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*+A$D5CA76AC57AB3""965287AC33B57A52A697525 ********************************************************************************************** 14A 6+2+21$CB1 9D 1 CB1CB1#9 BECB +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 7?1 6+2+61$CB1 CB1CB1E9CB1!C!9DCB +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 7@1 6+2+71$CB1 CB1DCE9CB<1CB1 CB1C19ABC9FCB +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 7@1 *A5AE98DC37E5AC33B3(5****************************************************************************************************************************** 08A *1ADAE73832AC33B3(5AB57AE36DC57A5D6,857 ****************************************************************************** 04A 6+7+213CDC91CD1 CB1EE ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 9:1 6+7+61$CB1&&A9CDECB1BBCB1C1 CB+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 9:1 6+7+71$CB1CB19B ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 9?1 6+7+91=9EDB1DEC9DCB +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 9@1

3. L’INFLUENCE DU CYCLE CELLULAIRE ET DU CYCLE NYCTHEMERAL SUR LA COMPOSITION LIPIDIQUE ... 48

1*+A)2"C525ABA,C5A5CCCD365 ************************************************************************************************************************* 03A

7+2+210DE9JC11(C1C9C1 9119BBDC +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ 9F1 7+2+61$1! BDEC1 )C1C119BBDC1B!E)C ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ :21

1*A)2"C525ABA,C5A2,8 9E96DCAF<6I238************************************************************************************************* 4A

4. CONDITIONS ENVIRONNEMENTALES ET PRODUCTIONS QUALITATIVE ET QUANTITATIVE DE LIPIDES ... 52 0*+*AE9296DC3897 **************************************************************************************************************************************************** 41A 0*ABD8D832AADC3ED8D832A/ ********************************************************************************************************************* 40A 0*1A)2"C525AB5ACDA25286D832A52A2386D85ABAE3C35************************************************************************************* 40A 9+7+21$19CDC1CD1'EC1 9F)C1D199;E1C11FBD1C9C ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ :91 9+7+61$19CDC1CDE9LDC1DC1!ED1C1E9(A9CB1CE1C1B9CB +++++++++++++++++++++++++++++++++++++++++++++++++++++++ :@1 9+7+71$19CDC1CD1'EC1 9F)C1DC1!DED1C1EFEA1 #EB(DE#AE)C ++++++++++++++++++++++++++++++++++++++++ :F1 9+7+91$19CDC1CD1'EC1!&C11! BED1CD1C19B1CB1(A9 CB ++++++++++++++++++++++++++++++++++++++++ ?I1

0*0A)2"C525AB5ACDACE3.65********************************************************************************************************************************** 5+A 9+9+21&ED1CB1BBCB1C1 CB +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ ?61 9+9+61&ED1C1DBE9ED1C1CB1 CB+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ ?61 0*4A)2"C525AB5ACDA85E96D865 ************************************************************************************************************************* 51A 5. CONCLUSION ... 64 1

1 791

1 7:1

1. Introduction

0CEEC1 9EC1 &9DE1 DC1 CB9 ED1 B(D E)C1 !B1 DD1 C#BEFC1 1 9JC1 8A1 91 CB1 &&A9CDECB1 BBCB1 C1  CB1 DB1 C1 !AEB!C1 C9C1 1 #(E DED+1 DFC9BC!CDE<1 DB1 FDB1 9C#C9#A1 !!CDE1 CB1 &&A9CDECB1 BBCB1 FCDE1 AFC91 CD1 &DED1 CB1 DE9DECB11!C<1DE!!CDE1DECDBEA1!DCBC<11EC! A9E9C1CE11B DEA1CB1 DE9!CDEB+1$1EEA9E9C1B1C1B8CE1AEDE1C 1 B1DDEC1B91CB1 DECB1B A9C9CB<1 D1C9ED1D!9C1C1DB1EEDB1(1&E19A&A9CDC+11

2. Les lipides du phytoplancton

5CBE1 C1 )D1  C1 K1 $CB1  CB1 BDE1 CB1 BBEDCB1 #(9 #CB1 ,1 ! # E)CB-1 BCB1DB1CB1BFDEB19D)CB+10CBE1BFCDE11 9ABCDC1D1! BDE1 9EC9<1 C1 9B<1 )1 DDC1 BCB1 9 9AEAB1 #(9 #EA1 1  CB+1 1 CBEC1 BC9B1 EA9CB1C1 CB1)1CCB!;!CB1CDCDE1C1D!9C1! BDEB11BE9E9CB1E9.B1 &&A9CDECB+1 "D1 CE1 AC91 BC9B1 BBCB1 C1  CB1 %1 CB1 CB1 9B<1 CB1 (A9 #B # CB1 , #B # CB-<1 CB1 (A9( CB1 ,1 ( CB-<1 CB1 B #D CB<1CB1BEA9B<1CB1 9ADB1CE1CB1((A9B1,DE1CB1E9(A9CB-+1DB1CEEC1 AEC<1 DB1 DB1 DEA9CBBC9DB1 CBBCDECC!CDE1 1 (A9 CB1 , #B # CB<1 ( CB1 CE1 E9(A9CB-1 )<1 !!C1 C91 D!1 D)C<1 BDE1 DBEEAB1 D1 B)CCEEC1 (A91B91C)C1BDE1CBEA9&AB1CB1CB19B+11

2.1 Rappel sur les différents lipides en présence

EC,C33B57A 58A  7 C33B571 BDE1 CB1 ! BDEB1 BE9E91 CB1 !C!9DCB+1 B1 AEBBCDE1 911CC1CE1 91CB19DECB1DC199.9C1)1CBE1B19DC1C11! CEA1 C11FC+1B1BDE1BB1! )AB1DB1C1D!9C1!ADB!CB1C9CB1CD1 99DE1DC1 !E9C1 91 C1 D!9C1 9CBBB1 E(E)CB<1 CD1 D&CDADE1 EFC!CDE1 CB1 9 9AEAB1 &DEDDCCB1CB1 9EADCB1BBACB11!C!9DCB+1CB1A9FAB1C1CB1 CB1BDE1BB1

1 7?1

CB1!CBBC9B1BCD9CB1DE9C9CB1C1 9C!.9C1! 9EDC1, CB1E&B1%187<18<1 2<1 CE+-+1 $CB1 863C,963B571 BDE1 CB1 !ACB1 C1 9ABC9FC1 )1 CFCDE1 ;E9C1 EBACB1 91&9D91C1ADC9C1CE11 F919AEC911CCB+111

