petroulas biodiesel

1

IV:

: .

2012

3

.......................................................................................................... .7

1: - -

1.1 ...............................................................................................9

1.2 .................................................10

1.3 .........................................................................13

1.4 ...............................................................17

1.5 ..................................................19

1.6 ..............................................................21

1.7 ...........................................................................................26

2: -

2.1 ....................................................................................................30

2.2 ...............33

2.3 ...........................................................................................41

2.3.1 ...............................43

2.3.2 ......................49

2.3.3

..................................................................................................50

2.4 .................................................................................57

2.5 .................................................58

2.5.1 .................................................61

2.5.2 ........66

2.5.3 ...............................66

2.6 ...............................................76

2.7 ..........79

4

3:

3.1 ..................................................................................................82

3.2 ............................................82

3.3 .....................................................................................85

3.3.1 ..........................................................87

3.3.2 ...........................................87

3.3.3 ...89

3.3.4 .......................90

3.3.5 .............91

3.3.6 ..............................................................93

3.4 ........................94

3.4.1 .............................................94

3.4.2 .......96

3.4.3 ..............................................96

3.4.4 .............................................98

3.4.5 ........99

3.5 ................................................................................................101

3.5.1 ..........101

3.5.2 ...................103

3.5.3 ..........................105

3.5.4 ........................................................108

3.5.5

..........................................................................................109

3.5.6 ..........................................................................111

3.5.7 .............................................................113

5

4:

4.1 ............................................115

4.2 .......................................116

4.3

SCO...................................................................................................................123

4.4 ......................................................................129

4.41 ..........................129

4.4.2

.................................................................................................131

4.5 .........................134

4.6 ..................................................134

4.7 .....................................135

4.8 ............................................137

4.8.1 Microdiesel....................................................140

4.8.2 ................................141

4.9

........................................................................................................148

5:

5.1 ..................................................................................................151

5.2 ........................................152

...........................................................................................................159

7

,

.

,

,

.

, ,

ASTM D6751.

.

,

,

,

(. Jatropha).

.

.

,

, , .

.

, ,

.

,

(Single Cell Oils, SCO).

,

20%. ,

, , , .

, ,

SCO, .

.

.

9

1: - -

1.1

.

.

:

IPCC (Intergovermental Panel on Climate Change):

, (.. )

/

,

.

[1].

UNFCC (United Nations Framework Convention on Climate Change):

[1].

: ,

, ,

, . 1995

1850. 1956

2005 1906-2005.

1961-2003 1,8mm/

1993-2003 3,1 mm/. ,

2,7%

7,4% .

1980 3. ,

50 ,

,

, . ( 1)

10

1 : .

,

.

1906-2005 (

50%).

.

.(IPCC,

Climate change 2007: Synthesis Report, .40)

1.2 -

,

, , ,

.

11

, ( 30

).

(forcings) [2].

(external forcings) ,

, ,

[3].

.

:

1. (.

)

2. (.

)

3. (

)

.

m2 , 1.370 Watt,

( 2).

30%

. 2/3

, (aerosol). To

1/3

.

.

240 Watt/m2. ,

. , .

240 Watt/m2

-19.

(GreenHouse Gases, GHG).

12

2: .

,

.

. ,

(evapotranspiration) ,

(IPCC, Frequentlyasked questions 94)

GHG

15.

( 2). GHG

.

,

[2].

13

1.3

1970-2004 GHGs 70%

( 1750)

( 3).

GHGs (aerosols)

, ,

.

GHGs ,

, .

,

, (radiative forcing [60]).

- .

/radiation

.

. /forcing

.

:

,

Watt/m2.

, -

. ,

.[2]

14

3: (a) GHG 1970-2004.

: CO2, CH4, N2O, HFCs, CFCs, SF6. (b)

GHGs 2004 CO2,-eq. (c) GHGs

2004 CO2,-eq (IPCC, Climate change 2007: Synthesis report, .36).

:

:

, ,

. CO2

[2]. 10.000

CO2 280 ppm (

). 2009 CO2 386.8 ppm

[6].

: ,

. ,

[2].

:

.

, [2].

15

:

.

CFC-11 CFC-12.

.

.

[2].

:

.

,

[2].

: .

,

. ,

.

.

[2].

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, .

()

().

.

(black carbon). ,

. aerosol

,

[2].

16

, GHGs

: CO2 ( ), CH4 (),

2 ( ), CFC-11 CFC-12.

GHGs

/ ,

. , CO2, CH4, 2,

, 1750

. 650.000 , CH4 CO2 2005

.

CO2 280ppm 2005 379 ppm. 1995-

2005 1,9ppm/ 1960-2005

1,4ppm/. , .

715 ppb 1990

1732 ppb 2005 1774 ppb.

90 . 2

270 ppb 319ppb 2005.

,

.

1750

: +1,6W/m2 ( 4) [1].

17

4 : , ,

. 2005 1750.

(IPCC, Climate change 2007: Synthesis

report, .39).

1.4

IPCC (Special Report on Emission Scenarios, SRES)

( 5).

,

(mitigation) .

GHGs

,

.

. ,

18

,

(adaptation) (mitigation).

[4].

SRES : 1,

2, 1, 2

GHGs. SRES

.

1: ,

,

(convergence) ,

. 1 3

: 1FI:

, 1: , 1:

[1,4].

2: ,

[1,4].

1:

, 1,

,

[1,4].

2: ,

.

