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.9):`+F1c)+-*,:;2-+C)#)<9+5-789)*1)-1+=.>;,5-).-3)+-?413)*1):-.-)*1)-7@:-+)d):9.3F,F;F,K-)1)09;:9)

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*1)-F,K,*-*13)1.4,5\F,:-3)-339:,-*93)])<9+5-789)*9)-1+=.>;,5-)15)+-?413)*1):-.-)*1)-7@:-+():95)

=.<-31) .9) 0-012) *-3) M,*+92-313) *1) 0-+1*1) :12;2-+) 1) 15) -26;5-3) 0+9F1?.-3) +12-:,9.-*-3) ]) /Z%C)

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0+9F1N5,:-) 1) -F,K,*-*1) 1.4,5\F,:-) *-3) 62,:93,2) M,*+92-313) 1) 9;F+-3) 0+9F1?.-3) >;1) -F;-5) .-)

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B) 1) OGC) _) -F->;1) 9:9++1) 0+,.:,0-251.F1) 39V+1) -3) 01:F,.-3) 1) 9) ~H62;:-.9C) Z9.F;*9() 93) *-*93)

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31W-5)*16+-*-*93)-9)29.69)*9)0+9:1339()15V9+-)1.4,5-3)1301:?<,:-3)*1F1:F-*-3)0933-5)-F;-+).-)

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.93) 316C) A) 1) BC) Z9.F;*9) >;-.*9) +1F,+-H31) -)5-,9+) 0-+F1) *-3) 01:F,.-3) 1) M15,:12;29313) -0`3) 0+dH

F+-F-51.F9):95)$-_v()-)3-:-+,<,:-789)d)5-,9+).93)31651.F93)B()O)1)Q()*1K,*9)-9)5-,9+)-:1339)1)-)

5-,9+)>;-.F,*-*1)*1):12;2931C)#3)62,:93,2)M,*+92-313)1.:9.F+-*-3).13F1)F+-V-2M9)3;61+15)>;1)13F-3)

-F-:-5)-)0-+1*1)*1);5)1301:?<,:9):9.W;.F9)*1):d2;2-3)*9):`+F1c)>;1)*\)9+,615)-9)-1+=.>;,5-C))S\)

.9) <,5) *9) 0+9:1339() >;-.*9) M\) 2,31) :12;2-+() -26;5-3) 0-+1*13) *1) :d2;2-3) +15-.13:1.F13) 389)

+1:-2:,F+-.F1)])M,*+`2,31()0+9K-K1251.F1)*1K,*9)-)3;-)-+>;,F1F;+-)1):95093,789C))e3F1)F+-V-2M9)F+-4)

,.<9+5-7X13)0-+-)9)*131.K92K,51.F9)*1)<;F;+-3)F1:.9296,-3)0-+-)-)0+9*;789)*9)1F-.92)*9)1F-.92)

:12;2`3,:9)*1):-.-H*1H-7@:-+C)

)

/020B:08$ @W0B7L) -1+=.>;,5-() 0-+1*1) :12;2-+() 62,:93,2) M,*+92-313() 0+9F1N5,:-() 59+F1) :12;2-+)

0+96+-5-*-C)

$

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Page 12: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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+,01.,.6)-.*)31.13:1.:1()<9+),.3F-.:1(),.K92K1)FM1)-:F,K-F,9.)9<)<;.:F,9.-2)59*;2,)FM-F):;25,.-F1),.)

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:122)�-22)59*,<,:-F,9.3()<9+5,.6)-,+)30-:13),.)FM1):9+F1c)9<)FM1)+99FC)DM1)<9+5-F,9.)9<)-1+1.:M^5-),.)

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:95093,F1)FM-F)3;++9;.*3)FM1)-,+)30-:13C)DM,3)3;6613F3)FM-F)0-+F)9<)FM1)092^3-::M-+,*13)0+131.F),.)

:122)�-223)-+1).9F)*16+-*1*)*;+,.6)FM1)0+9:133()-2FM9;6M)301:,<,:)1.4^513)M-K1)V11.)*1F1:F1*)FM-F)

:9;2*) -:F) 9.) 092^3-::M-+,*1)59V,2,4-F,9.() 3;:M) -3) FM1) !vAk) <-5,2^C) Y;+FM1+() ;.*1+) 0+1F+1-F51.F)

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FM1)9FM1+)M-.*()�M1.)593F)9<)FM1)01:F,.3)-.*)M15,:122;29313)-+1)+1F+,1K1*)V^)0+1F+1-F51.F)�,FM)

