routes to cellulosic ethanol: biotechnological solutions for the sustaineble improvement of cell...
DESCRIPTION
Presentation of Marcos S. Buckeridge for the Workshop on Hydrolysis Route for Cellulosic Ethanol from Sugarcane. Apresentação de Marcos S. Buckeridge realizada no "Workshop on Hydrolysis Route for Cellulosic Ethanol from Sugarcane" Date / Data : February 10 - 11th 2009/ 10 e 11 de fevereiro de 2009 Place / Local: Unicamp, Campinas, Brazil Event Website / Website do evento: http://www.bioetanol.org.br/workshop1TRANSCRIPT
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Routes to cellulosic ethanol: biotechnological solutions for the sustainable improvement of
cell wall degradation
Marcos BuckeridgeDepartamento de Botânica
Instituto de Biociências – [email protected]
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Carbon...............................................45%Oxigen...............................................45%Hidrogen..............................................6%
Nitrogen.............................................1.5% X 6.25 = 9.4% (10%)Potassium..........................................1.0% XCalcium..............................................0.5%Magnesium.........................................0.2%Phosphorous......................................0.2% XSulfur..................................................0.1% XSilicium...............................................0.1%
Boron..................................Manganese.........................Chloride.............................. XIron..................................... XSodium............................... XZinc.................................... XCopper............................... XNickel................................. XMolibdenium.......................... X
96%
3.6%
0.4%
Obtained from CO2 and water
Macronutrients
Micronutrients
Cellulose, hemicelluloses & pectins
96-10%=86%
Pectins = 0.7%
Pectins? = 0.7%
Pectins = traces
Lipids are approximately 15% of plant tissues
Thus, the wall corresponds to ca. 70 % of the plant
In sugarne = leaves contain 68% and stem 50% plus 18% of sucrose
Proteins and Nucleic acids
The wall in the context of plant composition
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Cell walls from leaves (left) and root (below) of
legumes
A
B
ML
ML
S1S2
S3
PC
PC
2µm
V
A
PP
C
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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Cellulose: the most abundant polymer on Earth. Photograph by Cesar Gustavo Lisboa e Marcos Buckeridge, 2005
H bridges Glycosidic linkage beta-(1,4)
A B
paper
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MANGLC
HOMOGALACTURONNAN
AGA
alpha-(1,4)
methyl
AGA AGA AGAAGA AGA AGAAGA
-
-
-
-
-
-
--
-
Egg boxes divalent ion, maily calcium and magnesium induce the formatio of gels in regions that are not methylated of homogalacturonan
A
B
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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GalA RHA
RHAGalA
GalA RHA
RHAGalA
GalA RHA
GalA
ARA
GAL
GAL
GAL
GAL
GAL
GAL GAL GAL
beta (1,3)
beta (1,6)
alpha (1,6)
alpha (1,5)
alpha (1,2)
alpha (1,4)
arabinogalactan I
beta (1,4)
ARA
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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Polysaccharideo
Lignin
LigninLignin
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THE ARCHITECTURE OF THE CELL WALL
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Design: Wanderley dos Santos
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Type I Type II
microfibril
Hemicellulose strongly likd to cellulose
Hemicelluluse loosely bound to cellulose
Pectins
Proteins
Ferulic acid
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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WALL BIOSYNTHESIS
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Biossíntese da celulose: o único polímero feito no
plasmalema
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AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAA AAAAAA
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Com exceção da celulose, os demais polímeros da parede
são feitos no complexo de Golgi
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Hemicellulose biosynthesis
Buckeridge et al. 2004, Cereal Chemistry, Vol. 81 pg. 115
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PLANT DEVELOPMENT AND WALL DEGRADATION
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THE FOUR GENERATIONS OF BIOTHANOL
Cane
Sucrose
Cell Wall
acid
Enzymes
glucose, xylose e arabinose
BIOETHANOL
Cane genome Fungal genome Enzyme structure
1
1
4
4
2, 3 e 42
4
3
Rot
as p
ara
o et
anol
cel
ulós
ico
–M
arco
s B
ucke
ridge
, msb
uck@
usp.
br
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National Institute of Science and Technology of BioethanolCNPq, FAPESP
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DIANOITE
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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Cell expansion in papaya during development
50µm
50µm
A
BPC
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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A B
ATACKS OF XTH AND EXPANSIN RELINK OF XYG BY XTH AND INTUSSUCEPTION
Mic
rofib
ril 1
Mic
rofib
ril 2
Mic
rofib
ril 1
Mic
rofib
ril 2
New
Mic
rofib
ril
expa
nsin
Buckeridge et al. 2008. Parede Celular, Cap 9 in Kerbauy G.B. Fisiologia Vegetal. Guanabara Koogan
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Tiné, Braga, Hahn, Freshhour & Buckeridge, unpublished results
Cotyledons of Hymenaea courbaril (jatobá)
Storage walls can be very complex
M1 antibody binds specifically to fucosylated XGs, which are present only in primary cell walls
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Xyloglucan
XGOs
Xyl
Sucrose
Degalactosylated XGOs
XTH
hcbetagal
beta glucosidase
Glc
alpha xylosidase
Gal
P-sugars ?
sucrose synthase
Auxin
DNA
mRNA
auxin-conjugate
LIGHT
NPA treatment
Shoot excision
Sucrose
GROWTH
Starch
P-sugars
sucrose synthase
invertase
Pentose P pathway ?