, 2,

1

1 [1,4].

19

5: GHGs ( GtCO2-eq per year)

: 6 SRES ,

80% (post-SRES)

post-SRES. CO2, N2O, CH4, F-gases (IPCC, Climate

change 2007: Synthesis report, .44).

1.5

0,2/ SRES.

GHGs

2000 0,1/.

, GHGs ,

21 ,

20.

, .

:

20

:

, .

20-30%

,

2080

.

,

,

.

,

, , ,

.

(.

)

,

- , ,

.

, 2020 75-250

. 50%

21

. 2080

5-8%.

2050

, ,

.

.

.

,

[1].

1.6

,

, , , .

(. , ,

, ) . 1850

(, , ),

.

GHGs .

,

. ,

,

41 63 .

. ,

, (CCS) ,

(renewable energy sources RES) [6].

(renewable energy, RE) ,

, . ,

, , [7].

22

.

3 : () ()

() .

, , .

GHGs.

:

,

,

,

,

,

,

[8].

,

, . RE

,

[6].

RES .

2040 1.

[8]. 6

2008.

23

1

2040 [8]

2001 2010 2020 2030 2040

Total

consumption

(million tons oil

equivalent)

10.038 10.549 11.425 12.352 13.310

Biomass 1.080 1.313 1.791 2.483 3.271

Large hydro 22,7 266 309 341 358

geothermal 43,2 86 186 333 493

Small hydro 9,5 19 49 106 189

wind 4,7 44 266 542 688

Solar thermal 4,1 15 66 244 480

photovoltaic 0,1 2 24 221 784

Solar thermal

electricity

0,1 0,4 3 16 68

Marine

(tide/wave/ocean)

0,05 0,1 0,4 3 20

Total RES 1.365,5 1.745,5 2.964,4 4.289 6.351

RES contribution

(%)

13,6 16,6 23,6 34,7 47,7

:

:

, , ,

, ,

/ [9].

, .

.

boilers,

,

24

. -

(lignocelluloses)

(research and development, R&D) [6].

[8].

:

/ . ,

, ,

,

. , R&D

. [6,8]

:

.

.

,

, [6].

:

.

(onshore)

(offshore). .

.

.

[6,8].

:

. ,

, .

[6].

25

: , ,

,

.

,

[6].

:

.

.

.

.

, [8].

6: 2008 (IPCC, Special report on

renewable energy sources and climate change mitigation, 6)

26

1.7

, , .

, ,

.

.

, .

.

, ,

. .

.

. flexi-fuels

100%.

.

.

. [10].

.

.

[11].

1, 2

3 . 2 3

(advanced biofuels) [12]. ( 1

2 [38]).

1 : 1

(), , ..

.

.

27

,

.

1 , ,

. FFVs (Flexible Fuel Vechicles) .

, 1

.

. ,

. ,

.

. ,

GHGs

,

.

[11,12].

2 :

. 2

, , , , .

,

.

. ,

. ,

( ).

. 2

(carbon neutral)-

, 2

( ). ,

. 2

28

(R&D) [11,12]. 7

2 :

7: 2 (S.N. Naik, Renewable and sustainable Energy

reviews, .588)

1

2 ( 8).

[11].

3 : 2

.

.

1 2 . ,

3 , (),

. ,

. , ,

[11,12].

29

2 . 3

[39,12].

.

8: 1 2 (S.N. Naik, Renewable and

sustainable Energy reviews, .580)

30

2: -

2.1

.

(diesel blend).[13]

, :

: ,

,

ASTM D 6751 [14]. ( 2 ).

:

:

, 100

ASTM D 6751 [14].

: ASTM D 6751

.

[14].

31

2 : biodiesel (B100) ASTM 6751-

100 Dr. Rudolf Diesel

.

, , ,

.

, ,

.

,

. 10-17

. , (acrolein),

32

.

[25].

.

, ,

.

,

. ,

[13,17].

.

, , .

. ,

-.

. ,

, ,

. ,

, (Fatty-Acid

Methyl Esters, FAME)

(FAEE). ,

.

. FAME FAEE

, FAEE

(cloud point) (pour point)

FAME.

[13,15].

: ()

, ()

(tallow lard) () (yellow grease) ()

jatropha oil . ,

.

33

(rapeseed).

.

, GHGs,

,

.

, [13].

2.2

.

, , (CN), (cloud point CP),

(pour point PP), , (distillation

range), , ,

, (high heating value HHV).

:

:

, .

.

,

. (-

) 100 --

15 [18].

:

(ASTM D-2500) [18].

:

,

(ASTM D-97) [18].

: (flash point)

.

,

[18].

34

3 .

, , 423 ,

( 337 ) ,

, .

4 ASTM

. 5

( ).

,

,

[13].

3: (A.Demirbas,Energy Conversion and management, .15)

35

4: (S.N. Naik, Renewable and sustainable Energy

reviews, .210)

5: (EN 14214) (A.Demirbas,Energy Conversion and

management, .26)

.

36

, .

.

.

.

, .

.

. ,

417

224.

.

, ,

- , , ,

, . , -,

, ASTM

. ,

.

.

.

.

, 16-18

. ,

.

.

.

.

. (CP)

.

CP 0

37

15.

. (PP).

.

. CP

,

,

(cold fiter plugging point CFPP)

. ,

20ml

60 sec [18].

. ,

(.. -34). , ,

.

(oxidation potential).

. 11% .