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Page 13: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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Page 14: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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<9+5-789)*1)-1+=.>;,5-)+-*,:;2-+)EeK-.3)BTTOGC)e3F1)@2F,59)<9,),*1.F,<,:-*9)15)+-?413)*1):-.-)*1)

-7@:-+() :959) +13;2F-*9) *-) 59*,<,:-789) *1) 0-+1*1) :12;2-+) ,.:2;,.*9) -) M,*+`2,31) *1) 01:F,.-3) *-)

2-512-) 5d*,-) 1) ~H62;:-.9) E[1,F1) BTAOG() 09+d5) -3) 1.4,5-3) +1309.3\K1,3) 09+) 13F-3) 1) 9;F+-3)

59*,<,:-7X13)-,.*-).89)<9+-5),*1.F,<,:-*-3C)

"#-4(52*+"P%;?JEDEKL:Q%:I:CCRDIEA%?%INA<<EJEIAKS?<%

_)-1+=.>;,5-)d);5)F1:,*9)1301:,-2,4-*9)>;1)09*1)31)<9+5-+)15)<92M-3()01:?9293():-;213)1)

+-?413):9.3,3F,.*9)15)130-793) ,.F1+:12;2-+13) ,.F1+2,6-*93) 29.6,F;*,.-251.F1()>;1) <-:,2,F-)-)*,<;389)

*1) 6-313)0129) F1:,*9) E#+53F+9.6)AjkjGC) /9*1)-F;-+) F-5Vd5):959)*,<;39+)*1)9;F+93) 6-313) :959)

51F-.9()*,`c,*9)*1):-+V9.9)1)1F,21.9) EZ9251+)BTTOGC) g)1.:9.F+-*9)0+,.:,0-251.F1)15)+-?413)15)

+13093F-)-*-0F-F,K-)-);5-)-502-)6-5-)*1)<-F9+13)-5V,1.F-,3)>;1),.:2;,)M,0`c,-)ES;3F,.)1)#+53F+9.6)

AjjA() %+1�) AjjkG() +13,3F=.:,-)51:L.,:-) Ev1) 1F) -2C) Ajj|VG() <-2F-) *1) .;F+,1.F13) E%+1�) 1F) -2C) Ajlj()

b9.,.63)1)f1+3:M;+1.)AjlT()I9;+-.,3)1F)-2C)BTTOG)1)-)*1331:-789)E%+1�)AjkjGC))

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! "&!

'89)1.:9.F+-*93)*9,3)F,093)*1)-1+=.>;,5-s)9)0+,51,+9)d)*1.95,.-*9)-1+=.>;,5-):9+F,:-2()

:959)9)1.:9.F+-*9)15)+-?413)*1)-++94)EV,10)%3)-.8)G()5,2M9)EW$)%2)13G():1K-*-)EX",*$42%84#+),$G()

F+,69)E7,.-.'42%)$3-.842G)1):-.-)*1)-7@:-+)E#+53F+9.6)Ajkj()eK-.3)BTTO()S-:U39.)1)#+53F+9.6)Ajjj()

$,3M,;:M,) 1F) -2C) BTAB() [1,F1)BTAOGC)_)9;F+9) F,09)d);5) F1:,*9)V+-.:9)1)1309.W939)0+11.:M,*9) :95)

130-793) *1) 6\3) >;1) 31) <9+5-) .9) :-;21() M,09:`F,293() +-,4) 0+,.:,0-2() +-?413) -*K1.F?:,-3) 1) .`*;293)

+-*,:;2-+13) *1) 216;5,.93-3() :959) -) 39W-) E?#1'./$% 2)GG() J$3S)/.)% ,"3-,)-)) 1) Y.2./),.)% Z4/'$))

:;2F,K-*-3)15):9.*,7X13)*1)-2-6-51.F9) E#+,U-*9)AjRQ()Ä-2U1+)1F)-2C)AjlO()'-+-3�-F,)1F)-2C)AjjB()

P9:M,4;U,)1F)-2C)BTTTGC))

_) -1+=.>;,5-) 09*1) 31+) +13;2F-*9) F-.F9) *1) ;5) 0+9:1339) 13>;,4`61.9) >;-.F9) 2,3?61.9)

E#+V1+)AjBT()S-:U39.)1)#+53F+9.6)Ajjj()'1-69)1F)-2C)BTTRGC)_)-1+=.>;,5-)13>;,4`61.9()61+-251.F1)

:9.3F,F;F,K9() 9:9++1) >;-.*9) 130-793) 6-39393) ,.F1+:12;2-+13) 389) <9+5-*93) *;+-.F1) 9)