2.1.1 Les principaux lipides des chloroplastes

$CB1 ( CB11 #(E DED1BDE1BAB1 DB1CB1!C!9DCB1CB1#9 BECB+1 $CB1 C1 ( CB1 9D 1 BDE1 C1 !DEB(((A91 ,22-1 CE1 C1 EB(((A91 ,22-1 ,>9C1 2-+1 $C91 BE9E9C1 9E.9C1 C91 D&.9C1 CB1 9 9AEAB1B A&)CB+1$C1221CBE1D1 C1)11ECDDC1B1&9!C91CB1BE9E9CB1C1E( C1 #CD1 <1 9B1 )C1 C1 221 &9!C1 EJE1 CB1 BE9E9CB1 C1 E( C1 #C1  )C1 ,1#CE9EF9A16II@-+1$C1CD1CDE9C11BE9E9C1CB1 CB1CE1C9B1&DEDB1DB11!C!9DC1 CE1DB11CC1BC91BEA1 B1D1,F91BB13C12-+10C1CD1 CBE1DC1! 9EDC1 9C1 91! 9CD9C1CB1DBA)CDCB1CB1!&EDB11 9&1 )C1 9F)ACB1 91 9ED1C1DFCCB1DEDB1CDF9DDC!CDECB+1

$CB1 #9 BECB1 BDE1 BB1 ! BAB1 C1 B&)DFB(((A91 ,652-1 ,>9C1 2-1 )11ECDDC1B1&9!C91CB1#CB+1

1

1 7@1 A365A+A-ADA786865AB57AC,C33B57AB57AA C6CD7857*A57ADCD8C33B57AE=EA58A=E=EA 25A728AD7A D697*A5A7C"C33B5A%9=EA578A D69A29D8345E528*A=D6.7A:D6JBAF+775*A

$CB1 #B # CB1 9A DA9DEB1 DB1 CB1 !C!9DCB1 CB1 #9 BECB1 BDE1 C1 #B #E((A91 ,82-<1 CE1 1 #B #E(#DC1 ,80-1 ,CDFC C1 CEC9DC1 CBFC!CDE-+1 $C1 821 , C1 DD)C-1 DB1 )C1 1 801 DE1 BB1 ECDDC1 B1 &9!C91 CB1 #CB+1

2.1.2 Les lipides des autres membranes

$CB1 #B # CB1 BDE1 CB1 ! BDEB1 9D 1 C1 EECB1 CB1 E9CB1 !C!9DCB1 C1 1 CC+1 7D1 B1 C1 1 801 CE1 1 82<1 D1 E9FC1 9D C!CDE1 1 #B #E(AE#D!DC1 ,87-<1  C1 DCE9C1 ,'MEC9D)C-1 )1 B9DBC1 9D C!CDE1 CD1 BE9E9C1 9BEDC1 #CDC1<1CE11 #B #E(BC9DC1,86-1,#C-+1

0CB1 &&A9CDEB1  CB1 DE1 CB1 &9!CB1 !A9CB1 FC9BCB1 ,(D9)C<1 D)C<1 D)C1 DFC9BAC-1)1AEC9!DCDE11BE9E9C1CB1!C!9DCB+1$1! BED1C1CB1!C!9DCB1CD1  CB<1 !B1 BB1 CD1 9EADCB<1 CBE1 CBBCDECC1 91 BB9C91 C9B1 &DEDB1 ,&EA<1 C9!AEA<1 DBC9ED1 CB1 9EADCB1 !C!9D9CB<1 CE+-+1 NDC1 !&ED1 C1 1 9A 9EED1 CDE9C1CB1&&A9CDEB1! BDEB1CDE9LDC91DC1!&ED1CB1 9 9AEAB1CB1!C!9DCB+11

2.1.3 Les lipides neutres, des lipides de réserves

$CB1 E9(A9CB1 ,1 E9((A9B<1 32-1 BDE1 CB1 9D 1  CB1 C1 9ABC9FC1 1 #(E DED+1 B1 BDE1 BE/AB1 BB1 &9!C1 C1 CEECB1 EECCEECB1 DB1 C1 (EB1 CE1 BDE1 9&B1BC9FAB1DB1CB1#9 BECB1, BE-1C11CC1,$1D1$D16II2-1,8#E1 2-+1

1 7E1

1

> 8A +A-A '5CCC5A B1234567282A 7*A ED6(95A DA #3C5A 65BA 58A 756495A DA E3675A 2"DCA BA !*A 5A #3C5A 65BA "C6575A BD27A C5A <D25A BD27A C57A 52436225E5287A C57A C7A DCD3657A F863C,963B57A58ABD27AC5A65ABD27AC57A52436225E5287AC7ACD3657AFE5E6D257*A

0!!C1 91C1D!9C19DB!CB<1CB1E9(A9CB1CB1!9CB1C9(ECB1BDE1CB1 9ABC9FCB1C19DC1CE1ADC9C1)1 CFCDE1;E9C1!BACB1)D119BBDC1CBE1BE!AC1 ,9 #(1 6II2-+1 $CB1 321 BDE1 9B1 W(AB1 91 99C1 C1 381 91 9CB 9ED<1 DB1 )C1CB1!ACB19AE9CB1)1BDE1EBACB1 911B(DE#.BC1C1! BAB1DACBB9CB1B11 9BBDC1 , 9EADCB<1 B9CB<1 (A9 CB1 91 B(DE#.BC1 E9 33<1 CE+-+1 891 CC! C<1 (ED1 ! .EC1 DC1 !AC1 ECC1 )C1 C1 !EEC1 ,02?-1 AD.9C1 2I?1 !ACB1 381 ,4C99 EF9 A1 6II7-+1 0C1 BDE1 CB1 9ABC9FCB1 ADC9AE)CB1 CE9;!C!CDE1 CDBCB1 ,B1 BDE1 9AEB1CE1D#(9CB-+1ND19!!C1C1E9(A9C1DECDE1CDF9D1?1&B1 B1ADC9C1)D1 9!!C1C1(.DC1#(9EA1, (B#9C1)1BC9E1BB1C1BE/C-+1

C1 9ACDECB1 AECB1 DE1 !DE9A1 )C1 CB1 E9(A9CB1 DAECDE1 B1 BCC!CDE1 CB1 9EB1 &D1C1DB!C1 )C1!B1)*B1 FCDE1BB1DBEEC91CB1DEC9!A9CB1DB1 1 B(DE#.BC1 CB1 ( CB1 CE1 #B # CB1 !C!9D9CB1 ,U#'D2C99 EF9 A1 6III-+1 CB1 C A9CDCB1 C1 !9)C1 12901 B91B36576889 46EF1 DE1 !DE9A1 )C1 CB1 CB1 9B1 AECDE1 91 DEA9AB1 1 321 91 ;E9C1 CDBEC1 BE9AB1 FC9B1 CB1  CB1 9CB1CB1E9CB1! 9E!CDEB1C9CB+1DB1C9EDCB1DEDB1C19BBDC1,9C 9BC1 C11FBD1C9C1 9.B1DC1 #BC1C19CDC1DE9EFC<1 91CC! C-1CE1 91C9EDCB1 CB .CB1C1 #(E DED<1CB1(A9 CB1!C!9D9CB1 CFCDE1;E9C1CD1 9EC1BBC!AB1B1 9E91 CB1 CB1 9B1 CB1 32+1 0C9EDB1 CB1 9B1 ! 9EDEB1 &DEDDCC!CDE1 ,C9EDB1 8N>-1 BE/AB1 DB1 C1 &9ECB1 9 9EDB1 DB1 CB1 321 DC1 BDE1 B1 A9AB1 9B1 C1 1