. (HHV)

. H HHV (39-41 MJ/kg)

(46 MJ/kg) (43 MJ/kg)

(32-37 MJ/kg) [13,15].

.

/. 2, 5, 20,

100 2%, 5%, 20%, 100% .

(100), (20-30), (5) (2). 5

.

. ,

, .

,

38

[17].

5-20 6:

6 : ASTM 7467-10 5-20 [19]

. a US

Department of Energy (DOE) US Department of Agriculture (USDA)

[20,21]. :

(total energy

efficiency ratio, /

, ) 83,28% 80,55%

(total fossil energy efficiency ratio,

/

, )

39

.

78% .

35%

(particulate matter, PM)

32%

8%

3%

13%

79,0%

96%

(hazardous solid wastes) .

. [21]

(U.S.

Environmental Protection Agency, EPA).

9. (heavy-duty highway) .

(x),

(), (CO), (HC) [23].

40

9: [22].

20.

20

7.

7: 20 [22].

x +2.0 %

PM -10.1 %

HC -21.1 %

CO -11.0 %

8:

41

8: HHV (.... Yusuf,

Energy conversion and management, . 2746)

, x

10% . ,

(particulate matter PM), (HC),

. ,

(selective catalytic

reduction SCR) ,

[15].

2.3

,

.

() () ()

() .

, , , jatropha, ,

[13]. ,

. ,

.

,

. ,

.

42

[16]. , ,

9.

9 : (S.N. Naik, Renewable and sustainable Energy reviews, .202)

. ,

, .

.

:

.

.

(seeds) ,

.

. 10

[16].

43

10 : (S.N. Naik, Renewable and sustainable Energy reviews,

.202)

2.3.1 ( )

,

.

.

.

mole moles

( --, TAG). ( 10)

10 : [16]

, 90-98%

- -.

.

44

. , ,

.

. ,

.

.

,

[16].

, .

, (

).

[23].

.

. ( ,

, ) (

)

, .

( )

. ,

.

. , ,

.

(

/C ) ,

. ,

11 [16].

45

11 : , [16]

.

.

.

75% 25%

. C1223. [16]

, ,

, , , , .

,

, , . 12

.

12 : (S.N. Naik, Renewable and sustainable Energy

reviews, .203)

800-900

.

.

13 14

46

. 13 51

. .

, ,

.

, , , .

. , ,

. ,

. 14

75 .

[16]. 94-96%

,

[23].

26 Azadirachta indica,

Calophyllum inophyllum, Jatropha curcas, Pongamia pinnata

, .

47

13: (S.N. Naik, Renewable and sustainable Energy

reviews, .204)

48

14:

(S.N. Naik, Renewable and sustainable Energy reviews, .205)

49

,

, ,

. ,

(free fatty acids FFA).

0,05% FFA ,

FFA 0,3-0,7%, 2-7%, 5-30%

. FFA

.

, FFA

0,06%w/w 0,5%w/w.

FFA

.

[26].

2.3.2

,

.

:

. ,

,

/ (feedstock).

(FFA). %wt

.

FF %

[23].

.

[25].

FFAs.

.

50

. FF

.

.

.

.

.

.

.

. 1%

.

. ,

..

.

.

.

, , (waxes), .

.

(titre):

[26]. /

. [23].

,

[26].

2.3.3

. ,

.

, , ,

, , , , ,

51

(grape seeds) (high oleic

sunflower oil, HOS)

. 15

, ,

.

15: [27]

,

41214.

16 41214

16: (M.J. Ramos, Bioresource

technology, .264)

.

( , , ,

52

, , , ,

, )

(),

. .

(CN),

, (IV)

(CFPP) 16.

:

CN= XME(wt%) CNME

XME: , CNME:

16.

,

, . 11

.

11 :

(M.J. Ramos, Bioresource technology, .264)

.

UNE-EN 14214 CN 51.

53

, ,

grape seed. , 137

.

(C18:2) (C18:3).

, , grape

seed ( 17).

17: (wt%) (M.J. Ramos, Bioresource technology,

.263)

(C16:0) (C18:0).

. ,

.

. ,

,

.

(iodine Value) gr I2 100

gr .

.

. UNE-EN 14214 120gr I2 / 100gr.

, grape seed.

,

54

.

.

(degree of unsaturation, DU) 11.

,

DU:

, . , grape seed

.

(C16:0) (C18:0)

.

.

Rancimat ( rancimat

.

EN 14112.

[28].) 6 UNE-EN

142114. 110 .

.

.

() .

, ,

, ,

.

.

,

.

(CFPP). CFPP

16

.

(CFPP=10). (18:0) (16:1)

.

( 17)

55

CFPP. CFPP =17.

, (C22:0)

(C22:0).

.

,

. CFPP

. (Long chain saturated factor

LCSF(A))

.

.

CFPP LCSF(A) 12. CFPP

LCSF(A). CFPP

LCFS(B)

.

12: LCSF (A) LCSF(B) (M.J. Ramos,

Bioresource technology, .266)

( 13). 100%

, .

56

.

.

, , ,

, .

50wt%.

. CFPP

. , .

(

),

20-80wt%.

UNE-

EN 14214 , , .

( ).

. , ,

, ,

.

13: , .

: ,

: CFPP, :

(M.J. Ramos, Bioresource technology, .266)

57

2.4

. ,

. ,

, ,

,

[16].

() ()

( ) ()

. [17].

:

.

,

.

. (10-50%) ,

.