Starch
coty
ledo
n
hyp
ocot
yl
leaf
phy, cry ?
?
?
??
invertase
Brandão, Del Bem, Vincentz & Buckeridge. Journal of Experimental Botany, 2009 in press
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Cotyledons of Lupin: one enzyme does the job
Buckeridge et al. 2005. Annals of Botany, Vol.96:435.
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Sugar composition of sugarcane leaves
0
10
20
30
40
50
60
70
fucrham ara gal
glcxyl
0,240,21
12,58
2,32
14,52
69,88
Collaboration with EMBRAPA Bioenergy and INCT do BioetanolMaria Thereza Bazzo Martins, Amanda P. De Souza, Hugo Molinari & Marcos Buckeridge
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Parede celular e
senescência foliar
876
5
4
3 2
1
EMBRAPA agroenergiaPlantas de RB867515 coletadasMaria Thereza Bazzo
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0,00,10,20,30,40,50,6
+1 +2 +3 +4 +5 +6 +7 +8
Fuc %
0,00,10,10,20,20,30,30,40,4
+1 +2 +3 +4 +5 +6 +7 +8
Rham %
11,011,512,012,513,013,5
+1 +2 +3 +4 +5 +6 +7 +8
Ara %
1,8
2,0
2,2
2,4
2,6
2,8
+1 +2 +3 +4 +5 +6 +7 +8
Gal %
1011121314151617
+1 +2 +3 +4 +5 +6 +7 +8
Glc %
64
66
68
70
72
74
+1 +2 +3 +4 +5 +6 +7 +8
Xyl%
Do composition during leaf senescence?
Collaboration with EMBRAPA Bioenergy and INCT do BioetanolMaria Thereza Bazzo Martins, Amanda P. De Souza, Hugo Molinari & Marcos Bucker
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0,00
0,10
0,20
0,30
0,40
Base Middle Tip
Fuc %
0,000,050,100,150,200,250,30
Base Middle Tip
Rham %
1,51,71,92,12,32,52,7
Base Middle Tip
Gal %
11,0
11,5
12,0
12,5
13,0
13,5
Base Middle Tip
Ara %
12,0
13,0
14,0
15,0
16,0
Base Middle Tip
Glc %
64
66
68
70
72
74
Base Middle Tip
Xyl%
Do composition change along the leaf blade?
Collaboration with EMBRAPA Bioenergy and INCT do BioetanolMaria Tereza Bazzo Martins, Amanda P. De Souza, Hugo Molinari & Marcos Buckeridge
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From 1999 to 2001, the SUCEST genome program produced 238,000 ESTs from various tissues of the
sugar cane plant.
Since then we found:1) 469 cell wall related genes in different cane tissues
(Lima et al. 2001, GMB)1) Determined the chemical composition and structure of the cell wall
polymers of different sugarcane tissues
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How to modify the wall to obtain energy?
Microorganisms
Hydrolytic enzymes
Action on the bagasse wall
Free fermentable sugars
Change polymer structure
Change synthesis
Change wall architecture
Control of hydrolysis
Activation of endogenous hydrolysis
MODIFIED WALL
Ethanol
Fermentation
Increase wall and decrease sucrose
“Papaya Cane”or” Energy cane”Increase accessibility Increase accessibility
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THANK YOUMarcos Buckeridge
Departamento de BotânicaInstituto de Biociências – USP
http://bioethanolbrazil.wordpress.com
Hugo MolinariEMBRAPA agroenergia
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Sesbania virgata, a galactomannan storing seed
1 2A 2B
3 4
Barr
a=84
µm
Barr
a=34
µm
Barr
a=22
µm
Barr
a=40
µm
ex
me
en
e
ex
me
e e
end
sc
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Galactomannan degradation in S. virgata
0 1 2 3 4Time (days)
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PhD thesis Patricia Tonini
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Microfibrilas: 2-20 nmdiametro e 100 - 40 000 nm de comprimento
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B
D
Quais genes?(sequenciamento completo)
Expressão gênica
Proteômica
B
D
B
D
CANA ENERGIA
FermentaçãoHidrolases fúngicas
(modo de ação, cristalografia e
genética)
SinalizaçãoCelular
CO2
TemperaturaÁgua
Seqüências e propriedade das
proteínas
Controlar a arquitetura da parede
Estudar a variabilidade genética
Xilose ?