*131.K92K,51.F9()315)9)0+9:1339)*1)59+F1):12;2-+)0+96+-5-*-C)e3313)130-793)389)<9+5-*93)0129)

:+13:,51.F9) :12;2-+) *,<1+1.:,-2() :95) :d2;2-3) -*W-:1.F13) 310-+-.*9H31) ;5-3) *-3) 9;F+-3) 1.F+1) -)

2-512-) 5d*,-() 0+9K-K1251.F1) :95) -) *,613F89) -01.-3) *-) 2-512-) 5d*,-) E[--.) 1F) -2C) Ajlj() AjjA()

f,331+) 1F) -2C) BTTO() eK-.3) BTTOGC) J5) 1c15029) *1) -1+=.>;,5-) 13>;,4`61.9) 13F\) .9) 01:?929) *1)

J)+.--),.)%#)/'.&"#.))E':M;3321+)1F)-2C)AjjkGC)_)-1+=.>;,5-)13>;,4`61.9)d);5)0+9:1339):95021c9)1)

9+*1.-*9() 09+d5) 3;-) +16;2-789) 1) 93) 12151.F93) :M-K1) >;1) 21K-5) ]) 3;-) <9+5-789) 389) 09;:9)

13F;*-*93)EeK-.3)BTTOGC)))

_)-1+=.>;,5-) 2,3?61.9()9.*1)0933;15)5-,9+)0-+F1)*93)13F;*93()09*1) 31+) :9.3F,F;F,K9)9;)

,.*;4,*9) E09+) -2-6-51.F9) 9;) 1F,21.9G) 1) 1.K92K15) .1:133-+,-51.F1) 59+F1) :12;2-+) 0+96+-5-*-)

316;,*9)*-)*16+-*-789)*-)0-+1*1):12;2-+C)/9*1)31+)1.:9.F+-*9)15)130d:,13)*1):1K-*-)E#+,U-*9)1)

#*-:M,)AjRRG()F+,69)ED+9;6MF)1)%+1�)AjlTG()-++94)ES;3F,.)1)#+53F+9.6)AjjAG()5,2M9)Ev1)1F)-2C)Ajj|-()

!;.-�-+*1.-)1F)-2=)BTTA-G)1):-.-)*1)-7@:-+)E[1,F1)BTAOGC)

+:C9?%I?N>NAC%MC:OCA@A;A%T!1&U%

_) 0+,51,+9) 3,.-2) >;1) *131.:-*1,-) F9*9) 9) 0+9:1339) *1) <9+5-789) *9) -1+=.>;,5-) 2,3?61.9)

,.*;4,*9)d)-)+1*;789)*9).?K12)*1)9c,6=.,9).-3)+-?413();5-)K14)>;1).-)\6;-)-)3;-)*,<;389)*1)9c,6=.,9)

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! "'!

0-33-)-)31+)ATQ)K1413)51.9+)>;1).9)-+)E#+53F+9.6)1)%+1�)BTTBGC)#0`3)-)+1*;789)*-)9c,61.-789)M\)

9)-;51.F9)*9)1F,21.9)1)*9)Z-BÉ) :,F93`2,:9)1)-33,5)1.F+-)15)1c1:;789) F9*9)9)0+9:1339)*1)/Z%)1)

:9.31>;1.F1)<9+5-789)*9)-1+=.>;,5-)EY-61+3F1*F)BTATGC))

$-) <9+5-789) 2,3?61.-()5;,F-3) 02-.F-3) *,<1+15) .9) >;1) *,4) +1301,F9) ]) *101.*=.:,-) *1) ;5)

,.*;F9+) -5V,1.F-2) 0-+-) >;1) 9) -1+=.>;,5-) 31) ,.,:,1C) e5) -++94:) -) <9+5-789) *1) -1+=.>;,5-) .89) d)

,.*;4,*-() 09,3) -26;.3) F+-V-2M93) 593F+-+-5) >;1) .89) *101.*1) *1) M,0`c,-) 0-+-) 31+) <9+5-*9()

9:9++1.*9)51359)15)39293)-1+-*93)Eb-�-,)1F)-2C)AjjlGC)e.F+1F-.F9()51359)>;-.*9)d):9.3F,F;F,K9()

9)1F,21.9)d)+1>;1+,*9)0-+-)3;-)<9+5-789)Ea;U,^93M,)1)b-+-M-+-)BTAQG():9.F;*9()0-+-)9;F+-3)02-.F-3)

-) <9+5-789) *1) -1+=.>;,5-) 0-+1:1) 31+) ,.*101.*1.F1) *13F1) M9+5N.,9() :959) 09+) 1c15029) 15)