,

[16,13,17].

.

, , (surfactant)

(cosurfactant).

100-1000 .

, ()

, .

[17,13].

(Pyrolisis, thermal cracking).

. ,

, .

100 ,

58

.

.

, , ,

[17].

( )

.

[16].

,

. ,

[13].

2.5

14.

( 14a).

( 14 b).

( 14 c)

(, ) 14 d. [29]

59

14: () , (b) (c)

(d)

(, ,

) .

()

, ,

. ,

1846

Rochieder .

,

[13].

15.

60

15 : (.M. Shahid, Renewable and sustainable energy reviews,

.4733)

16

.

.

.

(FFA) .

FFA

.

, .

[13].

16 : [13]

, 1-8 .

61

,

, , amyl- (C5H11OH).

.

( ) ,

(, ).

,

. , -

, (2)-(4). (1)-(4)

.

(R1 COOH) + (ROH) (R1 COOR) + (H2O) (1)

+ ROH + RCOOR1 (2)

+ ROH + RCOOR2 (3)

+ ROH + RCOOR3 (4)

1 mole .

.

[13].

2.5.1

,

. , .

,

, CaO, MgO [13].

(methanolic sulphuric

acid H2SO4), (ferric sulphate Fe2(SO4)3 H2O),

(RS(=O)2-OH), (BF3).

.

.

- .

.

62

.

/. .

, ,

/

. 17

.

17 : (A.Demirbas,

Energy conversion and management, .21)

(1)

(2) (3).

,

.

18 .

,

(4)

(5) (6) [13].

63

18: (A.Demirbas,

Energy conversion and management, .22)

19.

19 : (S.N. Naik, Renewable and

sustainable Energy reviews, .207)

.

(, RO-)

.

.

. NaOH,

64

KOH, K2CO3 ,

, .

.

[13,16]. ,

. FFA

0,5% ,

FFA [30].

FFA .

, ,

[25].

. - ,

. 20

20 : [16]

. ,

[72].

, , .

,

[73].

. ,

()

.

65

. ,

.

, , .

M. Miehei

. C.antartica

[16].

2.5.2

99%. ,

, ,

- - .

.

,

.

. ,

. , -

.

(, ) , .

.

.

.

20-60.

()

() .

.

, (FFA),

, , , ,

, , .

66

.

FFA .

FFA

. ( 21)

, .

.

21: () FFA

(M.Szczesna, Renewable energy, . 1186)

solvent-free

( ) .

.

[30,31].

2.5.3

22.

.

. ,

, , ,

, pH

, .

67

22 :

(M.Szczesna, Renewable energy, .1187)

: solvent free ( 18 ).

18:

[30].

.

(regiospecificity)

.

:

68

- sn-1,3 specific ( R1 R3

- sn-2-specific ( R2 )

- non specific (

)

.

.

.

- ,

, .

, ,

, , ..

.

R.oryzae .

C18 C2-C4 .

,

.

( ) ,

50 .

( 23):

69

( ), , , , (,

), .

. ,

. (

) M.miehei, Mucor circinelloides.

solvent free T.lanuginose, .

Circinelloides .

,

.

M. miehei . solvent-free

2- C. Antarica .

. ,

, ,

23:

. (Mamoru Iso, Journal

of molecular catalysis B: Enzymatic,

.55)

70

, .

.

.

,

, , ,

. 19

.

19: (M.Szczesna, Renewable

energy,.1188)

. ,

. (

) (

).

: 24

(), (AG),

(DAG), (MAG)

7 hr, 50, 1:4 mol :mol 30%

C.antartica.

71

,

,

.

1:1 ( 40).

( )

.

--.

( : 4-5:1)

. solvent

free

. ,

, 1

.

, (:)

20 solvent-free,

40, 7 , Novozyme 435.

.

24:

(Oznur Kose, Bioresource

technology, .129)

72

20: : (Oznur

Kose, Bioresource technology, .128)

-

logP ( ).

.

.

.

.

,

, .

, (logP=4.7) n- (logP=4.0).

.

. ( 21)

73

21 : (M.Szczesna,

Renewable energy, .1190)

( ).

, . ,

/

. , -

.

, -

. ,

. , , , ,

. solvent

free .

74

, , , ,

30-55. solvent

free (30-50).

. ,

:,

.

25

P.fluorescens ( 25).

(w :

p/pH2O )

25:

P.fluorescens . (Mamoru

Iso, Journal of molecular catalysis

B: Enzymatic, .56)

75

(%). O

.

.

: w

.

( ),

.

w

. .

:

.

(

)

.

.

.

( )

.

FFA .

,

.

.

.

silica gel

76

.

,

[30].

, ,

(, ) ,

( , , ).

,

, .

.

2.6

BIOX

BIOX Dr. David Boocock

.

.

. ( )

( ).

, ,

90 min.

.

. BIOX 99%

ASTM.

[13,34].

,

, .

(supercritical methanol, SCM).

77

.

. 26

SCM

26: SCM: A: :

MeOH [29]

, ,

, ,

FFA .

.

/

.

SCM , -.

19-45 Pa, 320-350 :

40:1-42:1. 21

SCM

. 22

SCM.

78

22: SCM

(A.Demirbas, Energy conversion and management .23)

23 : [29]

, SCM .

-.

SCM .

.

nano MgO CaO SCM.

,

,

[13,29,25].

(microwaves)

.

.

79

. , 1mm 1m

300MHz 300GHz.