1 7F1 9C 9BC1 C1 1 9BBDC<1 !B1 9AEBAB1 91 &9)C91 C1 DFCCB1 !C!9DCB1 ,U#'D 2C99EF9A16II:-+11

$CB1 !9CB1 !9DCB1 AFCDE1 DB1 D1 CDF9DDC!CDE1 F9C1 DE1 CB1 !E CB1 !&EDB1B19E1EC9!C1,BDEA<1EC! A9E9C<1B DEA1CD1DE9!CDEB1CE1CD1 #EDB-1 CFCDE1 D9C1 DC1  EED1 CE1 DC1 9A9DBED1 CB1 !C!9DCB+1 $CB1 EECCEECB1  )CB1C1321BDE1F9BC!C!CDE1CB1B(BE.!CB1E! DB1DEC9DCB1)1 CFCDE1&9D91 9 C!CDE1 CB1 9 CB1 (CB1 B A&)CB1 ,78<1 4-1 BBC ECB1 C1 !&C91 CEH1 C1 BEBC91CB1!C!9DCB1DB1D1CDF9DDC!CDE1F9C+11 1 9 9 9 A A DC5DA+A-ADC37D832A58AA8,57AB5AE5E6D257A"6E957AD6AC57A6323DAC,96C33B57A 33B5A ,5AB5AE5E6D25A"6E95A DC37D832ABD27ACDA5CCC5A

221 4CD11 3#(/RCB1 221 4#C1 3#(/RCB1 6521 4#C1 3#(/RCB1 821 4#C1 0#9 BECB1 801 4#C1 3ECB1CB1!C!9DCB1 871 4#C1 3ECB1!C!9DCB1B&1#9 BECB1 861 4CD11 3ECB1!C!9DCB1B&1#9 BECB1

1 9I1

2.2 Le métabolisme lipidique

$1 ! BED1 CB1 &&A9CDECB1 !C!9DCB1 C9CB1 &&.9C1 BD&EFC!CDE1 BFDE1 C91 &DED1 ,CDFC C1 CB1 #9 BECB<1 E#(/RCB<1 9AE!1 CD B!)C-+1 $CB1  CB1 BDE1 B(DE#AEBAB1 DB1 FC9B1 ! 9E!CDEB1 C1 1 CC+1 $1 CC1 B BC1 C1 !ADB!CB1 C9!CEEDE1D1E9DB 9E1CB1&&A9CDECB1BBCB1C1 CB1)19AC1C91! BED11BCD1 CB1 &&A9CDEB1 ! 9E!CDEB+1 $CB1 FCB1 C1 B(DE#.BC1 CB1 &&A9CDEB1  CB1 BDE1 D1 #EC!CDE1DEA9ACB+1

$CB1  CB1 !C!9D9CB1 BDE1 CBBCDECC!CDE1 CB1 (A9 CB1 ! BAB1 D1 B)CCEEC1 E99DA1,(A9-1B91C)C1BDE1CBEA9&AB1C1CB19B1CD1 BED1BD21CE1BD61CE1DC1 E;EC1 9C1CD1 BED1BD71,>9C16-+11

1

A365AA-A% 9EDAAB2AC,96C33B5A

0#C'1 CB1 DECB1 B A9C9CB<1 CB1 CB1 9B1 BDE1 B(DE#AEBAB1 DB1 CB1 #9 BECB1 BB1 1 &9!C1 (081 91 D1 ! CC1 CD'(!E)C1  CA1 >61 ,>EE(1 1 6(DE#BC-+1 $CB1 9EB1&D1BDE1 9D C!CDE1C12?%I081CE1C12E%I08+1$C12E%I1CBE1ABE9A1CD12E1 %208+17DBEC12?%I1CE12E%21 CFCDE1;E9C1CBEA9&AB11(A97 #B #EC1 91&9!C91C1 C1 #B #E)C1 ,8-1 1 ;E9C1 #(9(BAB1 91 99C1 CB1 CB1 9B1 9CB1 ,>9C1 7-+10CB1C9DC9B1BDE1CBEA9&AB11CD'(!C11,0-<1 B1E9DB 9EAB1DB1C1(E B!C11 B1 CFCDE1 ;E9C1 CBEA9&AB1 1 81 ,U9 EF9 A1 6II9-+1 $1 >9C1 71 9ABCDEC1 D1 9AB!A1 C1 1 B(DE#.BC1CB1(A9 CB1FC1CB1&&A9CDECB1CD'(!CB1! )ACB+1

1 921 1

A365A1A-A57A69D8327AB5A86D27"568ABD,CABD27AC57A C6CD7857A58ABD27AC5A,8CD7E5AB57A CD2857*AF+AC,96C1 7 D85AD,C86D27"96D75&AE>AFAFA+D,CC,56C1 7 D85A D,C86D27"96D75AFAF1A 7 D83BD85A 7 D8D75AFAF0A 7 D83BD85A,83B,C86D27"56D75*A=D6.7A 6D8DA58AD7D@DAF+772*A

$CB1 (A9 CB1 B(DE#AEBAB1 DB1 C1 (E B!C1 ,(A9 CB1 C1 BE9E9C1 C9(EC-1 CBEA9&CDE1D1 C1 9B1 CD102E1CD1 BED1BD61 9B1)C1 C1CB1#9 BECB1, CB1 C1 99(ECB1D-1CBEA9&CDE1CB1CB19B1CD102?1,>9C19-+17D1 BED1BD2<1CB1CB19B1 CD102?1CE1CD102E1BDE1 9ABCDEB1)C1)C1BE1C91BEC1C1B(DE#.BC+10CEEC1BE9ED1B.9C1 )C1CB12831CB1#9 BECB1CE11(E B!C1DE1DC1&C1B A&EA1FC1C91BBE9E+1 1 DE99C<1 1 2((C97 #B #EC1 (E9DB&A9BC1 1 (E B!C1 9CDDLE1 B A&)C!CDE1 CB1 CB1 9B1 CD1 02E1 9B1 )C1 CC1 CB1 #9 BECB1 9CDDLE1 B A&)C!CDE1CB1CB19B1CD102?1,9E1D13B/12FF@-+1