130d:,13)*1)+16,X13)-2-6\K1,3):959)[4/'43%$&&4343)Ef,331+)1)I9615-..)BTT|GC)S\)0-+-)130d:,13)5-,3)

13F;*-*-3) :959) 9) 5,2M9() -) <9+5-789) *1) -1+=.>;,5-) d) ,.*;4,*-) 012-) V-,c-) :9.:1.F+-789) *1)

9c,6=.,9)1i9;)0129)-;51.F9)*1)1F,21.9) E%+1�)1F)-2C)Ajlj()%+1�)1F)-2C)BTTTG)>;1) ,.*;415)-)/Z%C))

e.F+1F-.F9() :959) K,3F9) 09+) 13313) F+-V-2M93() 3,.-2,4-.*9) 9;) .89) K,-) 1F,21.9() -3) :d2;2-3) +1:1V15)

.1:133-+,-51.F1)-)3,.-2,4-789)K,-):\2:,9()9)>;1)+13;2F-).-)/Z%)1):9.31>;1.F151.F1) <9+5-789)*9)

-1+=.>;,5-)2,3?61.9C)

#)/Z%)d);5)0+9:1339)9+*1.-*9)*1)3;,:?*,9):12;2-+)>;1).1:133,F-)*1);5-)-F,K-)1c0+13389)

6=.,:-() 9:9++1) >;-.*9) M\) ;5) :1+F9) .@51+9) *1) 1K1.F93) *9) *131.K92K,51.F9) 1) <-:,2,F-) :95) -)

+159789)*1)F,093):12;2-+13)1301:?<,:93)>;1).89)389)5-,3)1c,6,*93)E[-5)1F)-2C)BTTAGC)g):9.M1:,*-)09+)

9:9++1+)*;+-.F1)1K1.F93)*9)*131.K92K,51.F9)*-3)02-.F-3):959).-)31.13:=.:,-)*1)<92M-3)1)<+;F93()

.9)*131.K92K,51.F9)*93)12151.F93)F+->;1-,3()*;+-.F1)-)M,01+31.3,V,2,*-*1)15)+13093F-)-9)-F->;1)

*1)0-F`61.93)1)15)1K1.F93)5;,F9)31512M-.F13)-)13F1)9:9++15)*;+-.F1)-)<9+5-789)*9)-1+=.>;,5-)

39V)13F+13313)-5V,1.F-,3)F-,3):959)M,0`c,-)E/1..12)1)[-5V)Ajjk()Y-61+3F1*F)BTATGC))

_),.?:,9)*9)*131.K92K,51.F9)*9)-1+=.>;,5-)F15):959):-+-:F1+?3F,:-3)0+,.:,0-2)-)/Z%()>;1)

:950+11.*1) 93) 316;,.F13) 0+9:13393) :12;2-+13) 3;V31>;1.F13s) :9.*1.3-789) *-) :+95-F,.-()

<+-651.F-789) *1) %$#) 1) <9+5-789) *1) :9+093) *1)515V+-.-3) >;1) :,+:;.*-5) -3) 9+6-.12-3) E"11*)

BTTT() !;.-�-+*1.-) 1F) -2C) BTTA-GC) Y9+-5) 9V31+K-*93) 3,.-,3) *1) *16+-*-789) *9) .@:219) :12;2-+)

*;+-.F1) -) 59+F1) *9) :`+F1c) *-3) +-?413() 1) 13F13) 31) ,.,:,-5) *9) 1cF1+,9+) *9) :`+F1c) 1) -K-.7-5) 15)

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*,+1789)-9)13F129)Ev1.+^)1)%1-:9.)AjlAG().-)5135-)*,+1789)15)>;1)93)130-793)*1)-+)389)<9+5-*93)

.9):`+F1cC)

%;+-.F1)-)/Z%()-)3,.-2,4-789)09+)1F,21.9)d)-:950-.M-*-)09+);5)-;51.F9):,F93`2,:9)*1)Z-BÉ)

:95)093F1+,9+)-:,*,<,:-789)*9):,F902-35-()12151.F93)13F13)+1>;1+,*93)0-+-)>;1)31),.,:,1)-)59+F1)*1)

:d2;2-3) :9+F,:-,3) *1) +-?413) EP9+6-.) 1) %+1�) BTTQGC) Z9++9V9+-.*9) 133-) ,.<9+5-789() 15) -++94) <9,)

