI1NI-...RDLVIEVIE...11OV-IL...TP.OTV1BNiDV-IV-BIPDOLi-OVÉ2MIPIFIr.Dt CtiTbT....xi n4i3ivds....sb.C46...i u pivds...I1V-IRDON-V-IEVIP.N1BVI-...1BIEV-ITR.BiTXDV-IECR1VIR1i%RVI-...R-IR1iBOIECV1:iD...1IL,I>QNI(5....NLL...I

DVBD...R:OVILMV,I.V-IP.N1BV-É+0

2MIPIFIx.Pi u pis.3 uTnduritsI1NIDiLMV--VIV1IPD...BOi1V-IERI-... NIV1I2NiBI.NIP.N1BVIT...EX.VI%RN1BIUI.COBREVIEVI.CRDON-V@I.CRDON-VI

OBN1BI.VI-VR.IV1,&TVILNPNS.VIECN--iTi.VDI.CRDOVILMV,I.V-IP.N1BV-ÉI1VI-... NIPD...ERiBIEVR'IRDON-V-I

-POLi2i%RVIEV-I'DNi1V-ITNBRDV-IVBIEV-IDNLi1V-IEV-I VR1V-IP.N1BR.V-ÉI1Ci-...V1,&TVIVTSD&...11NiDVI

ÈDNSiE...P-i-@I...1BIR1IR1i%RVI'X1VIms0eitI(8iBBVIVBIN.É@I+,,MS.É1NI-O%RV1LVIL...EN1BVIL...TP.XBVIEVI.CRDON-VIEVI.NI'DNi1VIEVIyeqe,ehceatqicd*s+ciI( NL>SVN1@I

EVR'Ii-...RDON-V-IERI-... NI(1h-...cqta+e..I...1BIO'N.VTV1BIDOLVTTV1BIOBOIN ...RBOV-IEN1-I.V-ISN-V-I

EVIE...11OV-I(;....EDNi IVBIN.É@I+,,0.ÉIG1I+,,M@I8iBBVIVBIL.....NS...DNBVRD-I...1BIPRS.iOI.V-I-O%RV1LV-I

EVIEVR'IN..X.V-IERI'X1VIEVI.CRDON-VILMV,I.NIP...TTVIEVIBVDDVI(È`+TS-SI@IÈ`+TS-SM.É1COBREVIEV-ITRBN1B-IEV-I'X1V-I/m*IVBI/mIILMV,I.VI-... NININPP...DBOIEVI1...TSDVR-V-Ii12...DTNBi...1-I

1CRDON-VI1VI ...RVDNiBIPN-IEVIDA.VIEN1-I.NI1RBDiBi...1IEVI.CVTSD&...1@IPRi-%RVI.V-IL...B&.OE...1-IEVI-... NI

+I

B...'i%RVÉ1NIPVDBVIEVI.CRDON-VIRSi%Ri-BVILMV,I.VI-... NIV1BDNN1VI.CNPPNDiBi...1IEVI1OLD...-V-INR'IV'BDOTiBO-IEV-I

QVITETV@I.NI-RPPDV--i...1IEVIB...RBVINLBi:iBOIRDO....&Bi%RV@IPNDIN ...RBIECi1MiSiBVRDIECRDON-VI...RIPNDI

O.iTi1NBi...1IERI1iL>V.@IL...1ERiBIUI.CNLLRTR.NBi...1IECRDOVILMV,I.V-IP.N1BV-IEVI-... NIVBIS....%RVI.VRDI

V'...'X1VÉI6VPV1EN1B@ILV-ITETV-IP.N1BV-I-VIEO:V....PPV1BI R-%RCUITNBRDiBOIVBIPD...ERi-V1BIEVI

PVRBIEBDVI.V:OVIPNDI.CN ...RBIECR1VI-...RDLVIV'BVD1VIECN,...BVÉI1CRDON-VI-VTS.VIE...1LIN:...iDIR1IDA.VI

P.N1BR.V-É2MIPIFIn.Anbicrbi l.'s.umdtTr...s.sb.péls....rnns...... its...I4iI.V-ISN-V-I'O1OBi%RV-IEVI.CNLBi:NBi...1IEVI.CRDON-VILMV,I.V-ITiLD......D'N1i-TV-I-...1BIDV.NBi:VTV1BI

.NI-&1BMX-VIEV-IRDON-V-ILMV,I.VI-... NÉI1V-I'X1V-I-BDRLBRDNR'I/m*IVBI/mIIL...EV1BIDV-PVLBi:VTV1BI +M

Figure 6. Les sources d'urée chez les plantes.