1 961 1

1

A365A0A-A33B5AB5A786865A6D6,85A7,28 98379ABD27AC57A C6CD7857AFDAD 5A58AB5A 786865A5D6,85A7,28 98379ABD27AC5A,8CD7E5AFDAB6385A

1

$CB1 9D 1 #B # CB1 E9FAB1 DB1 CB1 EBBB1 FAAE1 DE1 DC1 BE9E9C1 E( C1 C9(EC<1 CC EA1 91 C1 821 BE+1 $1 B(DE#.BC1 CB1 #B # CB1 CBE1 !!DA!CDE1 BA 9AC1 CD1 71 FCB1 %1 CB1 #B # CB1 A9FAB1 1 (EDC1  #B #EC1 ,08-2<1 C1 A9FAB1 1 2<1 CE1 C1 9FCDDE1 A#DC1 C1 E;ECB1 9CB1  9ECDDE1 B1 E9CB1 #B # CB1 ,1#CE9 EF9 A1 6II@-+1 $C1 2<1 C1 BE9E9C1 C9(EC1 1 99(EC<1 CBE1 C1 9A9BC91 C1 1 B(DE#.BC1 CB1 ( CB1 ,652<1 221 CE1 22-+1 1 CBEC1 D1 C1 E( CB1 C1 ( CB1 %1 C1 E( C1 99(EC1 DE1 C1 B)CCEEC1 21 9FCDE1 CB1 BECB<1 CE1 C1 E( C1 C9(EC1 DE1 C1 21 CBE1 BB1 C1 1 80+1 $1 B(DE#.BC1 CB1 ( CB<1 AEDE1 BAC1 DB1 CB1 !C!9DCB1 C1 CDFC C1 CB1 BECB<1 9C)C9E1 D1 D1 !ADB!C1 ! 9E1 1 21C1E( C1C9(EC+1

$1 221 B(DE#BC1 E9DB&.9C1 D1 EBC1 B91 D1 2+1 1 CBEC1 BC9B1 #!CB1 C1 CEEC1CD'(!C1)1 9ABCDECDE1CB1&&DEAB1&&A9CDECB1 91CB1BBE9EB1,9DC1C9(EC11 99(EC-1 ,M9 EF9 A1 6II2-<1 CE1 )1 BDE1 EFAB1 &&A9C!!CDE1 CD1 &DED1 CB1 DEDB1 CDF9DDC!CDECB+1 0CBE1 DC1 CB1 FCB1 C1 9A DBC1 C1 1 ! BED1 CB1 !C!9DCB1 1 F9EDB1 C1 DEDB1 CDF9DDC!CDECB+1 1 CBEC1 BB1 &&A9CDEB1 #!CB1 C1 1 221 B(DE#BC<1 )1 8ECDE1 D1 EBC1 1 22+1 0CB1 #!CB1 BDE1 AC!CDE1

02E1102?1 02?1 X1 02E1102?1 02E1 X1

1 971 B A&)CB1 D1 BBE9E<1 CE1 DEC9FCDDCDE1 DB1 1 9AED1 C1 1 ! BED1 CB1 !C!9DCB+1

$(ED1CB1((A9B1,2-1 91D1(101 911((A91(E9DB&A9BC1 ,23-1 AEE1 8B)B1 9AC!!CDE1 DBA9AC1 !!C1 1 BCC1 FC1 C1 B(DE#.BC1 C1 E9((A9B1 ,32-+1 C1 DFCCB1 CD'(!CB1 C1 1 B(DE#.BC1 CB1 321 )1 E(BCDE1 1 E9DB(ED1C19 CB1(CB1CB1 #B # CB1CE1CB121FC9B1CB1321DE1AEA1A9ECB1 ,6E9E9EF9A12FF@<16E#1D1+16II9-<1CE1BC!CDE1DEC9FCD91C1!D.9C1E9.B1BD&EFC1 DB11B(DE#.BC1C132+1$CB1321BDE1CBBCDECC!CDE1CB1BE9E9CB11E( C1C9(EC+1

1 991 1 DC5DAA-A5256A52AC33B5AB5A+8A57.57AB5A ,8CD282*A H7.5A 33B5AFKA 3B7A75A $9"965257A

C4EEE221B +1,0#9 #(CC-1 269I1 ,6#CC#D9EF9A12FFE-1 5A733A21B +1 ,0#9 #(CC-1 6I1 ,6#CC#D9EF9A12FFE-1 536EA1B +11 ,0#9 #(CC-1 6E761 ,0#BE16II@-1 E345368293E3AA21 ,0#9 #(CC-1 7::91 ,0#BE16II@-1 A8EA1BD1,0#9 #(CC-1 29+91 ,4DEC(1D13C8C1 6II@-1 B67E281B +1 ,4 E #(CC-1 667E1 ,6#CC#D9EF9A12FFE-1 B36576881B +1,3# #(CC-1 F291 ,6#CC#D9EF9A12FFE-1 B5AE3A4F79F68436F9 ,49 #(CC-1 6I7I1 ,0#BE16II@-1 C86376A1B +1 ,0#9 #(CC-1 22621 ,6#CC#D9EF9A12FFE-1 12345672821,E#ED-1,EB-1 ,89(!DCB #(CC-1 6E+:1 ,6#CC#D9EF9A12FFE-1

1 9:1

2.3 La composition lipidique des micro-algues eucaryotes

$1 ! BED1  )C1 CB1 CB1 C9(ECB1 CBE1 9ADFDE1 CD1 DDC1 #C'1 D1 9D1 D!9C1CB .CB+10CB1AECB1DE1!!CDA11(11CDF9D1:I1DB19TC11AFC C!CDE1C1 &&A9CDECB1EC#D)CB<1CE1DE!!CDE1C11#9!E9 #C1'CBC1)1DB119CDBCDAB1B91 1! BED1CD1CB19B1CB1&&A9CDEB1 CB+10CB1AECB1DE1BFCDE1AEA1!EFACB1 91 ! 9EDC1 CB1 9DB!CB1 #(E DED)CB1 CD1 )E9C+1 1 BBBE1 C1 AEC9!DC91 C9B1 )EAB1 DE9EFCB1 DB1 C1 E1 C1 BACEDDC91 CB1 B1  9 9ACB1 91 #)C1 E( C1 ACFC+1"D1BCBE1DE!!CDE1DEA9CBBA1B11ECDC91CD1E9(A9CB1,#EC1FC91ADC9AE)C-1 CE1CD1C19B1 (DBE9AB1,CB19B1CBBCDECB-+11