20% 1% NaOH 65.

60 min 8

.

. 2 min

30 min. 100%

96%.

1 min

. ,

1% NaOH.

3 9, 7.19%

100%. (100%)

- 9. ,

, / [25].

2.7

, , , .

.

- .

, - .

3 mol

1mol 3 mol 1 mol

.

.

18:1 6:1

80

. , 6:1 12:1 .

.

,

, .

318-338 .

337.9 . ,

. ,

323

. 27

.

,

[13].

27:

.

:: 1:41. : huzelnut kernel oil (A.Demirbas, Energy conversion and

management .23)

81

(FFA)

.

, FFA

,

.

FFA ,

.

.

. 28,29

FFA [13].

28 :


82

29: FFA


3:

3.1

, , ,

,

. ,

.

, .

, ,

, ,

.

70-85% . ,

,

. .

3.2

, ,

,

83

.

(oleaginous microorganisms)

20%.

.

()

(Single cell oil, SCO).

, ,

( 24).

24 : (X.Meng, Renewable energy, .2)

,

.

, SCO

.

.

. ,

(, pH, )

. ( 25)

25: (X.Meng, Renewable energy, .2)

84

CO2

.

, ,

-.

-3 -6 .

(>80%) C16 C18,

, ,

.

24 .

1-

70% 90%

. .

,

.

,

.

24 .

20-40% ,

Arthrobacter sp. Acinetobacter calcoaceticus

40% 38% .

(

12-24 ) . ,

.

,

, (

70% ) , ,

.

.

. ,

.

( )

80. Rhodosporidum sp.,

Rhodotorula sp. Lipomyces sp.

85

70% .

Cryptococcus curvatus >60%

.

90% w/w

(%SFA: saturated fatty acids) 44%,

.

, :

(18:1) (18:2) , (16:0) (16:1)

. , Rhodosporidium toruloides

Y4 16 18

.

151.5 g/l 48.0% (w/w)

, 25 .

,

. [24]

3.3

, ,

.

(moulds) ,

TAG.

(filamentous fungi)

TAG

70% o .

. ,

.

70

.

.

86

.

1500 .

.

30

25% .

Yarrowia, Candita, Rhodotorula, Rhodosporidium,

Cryptococcus Lipomyces. Yarrowia lipolytica SCO.

yarrowia

. Y. Lipolytica

Saccharomyces cerevisiae,

. 38%

.

,

,

, -.

SCO Candita, Candita

curvata. .

, Lipomyces spp. TAG.

Lypomyces starkeyi

70%

, L-

.

Cryptococcus curvatus.

.

60% .

Rhodosporidium Rhodotorula.

Rhodotorula glutinis

. Rhodotorula glacialis

-3 20. Rhodosporidium

,

,

.

87

.

Mucor circinelloides

.

.

TAG

[56] .

3.3.1

(.. )

.

. ,

,

, . ,

. -

.

TAG

.

.

(. -CoA) NADPH.

Saccharomyces cerevisiae

Yarrowia lipolytica - -

.

3.3.2

-CoA

-CoA 2

88

. -CoA

. -CoA

( 30).

.

AMP

(ammonium) .

AMP

.

. ATP

(CL) -CoA ( 30). ACL

-

. - de novo

ACC (-CoA )

-CoA -CoA.

30 :

(M.Rossi, Biodieselfeedstocks and processing technologies, .76)

, NADPH

. NADPH

89

(pentose phosphate) ,

NADH NADPH

, (malate dehydrogenase,

MDH) (malic enzyme ME).

-

. Mortierella circinelloides

, Yarrowia lipolytica

. (C14:0)

(C16:0) .

.

3.3.3

-CoA de novo

(SE) .

TAG SE

(lipid bodies, LB).

.

(lipid bodies, LB).

.

. ,

,

(bud)

. ( 31)

TAG ( 90%) .

Y.lipolytica .

S. cerevisiae 15%

TAG SE.

90

31: . AG

SE . (i-iv)

(v) (M.Rossi, Biodiesel feedstocks and processing technologies, .79)

- , .

/

. ,

TAG SE - .

3.3.4

,

S. cerevisiae Y. lipolytica. S. cerevisiae

, Y. Lipolytica

TAG.

91

.

-

TAG, .

-,

Y. Lipolytica 65%.

3.3.5

.

TAG .

, ,

. ,

/ .

- (C/N)

.

. C/N

Y. Lipolytica

Rhodotorula, Candita, Cryptococcus, Mortierella

.

Y. Lipolytica -

C/N ]

. C/N ,

Y. Lipolytica: (i)

(ii) , (iii) (iv)

. ( 32)

92

32:

.

, .

(M.Rossi, Biodieselfeedstocks and processing

technologies, .81)

( 32(i)).

,

( 32ii). ,

, ,

( 32iii).

,

, ( 32 iv)

93

3.3.6

.

.

. ,

,

SCO .

TAG. , (.

R.toruloides) .

, , , ,

. ,

.

,

.

.

,

.

,

. TAG

.

Yarrowia

lipolytica,

, .

Rhodotorula glutinis

Candita freyschussi .

/

, .

Cryptococcus curvaus.

94

. [35]

3.4

.

,

, ,

. , ,

. - (poly--

hydroxy-bytyrates) - (PHA). PHA

(inclusions) . , PHA,

, (wax esters) o

Acinetobacter . TAG

TAG. TAG

TAG.