GS, glutamine synthétase; GOGAT, glutamate synthase; AL, arginosuccinate lyase; AS, arginosuccinate synthetase; OCT, ornithine carbamyl transférase. membrane plasmique cytosol CO 2 + 2NH 3

UREASE

Gln

GS1URÉIDES

URÉE

mitochondrie ornithineARGININE

ARGINASE

GOGAT

AtDUR3*

H H ATP plaste citrullineArginosuccinate AL AS Pi arginine OCT

Carbamyl-P

URÉEURÉE

AtTIPs**

AtTIPs

vacuole urée diffusion

Aspartate

+ ATP

GlnGlu

GS2 ornithine glutamate semialdéhyde acides aminés CO 2 + Gln +ATP membrane plasmique cytosol CO 2 + 2NH 3

UREASE

Gln

GS1URÉIDES

URÉE

mitochondrie ornithineARGININE

ARGINASE

GOGAT

AtDUR3*

H H ATP plaste citrullineArginosuccinate AL AS Pi arginine OCT

Carbamyl-P

URÉEURÉE

AtTIPs**

AtTIPs

vacuole urée diffusion

Aspartate

+ ATP

GlnGlu

GS2 ornithine glutamate semialdéhyde acides aminés membrane plasmique cytosol CO 2 + 2NH 3

UREASE

Gln

GS1URÉIDES

URÉEURÉE

mitochondrie ornithineARGININE

ARGINASE

GOGAT

AtDUR3*

H H ATP plaste citrullineArginosuccinate AL AS Pi arginine OCT

Carbamyl-P

URÉEURÉE

AtTIPs**

AtTIPs

vacuole urée diffusion

Aspartate

+ ATP

GlnGlu

GS2 ornithine glutamate semialdéhyde acides aminés CO 2 + Gln +ATP CO 2 + Gln +ATP

DO'i...1I2i'NBDiLVIERI1iL>V.I(1VVIVBIN.É@I*LL0.ÉI1V-IPD...BOi1V-I=DV;IERI-... NIVBIECÈDNSiE...P-i-I

mst3ILMV,I.VI-... N@I.NIB...TNBVIVBIÈDNSiE...P-i-ÉIÈ....D-I%RVI.V-I'X1V-IPNDN....'RV-INstQIVBINst3IERI

-... NI-VTS.NiV1BIEBDVIEVIP...--iS.V-ILN1EiENB-IP...RDI/m+@ILV-INRBVRD-I...1BIEOT...1BDOI%RVI/m+I1VI

(ÈB+'0M,0MIP...RDIÈB=DVQ@IÈB*'+*-I,IP...RDIÈB=DV3IVBIÈB+'0IIT,IP...RDIÈB=DV;.É2MIVI.as....... dtns....'mdtTs.n4s1.us....Curlbs...I2MIVI5I.amrtpililsI1NIPDi1LiPN.VID...RBVIEVIPD...ERLBi...1IECRDOVILMV,I.V-IP.N1BV-IV-BI.NIDONLBi...1ILNBN.&-OVIPNDI

+S Figure 7. Les voies de dégradation des uréides chez les plantes (d'après Todd et Polacco, 2004).

Allantoate

amidinohydrolase allantoïne allantoate uréidoglycolate glyoxylate CO 2 + 2NH 3 CO 2 + 2NH 3 CO(NH 2 2 uréaseallantoïnase

Allantoate

amidohydrolase uréidoglycolate amidohydrolaseUréidoglycolate urée-lyase CO 2 + 2NH 3 CO 2 + 2NH 3 CO(NH 2 2 uréase

L...TP...-OITN VRDIEVIBDN1-P...DBIVBIEVI-B...L>N'VIEVI.CN,...BVILMV,IEVI1...TSDVR-V-IV-PXLV-I:O'OBN.V-ÉI1CRDOVI-CNLLRTR.VINRIL...RD-IEVI.NI'VDTi1NBi...1IVBIERIEO:V....PPVTV1BIEV-IP.N1BR.V-@I

DO-VD:V-IPD...BOi%RV-IEVI.NI'DNi1VI(Y...1iNIVBIN.É@I*LLMBI;....EDNi IRI5....NLL...@I*LLL.ÉI1CRDOVI

.V-IVTSD&...1-IEVI-... NIV1IEO:V....PPVTV1BI(2iLN..V2IRI4MV.P@I*L-L.@INi1-iI%RCR1VI2NiS.VINLBi:iBOIquotesdbs_dbs26.pdfusesText_32

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