2.3.1 Teneur en lipides totaux

$1 ECDC91  )C1 CBE1 DDC1 91 D1 9D1 D!9C1 CB .CB+1 $CB1  CB1 DBEECDE1 CDF9D1 2I1 B1 9IY1 1 B1 BC1 DC1 C1 CD1 #BC1 C1 9BBDC+1 "D1 BC9FC1 D1 CB1 &&A9CDCB1DEC9B A&)CB1! 9EDECB1,3C16-+1

9A1C19D1D!9C1AECB1DECB1DB1C1!DC<1BC1D1E9.B1&C1 9CDEC1 CB1 !C9B1 *CB .CB1 DDCB1 1 AEA1 D(BA+1 0!!C1 DB1 C1 FC99DB1 ,F91 0# E9C1 :-<1 1 ECDC91 CD1  C1 CB1 !9CB1 A CD1 C1 C9B1 DEDB1 C1 9BBDC<1 C1 )1 9CD1 &&C11! 9BD1CDE9C1CB1AECB+1

2.3.2 Les différentes classes de lipides

$1 ECDC91 CD1 BBC1 C1  C1 CB1 !9CB1 CBE1 BBC'1 C1 DDC+1 2( CB1 CE1 #B # CB1 DE1 DC1 ECDC91 99AAC1 B1 1 EC1 C1 1 CC1 CE1 B1 DDC1 C1 BCB1 9DECB+1 0!!C1 DB1 C1 FC99DB1 CD1 7<1 1 ECDC91 CD1  C1 DCE9C1 A CD1 C 1 CB1 DEDB1CDF9DDC!CDECB+1$CB1&&A9CDECB1CB .CB1BC1BEDCDE1B9EE1 91C91 EA1 B1C!!BDC91CB1E9(A9CB1CD1 A9C1C1BE9CBB+11

1 9?1

2.3.3 Les acides gras

$1! BED1CD1CB19B11 #(E DED1A CD1C1CB .C1AEAC1CE<1!!C1 91 CB1 BBCB1 C1  CB<1 CB1 DEDB1 CDF9DDC!CDECB+1 $CB1 B CE9CB1 C1 9B1 C1 D!9CBCB1CB .CB1BDE1DDB1,UECB12F@I<13#! BD9EF9A12FFI<1=B1D19E(12FF7-+1 0C9EDCB1 BBCB1 1 #(E DED1 DE1 CB1 ! BEDB1 E9.B1 9EA9BE)CB+1 891 CC! C<1 CB1 E!ACB1 ,49 #(CC-1 BDE1 9#CB1 CD1 2?%2O@<1 2?%I<1 6I%:O71 CE1 2?8N>1 ,"9EE1 2F@7<1 " EC1 2F@9<1 ('9 EF9 A1 2FEF<1 =B1 D1 9E(1 2FF7-+1 $CB1 D #(ACB1 BDE1 BFCDE1 9EA9BACB1 91 C1 &9EB1 E1 C1 2?%I<1 66%?O7<1 6I%:O71 CE1 2E%9O71 ,DB99 EF9 A1 2FFF-<1CE1 911 9ABCDC1C12E%:O71,1BC #12F@9<1('1D1/!D12F@?-+1 $CB1CB1 9B1 CB1 B1 !!DB1 #C'1 CB1 89(!DAB #(ACB1 ,4 E #(CC-1 BDE1 C1 BE9AB1 %1 29%I<1 2?%I<1 2E%2OF+1 $CB1 #9 #(ACB1 BB.CDE1 !89E9C!CDE1 1 2?%I<1 2E%7O71 CE1 2?%7O9+1 $BCDC1 C1 6I%:O71 CE1 C1 66%?O71 DB1 CEEC1 BBC1 C1 #(E DED1 1 AEA1 EBAC1 !!C1 !9)C91 B A&)C<1 91 C1 #(E DED1 !9D1 CBE1 8BEC!CDE1 DD1 91 F91 CB1 E1 ACFAB1*CB19B1 (DBE9AB1CBBCDECB+11

"D1 E9FC1 C1 &9EB1 E1 C1 (DBE9AB1 CBEA9&AB1 B91 CB1 ( CB1 91 1  9E1 CB1 !9CB1 C9(ECB1 ,U#'D2C99 EF9 A1 6II6-+1 $CB1 ( CB<1 DBEEDEB1 CB1 !C!9DCB1 CB1 #9 BECB<1 DE1 D1 9JC1 BE9E9C1 CE1 &DEDDC1 ! 9EDE1 DB1 CB1 #EB(BE.!CB1 CE1 CB1 9AEDB1 #EB(DE#AE)CB1 ,Z9!DD1 D1 4CDDD1 6II6<1 2BB1 D1 S#C!1 6I2I-+1 $CB1 CB1 9B1 (DBE9AB<1 )1 D&.9CDE1 B1 1 !C!9DC1 B1 &EA<1 BC!CDE1AC!CDE1;E9C1DC1! 9EDC19C1 91C1D1&DEDDC!CDE1CB1CD'(!CB1 ! )ACB1 DB1 1 #EB(DE#.BC1 ,1DCB1 6II@-+1 $CB1 ( CB1 BDE1 CB1  CB1 )1 DE1 C1 E1 DBE9ED1 C1 B1 ! 9EDE+1 $CB1  CB1 DCE9CB1 ,32-1 BDE1 ADA9C!CDE1 C1 DBE9AB1 !;!C1 B1 C9EDCB1 CB .CB1 9E.9CB1 DECDDCDE1 CB1 321 E9.B1 9#CB1 CD1 CB1 9B1 (DBE9AB1,U#'D2C99EF9A16III<1U#'D2C99EF9A16II:-+1

1 9@1 7D1 B1CB1&&A9CDCB1DEC9B A&)CB<11&E1DEC91C1&9ECB1&&A9CDCB1DE9B A&)CB+1"D1 BC9FC1 CD1 C&&CE1 CB1 &&A9CDCB1 ! 9EDECB1 C1 ! BED1  )C1 *DC1 AEC1 B1 *E9C+1 0!!C1 DB1 C1 FC99DB1 B1 D<1 CB1 DEDB1 C A9!CDECB1 , CDDE1 !B1 BB1 FDE1 C A9CDC1 %1 C1 #BE9C1 D1 C1 1 E9C-1 DE1 DC1 D&CDC1 !8C9C1 B91 1 ! BED1  )C1DC1CC1C1 #(E DED+1