9V31+K-*9) >;1) M\) ;5) -:,*,<,:-789) .9) 0v) :,F902-35\F,:9() >;1) 31) ,.,:,-) 15) :d2;2-3) ,392-*-3)

1301:,<,:-3)*9):`+F1c)5d*,9)-.F13)*1) <9+5-+)9)-1+=.>;,5-)Eb-�-,)1F)-2C)AjjlGC)e3F-)3,.-2,4-789)d)

31512M-.F1) ]>;12-) >;1) 9:9++1) >;-.*9) M\) -) <9+5-789) *93) 12151.F93) F+->;1-,3) EP9+6-.)1)%+1�)

BTTQGC)

) #) :9.:1.F+-789) *1) Z-BÉ) :,F93`2,:9) 2,K+1) d) <+1>;1.F151.F1) 5-.F,*-) 1.F+1) ATT) -) BTT) .P)

EI;3M)AjjRGC)/9+d5()133-):9.:1.F+-789)09*1)-;51.F-+)09+):-;3-)*-)-<,.,*-*1)*9)Z-BÉ)])-26;5-3)

0+9F1?.-3():959)09+)1c15029()-3):-259*;2,.-3)E'-.*1+3)1F)-2C)AjjjGC)_)+1F,:;29)1.*902-35\F,:9()-)

0-+1*1) :12;2-+() -3) 5,F9:N.*+,-3() K-:@9293) 1) 93) :29+902-3F93) F=5) -) :-0-:,*-*1) *1) 31>;13F+-+) 1)

-+5-41.-+)9)Z-B)15);5-)13:-2-)*1)5,2,592()9)>;1)d)5;,F9)5-,9+)*9)>;1)-):9.:1.F+-789).9):,F9392)

EI;3M)AjjR()D+1�-K-3)1)P-2M9)AjjlGC)e5):d2;2-3)-.,5-,3()93)0+9:13393)*1)3,.-2,4-789)K,-)Z-BÉ)389)

512M9+)13F;*-*93C)$13F13()-3)5,F9:N.*+,-3)*131501.M-5);5)0-012) :1.F+-2).-):-3:-F-)*1)59+F1)

:12;2-+() ,.F16+-.*9) -2F1+-7X13) 51F-V`2,:-3) E>;1*-) .9) 0v() -;51.F9) *-) :9.:1.F+-789) *1) Z-BÉ) 1)

-2F1+-7X13) .9) 13F-*9) 1.1+6dF,:9G) :95) -) /Z%C) e5) 316;,*-) 9V31+K-H31) ;5) -;51.F9) *-)

01+51-V,2,*-*1) *-) 515V+-.-) 5,F9:9.*+,-2) 2,V1+-.*9) 9) :,F9:+959% '% 1) ;5) <-F9+) *1) ,.*;789) *1)

-090F931)>;1):9.*;4)])-F,K-789)*1):-30-313)9;):,3F1?.9H0+9F1-313)EY9+F13)1F)-2C)BTTA()f,+92-,.1.)1F)

-2C)BTTBGC)e5)1c01+,51.F93):95)F+,69()<9,)9V31+K-*-)-)-V1+F;+-)*9)09+9)5,F9:9.*+,-2)1)2,V1+-789)*9)

:,F9:+959% ') -01.-3) .-3) :d2;2-3) :95) <-2F-) *1) 9c,6=.,9) 1) >;1) <9+-5) :,+:;.*-*-3) 09+) -2F-3)

:9.:1.F+-7X13)*1)Z-BÉ()+1<9+7-.*9)-)0-+F,:,0-789)*9):\2:,9).-)3,.-2,4-789)0-+-)-)/Z%)9V31+K-*9)15)

:d2;2-3)-.,5-,3)EY9+F13)1F)-2C)BTTA()f,+92-,.1.)1F)-2C)BTTBGC)

Z959)9)0+9:1339)*1)/Z%)>;1)0+1:1*1)<9+5-789)*9)-1+=.>;,5-)F15)5;,F-3):-+-:F1+?3F,:-3)

15) :95;5) :95) -) /Z%) *93) 12151.F93) F+->;1-,3() 3;61+1H31) >;1) ;5-) K-+,1*-*1) *1) 1.4,5-3)

M,*+92?F,:-3) 13F1W-5) 1.K92K,*-3) 15) -5V93) 93) 0+9:13393) EP9+6-.) 1) %+1�) BTTQGC) e5V9+-) M-W-) -)

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*1F1+5,.-789) *-3) :-+-:F1+?3F,:-3)59+<92`6,:-3) *9) -1+=.>;,5-) 15) *,K1+3-3) 02-.F-3() -) 3,.-2,4-789)