ROTAS PARA O ETANOL LIGNOCELULÓSICO
ETANOL
Mitigação e adaptação
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A parede celular é composta por
polímero que são uma combinação
de monossacarídeosPode-se comparar
o número de combinações possíveis em
relação a outras macromoléculas
Considerando uma sequência..A-B-C-D-E-
Ácidos Nucléicos:
Quatro pares de bases ......45 = 1024
Proteinas:
Vinte aminoácidos....205 = 3,200,000
Carboidratos:
Dez monossacarídeos.... 105 = 100,000Quatro hidroxilas (hexoses) ou três (pentoses).... x 3.55 = 525
Ligações alfa ou beta... X 25 = 32
NÚMERO TOTAL DE POSSIBILIDADES = 1,680,700,000
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Estrutura e conformação da
celulose
cellulose
amilose
dextranos
glucanos
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Então, a parede poderia ser como um cristal líquido.....
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Microfibrila de celulose
Xiloglucano
Ramnogalacturonano I
Arabinogalactano
Extensina com tetrassacarídeos de arabinose (losangos)
Ponto em que foi proposta ligação covalente entre xiloglucano e pectina
Ponto em que foi proposta ligação covalente entre extensina e pectina
Figura 10 . Modelo de parede celular proposto em 1973 por Peter Albersheim e colaboradores. Neste, as ligações covalentes (exemplos circundados) seriam a principal forma de manter os diferentes polissacarídeos em interação. Note porém, que a interação não covalente entre xiloglucano e celulose já havia sido proposta neste modelo.
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MMMMM
MGMGM
MMMMMG
GMMMMM
G
G G GMMMMM
G G
G G GMGMGM
A AMMMMM
A A
Har
dnes
s
Water retention
Coffee, Palms
Legumes, Ferns
Orc
hids
,B
rom
elia
ds
Orc
hids
, B
rom
elia
ds,
Fern
s,
Ast
erac
eae
Legumes, Ferns
GGGG?
(cellulose)
Alp
ha g
alac
tosi
dase
Gal
acto
syl t
ranf
eras
eCesA Csl 3
Csl 4
Galactosyl transferase
Acetyl transferase
XG
XXGG XXXXG
LG
FG XG
SG
JG
XXXG
PP,PT,TP
Cel
lulo
se-X
g bi
ndin
g co
ntro
l ?B
eta
gluc
an
Xylan
(Csl n)
GAX
Ara
tran
(Csl
6)
Evolution of the cellulose-hemicellulose
domain in plants
Epiphytism
(Csl
5)
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GLC GLC GLC GLC GLC GLC GLC GLC
beta-(1,4) CELLULOSE A
B
GLC GLC GLC GLC GLC GLC GLC GLC
XYL XYL
XYL
GAL
FUC
alpha (1,6)
beta (1,2) alpha (1,2)
Action of XTH e cellulases
XYL
XYL
XYL
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Xyloglucan conformation
Bacterial cellulose
Without Xg With Xg
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XYL XYL XYL XYL XYL XYL XYLXYL
GalA
beta-(1,4) ARAARA
GLC
GLC GLC GLC GLC
GLC GLC GLC
beta-(1,4)GLC
beta-(1,3)
GLUCURONOARABINOXYLAN (GAX)
MIXED LINKAGE GLUCAN (MLG)
alpha-(1,3)
alpha-(1,6)
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MAN MAN MAN MAN MAN MAN MAN MAN
beta-(1,4) PURE MANNAN
MAN GLC MAN GLC MAN GLC MAN GLC
beta-(1,4) GLUCOMANNAN
acetyl
MAN MAN MAN MAN MAN MAN MAN MAN
GAL
GAL
beta-(1,4)
alpha-(1,6)
GALACTOMANNANO
GAL
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Figura 12. Modelo descontínuo da parede celular. Neste desenho a parede é mostrada em “corte transversal” ao eixo das microfibrilas. Ao invés de se sustentar por ligações covalentes, a parede celular primária e composta por três domínios independentes: Pectinas, Celulose-hemicelulose e Proteínas. Os três coexistem independentemente, ou seja sem ligações químicas covalentes, mas por interações fracas (pontes de hidrogênio).
Pectinas Celulose-Hemicelulose Proteínas
DO
MÍN
IOS
PAR
EDE
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Cell wall deposition is not uniform
Buckeridge et al. 2004, Cereal Chemistry, Vol. 81 pg. 115