2.3.4 Variations interclonales

1

4CD1)C1#)C1CB .C1C1 #(E DED1 BB.C1DC1! BED1 )C1 9E.9C<111 AEA1!DE9A1)C1CB1&&A9CDCB1! 9EDECB1CBECDE1CDE9C1CB1B#CB1DC1!;!C1CB .C<1 &&A9CDCB1 )C1 D1 CE1 EE9C91 B1 CB1 FC9B&EDB1 ADAE)CB+1 "D1 1 91 CC! C1 1 !DE9C91 )1 CBEE1 DC1 9DC1 F9EA1 #ADE( )C<1 DE!!CDE1 91 C1 E1 C1 9BBDC<11)DEEA1C1 C111! BED1CD1C19B+11 9E91DC1!;!C1B#C1 C1!.9C1D<1$D1EF9A1,6II:-1DE1!B1CD1E9C1?I1DCB1,BAB1B1 9E9121CC1D)C-1 DB1CB1DEDB1CDE)CB[1CB1&&A9CDECB1  EDB1BBCB1C1CB1DCB1BC1BDE1FA9ACB1 F91 CB1 ! BEDB1  )CB1 CE1 CB1 E1 C1 9BBDC1 E9.B1 &&A9CDEB+1 891 C9B<1 CB1 !CB9CB1 C1  CB1 )C1 DB1 FDB1 1 &9C1 1 (E!.E9C1 CD1 &1 B91 CB1  EDB1 C1 CCB1!9)ACB11GC19C1!DE9CDE119DC1F9EA1CDE9C1CB1DFB1DC1!;!C1  ED1C1,>9C1:-+10CEEC1F9EA<1! EC1B1CB1&&A9CDCB1ADAE)CB<1 CE1 BB1;E9C1C1 B1CB1&&A9CDCB1C1 BEDDC!CDE1CB19DB!CB1DB1C91(C1C9C1 ,F916+-+1$1 BEEA1 #ADE( )C1CB1CB1, EA1D1ADE( C1B1 99C1&&A9CDEB1 #ADE( CB-1C )C1BDB1EC1DC19DC1 9E1C11F9EA1CB19ABEEB1C11EEA9E9C+1

1 9E1

1

A365A 4A -A $9D683832A B25A A CD832A B5A 5CCC57A B12345672829 7CA 52A "2832A B5A CDA 25286D832A 3286D5CCCD365A 52A C33B57A FA&A 328527389A 65CD8345A B5A "C657525*A DA CD832A B5A5CCC57AD2DC,795A98D38A7E375A?A2A9CD365E528A2832A58A298D38AD7A7,2 623795*AD7525A B5A7,2 6237D832A75AD6D896375AD6AB57A7383225E5287ABD27AC5A,C5A5CCCD365AB3""965287A6A  D(5A5CCC5*AAA

3. L’influence du cycle cellulaire et du cycle nycthéméral

sur la composition lipidique

3.1 Influence du cycle cellulaire

5A ,C5A 5CCCD3651 CBE1 *CDBC!C1 CB1 #BCB1 91 CB)CCB1 DC1 CC1 BBC1 CDE9C1 C1 FBDB1 BCBBFCB+1 $1 FEA1 DC1  ED1 C1 CCB1 A CD1 1 C9A1 C1 9DED1 CDE9C1 9BBDC1 B!E)C1 CE1 FBD1 C1 BCB1 DFB1 ,3 D9 EF9 A1 6II2-+1 $D&CDC1 1 (C1 C9C1 B91 1 B(DE#.BC1 C1 CB1 CB1 !9CB1 1 AEA1 C1 AEAC+1 $CB1 EC#D)CB1 C1 E9CB1 B(D#9DBACB<1 DB1 )C1 CB1 !AE#CB1 C1 (E!AE9C1 CD1 &<1 C9!CEECDE1 AB9!B1 AEC91 C1 9!.E9C+1 $CB1 CF9CB1 BDE1 C B1 DEC! B1 CB1 9DB!CB1 !.CB1 91 AEC1 1 (C1 C9C<1 CE1 C1 BCB1 9AEDB+1 $1  9E1 CB1 D&9!EDB1B1C1B8CE1 9FCDDCDE1D1C1AEC1C1CB19DB!CB<1!B11BC!C1)C1CB1

1 9F1 !ADB!CB1!A9CB1AB11(C1 C9C1BCDE1E9.B1CD1DBC9FAB1CE1E9.B1B!9CB1 DB1EB1CB19DB!CB1C9(ECB1,=E12FF9-+11

3.1.1 Contrôle du cycle cellulaire par la croissance

$19BBDC1C11CC1CBE1C19ABEE1C11B(DE#.BC1C1!9!ACB1, 9EADCB<1 CB<1 (B#9CB<1 G-+1 0C9EDCB1 AECB1 B.9CDE1 )C1 CB1 E9DBEDB1 CDE9C1 CB1 #BCB1 1 (C1 C9C1 ,22H6<1 26H<1 CE+-1 BDE1 9AACB1 91 1 9BBDC1 B1 E9FC9B1 1 ! BED1 #!)C1CB1CCB1,3 D9EF9A16II2-+1ND1 DE1C1DE9JC1DD1CBE1CDE9AC1CD1 #BC1 61 ,9A ED1 C1 G-1 )1 BC9E1 DE9JAC1 91 1 )DEEA1 C1 9EADCB1 DE9C9CB1 1 91BD1E1C1B(DE#.BC1,C1BAC1D1 911DCDE9ED1CD1(DC1)1CBE1 9 9EDDCC1B11 ECDC91 EEC1 CD1 9EADC-1 ,8(!CDB1 D1 6#!E1 2FF@-+1 $1 ECDC91 CD1 B9C1 CE1 CD1  C1 99E1 F91 CB1 C&&CEB1 DCB1 B91 1 99CBBD1 1 (C1 C9C1 ,3 D9 EF9 A1 6II2-,>9C1?-+17D1C&&CE<1B1D1D#C11B(DE#.BC1C1 CB1CD122<1D1 9F)C1D199;E1C11 FBD1 C9C<1 9B1 )C1 B1 D1 D#C1 CD1 261 C1 (C1 C9C1 9BE1 B1 99CBBD1 ,UM/1 D1 SD1 6II:-+1 891 C9B<1 1 B(DE#.BC1 C1  CB1 BC1 9BE1 B1 *D1 )C1 1 FBD1 C9C1 ,CBE1 BC1 )1 BC1 BBC1 9B)C1 CB1 CCB1 BDE1 9CDACB1 CD1 'EC1 1 9B)D1 )C1 1 B(DE#.BC1 G-+1 1 1 AEA1 BA9A1 CBECDC1 D1 !ADB!C1 )1 AFC9E1 ,!!C1 91 CB1 9EADCB-1 1 )DEEA1 C1  CB<1 1 )DEEA1 D1 ! BDE1 B A&)C11CD9C11B(DE#.BC1C1 CB1CD1CC!;!C<1)1E1;E9C1B&&BDEC1 91CDE9C91 CD1 #BC16+1