>;1)21K-)-9)-F->;1)]))0-+1*1):12;2-+)d)09;:9):9.M1:,*-C)#:+1*,F-H31)>;1)-0`3)9):92-039),.F1+.9)*-3)

:d2;2-3()9V31+K-*9)15)5,:+93:90,-)121F+N.,:-)*1)F+-.35,3389)09+)Z-50V122)1)%+1�)EAjlOG()-)01+*-)

*-) ,.F16+,*-*1) *9) F9.902-3F9) 01+5,F,+,-) 9) -:1339) *-3) 1.4,5-3) >;1) 13F89) .93) K-:@9293) 1) .9)

:,F902-35-C)Z95)13F1)1K1.F9()-)*,3392;789)*-)515V+-.-):12;2-+)09*1+,-)13F-+):95021F-)*1.F+9)*1)

BQ)M9+-3)>;1)d)>;-.*9)31)9V31+K-5):d2;2-3),.*,K,*;-,3),.,:,-.*9)-)59+F1):12;2-+)EeK-.3)BTTOGC)

+:;EJEIAKS?<%=>?%:I:CC?@%DA%MAC?;?%I?N>NAC%;>CAD9?%A%J:C@AKL:%;:%A?CRD=>E@A%

#3) 0-+1*13) :12;2-+13) <9+.1:15) :-+-:F1+?3F,:-3) ,509+F-.F13) 0-+-) -3) 02-.F-3() 09,3) 389) 12-3)

>;1)*131501.M-5);5):9.W;.F9)*1) <;.7X13)1331.:,-,3() :959)09+)1c15029()9) <9+5-F9)*,<1+1.:,-2)

1.F+1) -3) :d2;2-3).1:133\+,9)0-+-) <9+5-+) F1:,*93)1)`+6893C) /9+) <9+5-+15)-) ,.F1+<-:1)1.F+1) :d2;2-3)

-*W-:1.F13() 0-+1*13) :12;2-+13) 5;,F-3) K1413) *131501.M-5) 0-0d,3) ,509+F-.F13) .-) :95;.,:-789)

,.F1+:12;2-+)1)012-)29:-2,4-789)01+,<d+,:-)*131501.M-5);5)0-012),509+F-.F1).-3),.F1+-7X13)02-.F-H

5,:+9+6-.,3593(),.:2;,.*9)+13093F-3)*1)*1<13-):9.F+-)-61.F13)0-F96=.,:93)Eb163F+-)BTATGC))

Z959)-)<9+5-789)*9)-1+=.>;,5-)61+-251.F1)d),.*;4,*-)09+)-2-6-51.F9)E13F+1331G()-26;.3)

F+-V-2M93) ,.*,:-5);5)-;51.F9).-3)0+9F1?.-3) +12-:,9.-*-3)-)*91.7-3i+13,3F=.:,-)1)-)59*,<,:-789)

*-)0-+1*1) :12;2-+) :959)~HA(O) 62;:-.-31() A(Q) 62;:-.9)0+9F1?.-) 3,.F-31() ~H62;:93,*-31) 1)51F,9.,.-)

3,.F-31()1)>;1)13F-3)1.4,5-3)09*15)13F-+)+12-:,9.-*-3):95)-3)59*,<,:-7X13)13F+;F;+-,3)9V31+K-*-3)

15)+-?413)*1)39W-)Eb9.6)1F)-2C)BTATGC)e.F+1F-.F9()15)13F;*93):95)02-.F-3)>;1)0933;15):-0-:,*-*1)

*1) <9+5-789) *1) -1+=.>;,5-) E:959) 9)5,2M9G() 389) 9V31+K-*-3) -2F1+-7X13) .-3) 01:F,.-3) *-) 0-+1*1)

:12;2-+)*;+-.F1)-)*13,.F16+-789)*9):`+F1c) +-*,:;2-+)15);5-)<-31)5;,F9)0+1:9:1)*-)59+F1):12;2-+)

E!;.-�-+*1.-)1F)-2C)BTTAVGC))

#2d5) *-3)5;*-.7-3) .9) :9.F1@*9) *1) 01:F,.-3) E!;.-�-+*1.-) 1F) -2C) BTTAVG()5;,F9) 09;:9)

F15)3,*9)*13:+,F9)39V+1)-3)62,:93,2)M,*+92-313)*1)0-+1*1):12;2-+C)#26;.3)13F;*93)<9:-5).-)+16;2-789)

5921:;2-+)*-)<9+5-789)*9)-1+=.>;,5-)1)-),*1.F,*-*1)*-3)1.4,5-3)1.K92K,*-3).9)0+9:1339)*1)/Z%)