3.4.1

TAG Gram

Gram . ,

. ( 26).

TAG .

95

26: TAG .

. Cdw : , n.r. not reported (H.M. Alvarez,Microbiol biotechnol,

.369)

TAG

Streptomyces, Norcadia, Rhodococcus,

Mycobacterium, Dietzia Gordonia. ( 26).

.

Streptomyces TAG 50-150 mg/l

60%

.

20% .

TAG

- -,

96

( 26).

Acinetobacter

TAG .

Nosostoccommune TAG

.

3.4.2

G. ,

, , n-, , -,

(coal). TAG

.

. TAG

Streptomyces Rhodococcus.

.

DGAT,

TAG Streptomyces. ,

, TAG .

, ,

TAG

. de novo

,

.

97

3.4.3

TAG :

(1) -

(2)

(3) -

sn-1,2 3 (glycerol-3-phosphate)

( 33).

.

R.opacus PD630,

2

3. TAG TAG

sn-2.

33 :

(TAG) reported (H.M. Alvarez,Microbiol

biotechnol, .370)

98

DGAT, ,

TAG

.

DGAT Mycobacterium, Streptomyces

TAG.

. , 5

.

, R.opacus PD630

TAG. DGAT

TAG, TAG.

:

(phospholipid : diacylglycerol acyltransferase, PDAT) -CoA.

PDAT

TAG.

Rhodococcus Norcadia

TAG.

3.4.4

,

.

.

. TAG

,

. TAG

.

TAG

(reductive form). TAG

. H -

TAG

(reducing equivalents) NAPD ,

99

. , - .

TAG .

TAG

. ,

TAG

.

. , TAG

. ,

TAG

.

, TAG

-C -CoA.

( 34).

34 : TAG

R.opacus PD630

.

(1)

.

()

reported

(H.M. Alvarez,Microbiol biotechnol, .372)

100

3.4.5 TAG

.

, ,

. ,

.

. TAG

80%

TAG .

, TAG

. TAG

. ,

.

. ,

.

TAG .

27 ,

. TAG

.

. ,

.

. ,

. TAG

. ,

.

,

, , , ..

.

101

.

, , [36].

().

E.coli

. , (

)

: ( 12-

24 ) [37].

27: , , . cdw:

, tr: , - : reported (H.M. Alvarez,Microbiol biotechnol, .374)

3.5

3.5.1

.

(cyanophycae)

(chlorophyta) (bacillariophyta).

102

, ,

[38].

200.000 800.000 , 35.000

[41].

. o

20%-50% . ,

.

24 ,

3,5 . ,

. ,

,

[37]. ,

,

.

.

.

.

.

:

30%(w/w) 49 132

.

, ,

,

(, ..).

,

, , .

. :

103

(bio-fixation)

, [38].

, 1 kg 1,8 kg CO2 [39].

NH4+, NO3

-, PO43-

.

,

, , ,

N:P .

,

,

.

.

.

, , , ,

.

,

, ,

, .

3.5.2

.

1% 70%

90% . 28

,

.

104

28: (T.M. Mata,

Renewable and sustainable energy

reviews,.221)

75%

.

20% 50%

. chlorella

.

,

,

.

.

,

.

7 C14:0,

C16:0, C18:1, C18:2, C18:3 .

, C16:2 C16:3 C20:5

Nanochloropsis sp. ,

105

.

, C18:1

.

29 .

,

.

29 : (T.M. Mata, Renewable and

sustainable energy reviews,.221)

3.5.3

,

.

(, ..).

35 .

,

(harvesting) , ,

.

106

35: [39]

.

,

.

, ,

,

.

:

-

107

-

( )

- (,

, , )

-

-

-

.

.

:

- :

CO2

- :

- :

CO2 .

- :

.

.

: , ,

, , , pH, ,

: (,

, ) :

, ,

108

(harvesting)

,

.

20-30%

.

-.

, (flocculation), , ,

(sendimentation), (flotation).

, ,

.

.

.

,

.

.

.

, bead mills, , .

, [38].

[39].

. (

[72]).

,

. .

, .

-,

Folch ( -).

109

.

.

(, ).

3.5.4

,

,

[41].

[42].

, (biorefinery)

. ,

,

.

,

. ,

.

10

15

. 30

. -

Chlamydomonas reihardtii ( 36).

110

.

DNA (particle

bombardment), (electroporation), Agrobacterium,

, .

.

. ,

[41].

3.5.5.

.

() . ,

.

.

omics

(genomics), (proteomics), (metabolomics),

36: Chlamydomonas reinhardtii

.

-

.

111

(transcriptomics) ,

, .

. ,

-

.

,

.

- - .

.

(cofactor)

. ( 37 )

, , .

-CoA (ACC)

-CoA

-CoA. ACC

Chlorella cryptic 2-3 .

.

.

,

. ,

. ,

.

112

37 : ER:

(R.Randakovits,Eukaryotic cell, .487)

:

-CoA,

- ,

,

. ACC

.

, . -,

.

- .

:

.

113

.

.

.

-

-CoA

.

.

ABC (

[43]). ABC

[41].

3.5.6

.

,

.

. 10%

1% 5%.

3,5

. ,

,

.

, ,

.

: .

(, , ).

[44] ( 38)

114

38:

[44]

3.5.7

,

,

,

.

:

.

.

115

(fatty acyl-CoA synthase)

. ,

Synechocystis sp. PCC8803

6.4nmol-1 OD750-1

. , PNAS

S .

