Influence du statut azoté et du cycle lumineux diurne sur le métabolisme lipidique d'Isochrysis sp. (Haptophyceae)

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Influence du statut azoté et du cycle lumineux diurne

sur le métabolisme lipidique d’Isochrysis sp.

(Haptophyceae)

Thomas Lacour

To cite this version:

Thomas Lacour. Influence du statut azoté et du cycle lumineux diurne sur le métabolisme lipidique d’Isochrysis sp. (Haptophyceae). Environnement et Société. Aix-Marseille Université, 2010. Français. �tel-00852596�

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I. L’ACCLIMATATION LIPIDIQUE DES MICROALGUES EN REPONSE A DES CHANGEMENTS DE CONDITIONS ENVIRONNEMENTALES : ETAT DES

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N-SUFFICIENT AND N-LIMITED CYCLOSTAT CULTURES OF ISOCHRYSIS SP. 1551

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1 291

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1. LES RESERVES ENERGETIQUES DU PHYTOPLANCTON ... 21

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2. LES GLYCOLIPIDES : DES ACTEURS DE LA PHOTOSYNTHESE ? ... 24 3. LES HUILES DU PHYTOPLANCTON COMME SOURCE DE BIOCARBURANT

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1

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Lumière

_{Photosynthèse}

Mise en réserve

sous forme de

sucre et TAG

Assimilation

de l’azote

Croissance

Respiration

Division

cellulaire

NO

3 Énergie, Pouvoir réducteur Énergie, Pouvoir réducteur Énergie, Pouvoir réducteur Énergie, Pouvoir réducteur

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biocarburant

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4. Organisation du manuscrit

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1 771

Sommaire

1. INTRODUCTION ... 35 2. LES LIPIDES DU PHYTOPLANCTON ... 35

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

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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

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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

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$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

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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

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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

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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