E';VV-,-M)1)'-:M3)BTTOGC)$9)F+-V-2M9)*1)v1)1F)-2C)EAjj|-G)<9,)+109+F-*9)>;1)-)-F,K,*-*1)*-)1.4,5-)

:12;2-31) d) -;51.F-*-).-) +16,89) -0,:-2) -.F13)*1)>;-2>;1+) 3,.-2) *1) -.9+5-2,*-*1)5,:+93:`0,:-)*-)

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Page 20: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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Page 21: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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Page 89: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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Page 98: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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0H,MK42KH7F0F2I4N32J/.234K/4210K42KN322I4KNHGM/7347/NJM/,3.32H2

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!D #$$$%@'$('9(D+,93N7H.L0HZ34JN/K3H.4\L"45#65N8_1421KHQ142FO2J+4

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17H./41GHP4H7J/NKI4-%4K/49/0,H4Q32HG03L73PH1K3P4KN1.2J/NKI49S;LXL]FG/234OH/28.KM3KHG4JN/G322I4,0FGFN/./R801.473K1O/0HG4JN/G322I4.HKN1K34KN1.2J/NKI4N3,F01KH/.4/]4N//K4P3Q30/J73.KI4N32J/.234K/4G/0PI4N32J/.234K/43.P/J0127HG4

N3KHGF0F742KN322I4N32J/.234K/4.HKN1K34IN32J/.234K/4210K42KN32I44R801.4OH/28.KM3KHG4JN/G322

1J/J012KI3RKN1G300F01N471KNHRI.FG03F2IJ01.KLK8J34G3004a100IJ01.KLK8J34Q1GF/03IJ012714

737ON1.3IJ0127/P3271

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1GKHQHK8I/R101K34/RHP12341GKHQHK8

!\ #$$$%&'((!9('+,b31K42M/GZ4G/,.1K34D(4ZS14JN/K3H.4!45#6#/01.F7408G/J3N2HGF749:6b#$L!4;-6!4#<6'4=>:?;%5@A

'+CC! (+\CE '+C\* (+\*\ (+!*(

1FK/JM1,8I4P3]3.234N32J/.234K/4O1GK3NHF7I4P3]3.234N32J/.234K/4]F.,F2I43.P/J0127HG4N3KHGF0F74F.]/0P3P4JN/K3H.4N32J/.23I4-%L122/GH1K3P4JN/K3H.4G1K1O/0HG4

JN/G322I4,0FG/.3/,3.32H2I4M31K41GG0H71KH/.4IM8J3N/27/KHG4210H.HK84N32J/.234I.3,1KHQ34N3,F01KH/.4/]4233P4,3N7H.1KH/.I4JN/K3H.4N3K3.KH/.4H.4-%40F73.I4JN/K3H.4K1N,3KH.,4K/4GM0/N/J012KI4N32J/.234K/4G1P7HF74H/.I4N32J/.234K/4G/0PI4N32J/.234K/4MH,M40H,MK4H.K3.2HK8I4N32J/.234K/4M8PN/,3.4J3N/RHP3I4N32J/.234K/4QHNF2I4N32J/.234K/4a1K3N4P3JNHQ1KH/.I4210HG80HG41GHP4OH/28.KM3KHG4JN/G322I4282K37HG41GcFHN3P4N32H2K1.G3

1J/J012KI4G3004a100I4GM0/N/J012K43.Q30/J3I4GH2L9/0,H4.3Ka/NZI4

G8K/2/0HG4NHO/2/73I43.P/J0127HG4N3KHGF0F740F73.I49/0,H41JJ1N1KF24IH.K3,N104K/4737ON1.3I473PH1K/N4G/7J03RI47HK/GM/.PNH10471KNHRI4.FG031N471KNHRI4.FG03/0F2I4J012714

737ON1.3I4J0127/P3271I4KM801Z/HPIQ1GF/01N4737ON1.3

T@;4OH.PH.,IdS-X423cF3.G34OH.PH.,IJN/K31234OH.PH.,IN3G3JK/N4

1GKHQHK8IFOHcFHKH.4JN/K3H.40H,1234OH.PH.,I_H.G4H/.4OH.PH.,

M31K42M/GZ4D(ZS14JN/K3H.4'^\Ib#;T'e\

!B #$fXX%'(''-(C+,b31K42M/GZ4JN/K3H.4\'L'45#65N8_1421KHQ142FO2J+4`1J/.HG149:6b#;\'L'4;-6!4#<6!4=>:?;%5@A

'+!!B (+BCB '+!"C '+((B (+\"*

G10GHF7L73PH1K3P42H,.10H.,I4G300F01N4N32J/.234K/4G10GHF74H/.I4P3L3KH/01KH/.I4P3]3.234N32J/.234K/4O1GK3NHF7Y4H.G/7J1KHO034H.K3N1GKH/.I4M31K41GG0H71KH/.I4/N,1.HG4