1

1

1

1 :I1 1 9 1 9

A365A 5A-A B.C5A 79CD83"A B5A D2A DE9 FCA F88+A 6A 5C3(56A EE528A B3""965287A 783EC3AA 54528A D436A B57A 5""587A B3""965287A 76A C57A 86D2738327A BA ,C5A 5CCCD365*A =D27A 5A EB.C5&A C57A 783EC3A 52436225E528DA A B9856E32528A CDA E73832A 52A ED6EC9C57A F765&A C33B5&A 689325A (3A 5CC5EBE5A 286CC5A 568D3257A 86D2738327A BA ,C5A 5CCCD365*A =D27A 58A 55EC5&A C5A 783EC7AA"D46375ACDA6B832AB5A6893257ADC67A(5AC5A783EC7AGA"D46375ACDECD832AB5A C33B57*A '57A A 6"3C7A B5A 6377D25A "D4637528A C5657732A B5A B3""965287A 69CD8567A BA ,C5A 5CCCD365AF5A-ACDA,C325AH&A69CD8635AB5ACDA86D273832AE+I%AC5A78632&A69CD856AB5ACDA86D273832A EI&A5286DL2D28AB57AB3""965257AB5A6377D25AB5ACDACD832AB5A5CCC5*A=D6.7AD2ADE9FCA F88+*A

1 :21

3.1.2 La composante lipidique de la croissance somatique

$1ECDC91 )C1CE11EC1CB1CCB1!CDECDE119B1C1C919BBDC<1CE19B)C1 CCB1DE1!A1BBC'1C1!EA9C<1DE!!CDE1!C!9D9C<1CCB1 CFCDE1BC1FBC9+161 D1 BB!C1 1 CC1 B1 DC1 B #.9C<1 1 )DEEA1 C1  CB1 , 9CB-1 DBEEE&B1 C1 1 !C!9DC1CEC9DC1E1;E9C1E#A9)C!CDE1!E AC1 912<?1 91D1C!CDE1C1F!C+1 0#C'1 *CB .C1 C1 D&CA1 67F5E43889 43588<1 1 )DEEA1 C1  CB1 9CB1 CBE1 !E AC1 91 2<@<1 9B1 )C1 1 )DEEA1 C1  CB1 DCE9CB1 CBE1 !E AC1 91 2<E1 ,UM/1 D1 SD1 6II:-+1 0CEEC1 !;!C1 AEC1 1 !DE9A1 )C1 !CDEED1 CBE1 DEDC1 91 CB1  CB1 DCE9CB<1CE1BDEDC1 91CB1 CB1 9CB1,B(DE#.BC19AEC1CD1 #BC16-+9

1

8BC9B1 )CBEDB1 9CBECDE1 CD1 BB CDB1 %1 1 FBD1 C9C1 CDE9LDCECC1 BCC!CDE1 1 9EED1 1 1 C1 E9(A9CB1 CDE9C1 CB1 C1 CCB1 &CB<1 1 CD1 CBECC1 AC!CDE1 ! DAC1 *DC1 DB!!ED1 C1 E9(A9CB1 , 9E1 ADC9C1 EBC1 91 1 FBD-K1\E1CB1 #BCB1C1 9ED1C1 CB119B11(C1C9C1K10!!CDE1 CBE1 !&AC1 1 ! BED1 CB1 &&A9CDECB1 !C!9DCB1 C9CB1 1 9B1 1 (C1 C9C1K110CB1B CEB1DE1AEA1 C1E9EAB+1

1

$CB1 AECB1 9ABACB1 B91 C1 !AEB!C1 CB1  CB1 CD1 CD1 FC1 C1 (C1 C9C1 CB1 !9CB1 BDE1 C1 D!9CBCB1 CE1 9ECDE1 1 B91 CB1 E9CB1 B(D#9DBACB+1 NDC1 CB1 !AE#CB1 91 ECD91 C1 E( C1 C1 E9CB1 DBBEC1 B1 CB1 B!CEE9C1 B1 CB1 DEDB1 CDF9DDC!CDECB1 ()CB1 ,(C1 !DC1 89HDE-1 DE1 CB1 C&&CEB1 BDE1 &&C!CDE1 BBCB1 1 (C1 C9C1 CD1 !;!C+1 7D1 2FE2<1 6#&9D1 D1 0#B#!1 ECDECDE1 C1 BA 9C91 CB1 C&&CEB1 1 (C1 89HDE1 CE1 1 (C1 C9C+1 891 C1 B1 B(D#9DBCDE1 DC1 E9C1 91D1(C189HDE1 B11B!CEECDE1B1D19A!C1DED+1$CB1CCB19CBEDE1CD1 #BC1 CDDE1DC1ADA9ED<11CFCDE1 BBC1C1BF9C1AFED1C11DCDE9ED1CD1

1 :61

 CB1C1DA CD!!CDE1D11(C189HDE+1B1DA CDDC1CB1CB1F9EDB1FBB FB11(C189H1DE1DC1 9E1 B1A!DE9AC+1NDC1E9C1 BBEA1BC9E1C1B(D#9DBC911 E9C1 91 CB1 CDEB1 #!)CB1 ,D'-1 )1 DE1 C1 C1 #DC1 F91 D1 C&&CE1 B91 C1 !AEB!C1 CB1  CB<1 )C1 B<1 BCB1 CB1 C&&CEB1 D#A9CDEB1 1 (C1 C9C1 99CDE1 ;E9C1BC9FAB+1

3.2 Influence du cycle nycthéméral (jour/nuit)

"D1 CE1 ECD91 CB1 CCB1 C1 #(E DED1 B(D#9DCB1 CD1 CB1 B!CEEDE1 B1 CB1 (CB1 89HDE1 )1 9(E#!CDE1 B!EDA!CDE1 91 EECB1 CB1 CCB1 CB1 9CBBB1 A CDDE1 9CEC!CDE1CE1D9CEC!CDE1C11!.9C+17D9C1DC1&B<1 C1AECB1DE1E9EA1CE1B CE+1 6/CD/1D109!C1,2FFI-1DE1!DE9A1)CD1 B1C1DE9JC911FBD1C9C<1C1(C1 89HDE1 DE9JC1 1 9ED1 C1 9EADCB<1 C1 B9CB<1 CE1 C1  CB1 1 !;!C1 EE9C1 )C1 1 DCDE9ED1 CD1 !CDEB<1 1 BE9E9C1 1 EFEA1 C1  9C1 #EB(DE#AE)C+1 B1 !DE9CDE1 )C1 CDDE1 C1 89<1 B9CB<1 9EADCB1 CE1  CB1 BDE1 9EB+1 $DC1 CB1 CD'(!CB1 9D CB1C11B(DE#.BC1C19B1,CE(019(BC-1CBE1381A CDDEC1CE1BC9E1 &&CEAC1 91 CB1 &CB1 DCDE9EDB1 381 DE9C9C1 CDDE1 1 DE+1 891 !9)C1 9E&<1B1!DE9CDE1AC!CDE1)C11B(DE#.BC1CB19B1CBE1CD1C&&CE1BE AC1 CDDE11 DE+10CBE1D1E#A9)C!CDE1CD1&D1C189DAC1)C1D1!CB9C1CB1 B19DCB1)DEEAB1C1  CB1 ,CE1 C1 E9(A9CB-1 ,>]9CB9 EF9 A1 6II6-+1 C1 &E9CB1 AECB1 CF9DE1 D&9!C91 CB1 9ABEEB+1