13312 mg/L 0.23 g

. , , . ACP-

, ,

. 14-18

[45].

4:

4.1

SCO ,

;

.

20

.

600 30

25% . Rhodotorula spp. Cryptococcus curvatus

40-70% , Saccharomyces

cerevisiae Candita utilis 5-10%

.

( ),

( )

.

, . ,

.

116

. (malate enzyme)

( ATP:citrate lyase, ACL) .

ACL

.

, ACL

. , ACL

,

.

.

.

, SCO.

-

CoA (CCase).

.

in vivo

. ,

ACCase E.Coli L21(DE3)

6

[24].

4.2

.

:

117

.

, .

(wild type). ,

Mortirella isabellina

( 35,9 g/L)

100 g/L

[24].

( DNA)

-

(wild- type) .

[24].

.

.

.

.

in vivo .

[46].

.

118

.

.

.

.

E.coli S. Cerevisiae

.

[47].

( 39).

.

.

: (1) (2)

.

119

39 : .

(Bouke de Jong,Coiot, .3)

omics. [48] omics

omes ( omes

). (1)

, (2)

(3)

(4) omes omics

[66]. omics

(genome-wide transcription analysis). H

120

.

omics

.

.

,

.

[48].

. .

.

: 1. 2.

3.

.

. (wax ester)

Pseudomonas citronellolis, (FABEs)

E.coli (FAMEs) S. Cerevisiae.

(heterologous expression:

[43].)

(pyruvate decarboxylase)

Zymomonas mobilis

Actinetobacter baylyi DP1, .coli,

FAEE1.28 g/L in vivo 26% .

,

. ,

(C18:0) (C18:1)

,

.

121

,

, .

. 90

,

. , 20

.

[24].

,

.

.

C:N

.

( C:N) >60%

. C.

currvatus 70 g/L (C16:0)

(C18:0) (C18:1).

Bacillus subtilis

(C:N=50) . C:N

16.3% 42.7%.

[24].

122

. ,

, .

.

.

. ,

[24].

,

(biorefinering).

. ,

, .

( 40) [49].

, (), /

. -

(. , ) (.

, , ) ,

[50].

40 : [49]

123

4.3

SCO

,

20% .

TAGs (poly--hydroxy-butyrates)

(alkanoates) . ,

, .

.

. ,

.

(

).

.

70% .

. -

-

, . ,

.

,

,

. :

- -CoA

,

(FAS)

- NADPH

-CoA

ATP: (ACL, - ) ( 1)

124

.

-CoA .

+ CoA + -CoA + + ADP + Pi 1

,

.

(TCA Krebs ) .

,

, TCA, AMP

( ).

. AMP

(AMP adeaminase).

.

-CoA.

41 [51].

125

41 : /

-CoA

NAPDH . : 1.

(pyruvate decarboxylase) 2. (malate

dehydrogenase) 3. (malic enzyme) 4.

5. (citrate synthase) 6. ATP: (ATP:citrate lyase)

7.Citrate/malate translocase. : -CoA+CO2.

NADPH: NADPH+NADP++ATPNAD+NADPH+Pi.

-CoA

NADPH . (C.Ratlede, Biochimie,

54)

- AMP

- AMP

126

-

-

ACL -CoA .

- -CoA

-CoA

,

, .

ACL .

ACL.

.

(highly reduced materials)

NADPH. 1 mol

C18 16 mols DPH. NADPH

(malic enzyme).

-CoA

TAGs

.

,

Lipomyces sp.

NADPH.

NADPH .

NADPH.

TAGs [51].

,

: -CoA

.

.

:

127

,

. ,

-CoA. ([52] 281-2) 42

TG o .

30

TAGs 31

.

42:

TAG

.

.

. (.Kosa,

trends in biotechnology,.54)

(a) (FA)

. -CoA (1a)

( )

- CoA 8 ( )

(b) TAG

(16) - CoA 8

(9)

(c) (9)

(18)

(short chain alcohol, SC)

128

30: (.Kosa,


31: (.Kosa,


129

4.4

4.4.1

43 TAGs

3 : (1) -CoA

-CoA (2)

- (3) TAG. ,

[54].

(committing step)

.

, -CoA

-CoA [52]. 43

-CoA (ACC, )

-Co -CoA.

ACC -CoA

-CoA .

ACC, -

.

. ACC -CoA

. , ACC .

ACC E.coli

(accA, accB, accC, accD)

.

ACC.

-CoA -CoA:ACP

(ACP) (FAS,

). I FAS ,

, FAS

130

. FAS

-CoA -CoA.

FAS

.

(C16:0) (C18:0) (

. )

[54].

[52].

, ,

FAS.

TAG

TAG

/ ( ) .

TAG .

. TAG

(G3P) -CoA (LPA).

GPAT (acyl-CoA:glycerol-sn-3-phosphate acyl

transferase). LPA

(PA phosphatidate). PA

. TAG DGAT (acyl-CoA:diacylglycerol acyl-

transferase) 3 -CoA .

[54].

( 43) [35].

131

43 : (.M.D.Courchesne, journal of

biotechnology, .34)

4.4.2

(Genetic engineering, GE) .

32.

.

132

32 : () (.M.D.Courchesne, journal

of biotechnology, .35)

-CoA (ACC)

CC

,

.

ACC accA, accB, accC, accD E. coli BL21

. ACC .

-CoA

ACC . 6

ACC

.