4. Conditions environnementales et productions qualitative

et quantitative de lipides

"D1 BE1 C B1 DEC! B1 )C1 CB1 DEDB1 CDF9DDC!CDECB1 CFCDE1 D&CDC91 1 ! BED1#!)C1CB1!9CB1,6 C#91D1DC912F9F-+10CEEC1 9 9AEA1 CE1;E9C1 EBAC1 91 &F9BC91 1 9ED1 C1 C9EDCB1 !ACB1 DEA9;E+1 DB1 CEEC1  E)C<1 1

1 :71 ! 9A#CDBD1 CB1 !ADB!CB1 C1 1 9AED1 91 CB1 DEDB1 CDF9DDC!CDECB1 C1 1 &ED1 1 9DC1 CE1 C1 B1 9EED1 CDE9C1 CB1 &&A9CDECB1 !ACB1 9D)CB1 , 9EADCB<1 B9CB<1 CB1!C!9D9CB<1E9(A9CB-1CBE19C+1DB1C1# E9C<1DB1ECDEC9DB1C1 ! 9CD9C1!!CDE1CBE1!&A1C1 9&1 )C11 #(E DED19B)C1CB1DEDB1 CDF9DDC!CDECB1,!.9C<1DE9EC<1EC! A9E9C-1#DCDE+1

4.1. Généralités

0#C'1 1  9E1 CB1 CB .CB1 #(E DED)CB<1 CB1 DEDB1 C1 9BBDC1  E!CB1 &F9BCDE1 1 9ED1 CB1  CB1 BE9E91 , #B # CB<1 ( CB-1 1 AE9!CDE1 CB1  CB1 C1 9ABC9FC+1 DFC9BC!CDE<1 CB1 AECB1 DE1 !DE9A1 )C1 CB1 E9(A9CB1 CE1 CB1 B9CB1 BDE1!AB1DB1CB1DEDB1CDF9DDC!CDECB1)1CDCD9CDE1DC19BBDC1&C1 ,9 #(16II2<12B#D1D149M16II?-+1

$CB1 DEDB1 C1 9BBDC1 &&CECDE1 CB1 ! BEDB1  )CB1 CE1 9EA)CB1 B A&)CB1 1&&A9CDEB1E( CB1C1!C!9DCB<1CE1 91FC1C1DBA)CDC<1C9B1 9 9AEAB+1"D11!DE9A1 )C1 CB1 CCB1 FAAECB<1 B!BCB1 B1 CB1 DEDB1 C1 9BBDC1 F9CB1 ,#DC!CDEB1 CDF9DDC!CDE<1 !EEDB1 &&CEDE1 1 B(DE#.BC1  )C<1 D&CED1 F9C<1 CE+-1 BDE1  CB1 C1 9ABC9FC91 *DEA9EA1 C1 C9B1 &DEDB1 !C!9D9CB+1 4CD1 )C1 CB1 !ADB!CB1 DE!CB1 )1 9ABBCDE1 C1 )C1 D1 CE1  CC91 C1 #!ABEBC1 !C!9D9C1 D1 9CBECDE1 !1 DDB<1CB1!ADB!CB1C19AED1C1C9EDCB1 9 9AEAB1&D!CDECB1CB1!C!9DCB1 DE1AEA1CDE&AB%1

11 &EA1 ,)1 CBE1 DE!C!CDE1 AC1 B1 1 C9!AEA-1 A CD1 C1 1 DC91 CE1 C1 DBE9ED1 CB1 #LDCB1 9BBCB1 CE1 C1 1 9ABCDC1 C1 C9EDB1  CB1 ,#CBEA9<1 B #D CB-1

11 EA1B1DEC991FC1CB1 9EADCB1!C!9D9CB1CD1!&DE1C91D&9!ED1 , EA1B1C1C91CB1 9EADCB1D<1B1CB19A 9C9<1B1!&C91C91EFEA<1CE+-1

1 :91

1$1 EA1B1B 9EC91CB1BE9CBB1 #(B)CB1,!DECD1C11BE9E9C1)D11BDEA1 #DC1 91CC! C-1

4.2 Adaptation ou acclimatation ?

1 &E1 &&A9CDC91 C1 #AD!.DCB1 )1 CFCDE1 ;E9C1 9CB DBCB1 CB1 !&EDB1  )CB1 DECB1 91 CB1 #DC!CDEB1 C1 DEDB1 CDF9DDC!CDECB1 %1  EED1 CE1 !EED+1 $!EED1 BC1 E9E1 91 CB1 #DC!CDEB1 #ADE( )CB1 BFCDE1 9AFC9BCB<1 C1 DE9C1 #(B)CB<1 #!)CB1 CE1 DE!)CB+1 $ EED1 BC1 E9E1 91DC1AFED1CDEC<1CE11  9E11EC! B199AFC9BC<11ADE( C1B91D19D1D!9C1 C1 ADA9EDB1 CD1 9A DBC1 B1 DC1 9CBBD1 DE9CC+1 CB1 !EEDB1 DECCB<1 CB1 9A99DC!CDEB1 DE9AD!)CB<1 1 CB1 E9DB&C9EB1 #9'DE1 ,9C 9ED-1 BDE1 BACEDDAB1CD1&DED1C1FDEC1AFE&1)B1AD.9CDE+1891CB1CB .CB1!!C1CCB1 1 #(E DED1 )1 DE1 CB1 EC! B1 C1 ADA9ED1 E9.B1 9EB<1 1 CBE1 9&B1 &&C1 C1 AEC9!DC91C1E( C1C1!&ED1,ADE( )C11 #ADE( )C-1)1CBE1!B1CD18C1DB11 9A DBC1B1DC1DE9DEC+1

4.3 Influence de la concentration en nitrate du milieu

$CB1 &9!CB1 'EC1 D9D)C1 CE1 9D)C1 CB1 B1 !!DA!CDE1 EBACB1 91 C1

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