,

E.coli.

ACC

133

.

.

. : (1)

ACC (2)

(bottleneck) ACC .

DGAT

DGAT TAG

-CoA.

Arabidopsis DGAT. 200-600 DGAT

3-9 TAGs.

DGAT DGAT,

,

TAG. H DGAT

TAG .

DGAT

.

.

.

ATP: (ACL), -CoA (ACS) .

,

(blocking competing pathways)

TAGs. -

.

TAG. ,

TAG.

TAG

.

CDP-

. ,

134

DGAT TAG. ,

.

-

PEPC (phosphoenolpyruvate carboxylase). TAG

- [54].

4.5

.

.

, .

, (

) [24].

4.6

. ,

. ,

,

,

pH. ,

. ,

DNA.

.

.

: ()

DNA.

135

()

.

, , . () omics

()

.

.

.

, ,

.

.

pH

.

. ,

.

,

.

,

.

hsp22.4 S.cerevisiae

51 4 .

.

.

,

.

: (rational) (combinatorial). H

.

[47].

136

4.7

:

(biochemical Engineering, BE), (genetic engineering,

GE) (transcription factor engineering, TFE).

(, pH, )

.

,

.

[54].

DNA

DNA

DNA.

,

[65]. ,

.

.

, ,

.

(Transcription factors, TFs). (transcription factor engineering TFE)

TFs

.

TFs DNA

DNA DNA -

137

. 50

DNA .

DNA

.

.

, , FE

( )

.

(bottlenecks)

. TFE.

G. ( 33) [54].

33:

(.M.D.Courchesne, journal of biotechnology, .38)

4.8

,

.

.

. ,

, E.coli, S.cerevisiae,

.

138

CoA

..

,

,

.

.

,

: (1) in vitro

(2)

.

.

. ,

,

,

.

.

,

. WS/DGAT (wax ester

synthase/acyl-CoA:diacylglycerol acyltransferase).

-CoA (fatty acid coenzyme A thiesters).

FAEEs

FAMEs. ,

139

(FAEEs) .

: E.coli

S.cerevisiae.

Kalscheuer et

al (. microdiesel).

. , . ,

, ,

E.coli .

-CoA. -

-CoA .

(wax synthase)

.

674mg/L. E.coli

11.6 mg/L.

E.coli ,

-CoA. ,

.

.

, E.coli,

.

/

,

.

, , E.coli.

- . ,

S.cerevisiae,

.

140

E.coli,

S.cerevisiae H1246, FAEE

(fatty acid isoamyl ester, FAIE),

WS/DGAT A.baylyi.

(wax synthase)(WS2) Marinobacter

hydrocarbonoclasticus S. Cerevisiae, .

WS2 wax synthase

WS/DGAT A.baylyi. ,

WS/DGAT, WS2 TAG .

TAG

. , WS2 Marinobacter hydrocarbonoclasticus

62 mg/L [37].

4.8.1 Microdiesel

2006

(FAEEs), Microdiesel, E.coli .

Acinetobacter baylyi ADP1

WS/DGAT (wax ester synthase/acyl-coenzyme

A: diacylglycerol acyltransferase) artfA.

(wax ester) TAGs

-CoA (fatty acid coenzyme A thioesters)

. WS/DGAT

:

, , - .

WS/DGAT

in vivo.

WS/DGAT

(physiological background)

,

.

141

E.coli, TAGs FAEEs (FAIEs)

Saccharomyces cerevisiae. ,

FAEEs FAIEs , FABEs E.coli,

,

WS/DGAT.

, FAEEs

.

WS/DGAT, A.baylyi ADP1, ,

.

FAEE E.coli

Zymomonas mobilis WS/DGAT

A.baylui ADP1. : artfA A.baylui

ADP1( pBBR1MCS-2::atfA)

WS/DGAT, pdc

adhB (alcohol dehydrogenase) (

pLOI297) Gram- Zymonomas

mobilis ,

FAEE. FAEE

(sodium oleate) . FAEEs .

GC-MS FAEE E.coli TOP10 (pBBR1MCS-2::atfA+

pLOI297) ,

(ethyl-oleate),

(ethyl palmitate ethyl pamlitoleate).

de novo

FAEE.

FAEE,

, ,

, FAEE

. ,

FAEE

0,26g/L 48h. 4,3 g/L ,

6,1% (w/w) FAEE.

142

, pMicrodiesel

. E.coli

pMicrodiesel FAEE.

FAEE.

pH 6-8.5 pH 7.

FAEE1,28 g/l

72h ( 5 )

4.9 g/l 26%(w/w) FAEE. ,

.

E.coli ,

FAEEs,

.

26% E.coli,

Microdiesel. FAEE

. de novo

, - ,

- WS/DGAT FAEE.

Microdiesel

, .

(70%

) , ,

TAGs.

, TAG

FAEE, Microdiesel

.

WS/DGAT (C10-C18).

WS/DGAT

.

.

H Microdiesel

.

143

.

, . Microdiesel

. Microdiesel

[55].

4.8.2

, Mucor circinelloides.

. 25%

,

. ,

- .

-

(RNAi).

,

,

M.circinelloides. :

M.circinelloides

(FAMEs)

M.circinelloides

FAMEs.

M. Circinelloides

. MU241

R7B leuA

.

,

M.circinelloides MU421

, . 3

144

.

: -, -- n-.

(19,9wt%) (1,7wt%).

(15-

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159

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