2FO2K1.G34OH/28.KM3KHG4JN/G322I4JN/K3H.423GN3KH/.I4JN/K3H.42K1OH0H_1KH/.I4N3,F01KH/.4/]473NH2K374,N/aKMI4N3,F01KH/.4/]473NH2K3742KNFGKFN104/N,1.H_1KH/.I4N32J/.234K/4G1P7HF74H/.I4N32J/.234K/4GM0/N1K3IN32J/.234K/4G/0PI4N32J/.234K/4M8PN/,3.4

J3N/RHP3I4N32J/.234K/473GM1.HG1042KH7F0F2I4N32J/.234K/4210K42KN322I4N32J/.234K/4a1K3N4P3JNHQ1KH/.

1J/J012KIG3004a100IGM0/N/J012K43.Q30/J3IGM0/N/J012K4

2KN/71IG8K/2/0I3.P/J0127HG4N3KHGF0F7I9/0,H4

1JJ1N1KF2I7HK/GM/.PNH/.I.FG03/0F2IJ012714

737ON1.3IJ0127/P3271IQ1GF/01N4737ON1.3

T@;4OH.PH.,IT@;12341GKHQHK8IF.]/0P3P4JN/K3H.4

OH.PH.,

MKJ9I7/03GF01N4GM1J3N/.34bKJ9

C( #$$$X%!$(!)'(+,b31K42M/GZ4JN/K3H.4\'LC45#65N8_1421KHQ142FO2J+4`1J/.HG149:6b#;\'LC4;-6!4#<6!4=>:?;%5@A

E+BD! (+\CB '+*'C (+*BE +,++(

G10GHF7L73PH1K3P42H,.10H.,I4G300F01N4N32J/.234K/4G10GHF74H/.I4P3L3KH/01KH/.I4P3]3.234N32J/.234K/4O1GK3NHF7Y4H.G/7J1KHO034H.K3N1GKH/.I4M31K41GG0H71KH/.I4/N,1.HG4

2FO2K1.G34OH/28.KM3KHG4JN/G322I4JN/K3H.423GN3KH/.I4JN/K3H.42K1OH0H_1KH/.I4N3,F01KH/.4/]473NH2K374,N/aKMI4N3,F01KH/.4/]473NH2K3742KNFGKFN104/N,1.H_1KH/.I4N32J/.234K/4G1P7HF74H/.I4N32J/.234K/4GM0/N1K3IN32J/.234K/4G/0PI4N32J/.234K/4M8PN/,3.4

J3N/RHP3I4N32J/.234K/473GM1.HG1042KH7F0F2I4N32J/.234K/4210K42KN322I4N32J/.234K/4a1K3N4P3JNHQ1KH/.

1J/J012KIG3004a100IGM0/N/J012K43.Q30/J3IGM0/N/J012K4

2KN/71IG8K/2/0I3.P/J0127HG4N3KHGF0F7I9/0,H4

1JJ1N1KF2I7HK/GM/.PNH/.I.FG03/0F2IJ012714

737ON1.3IJ0127/P3271IQ1GF/01N4737ON1.3

T@;4OH.PH.,IT@;12341GKHQHK8IF.]/0P3P4JN/K3H.4

OH.PH.,

MKJ9I7/03GF01N4GM1J3N/.34bKJ9

C' #$g9%@C(D")(D+, f127/.1K3=>:?;%5@A (+B!C '+("* (+B"( (+D'* +,+)'

`127/.HG41GHP4OH/28.KM3KHG4JN/G322I4031]423.32G3.G3I4/RHP1KH/.LN3PFGKH/.4JN/G322I4/R80HJH.473K1O/0HG4JN/G3224IN32J/.234K/4G1P7HF74H/.I4N32J/.234K/4]F.,F2IN32J/.234

K/4M/N7/.342KH7F0F2I4N32J/.234K/4/_/.3I4N32J/.234K/4210HG80HG41GHP42KH7F0F2I4N32J/.234K/4a/F.PH.,I42H,.104KN1.2PFGKH/.

G8K/2/0IJ3N/RH2/73'!L/R/JM8K/PH3./1K34

N3PFGK12341GKHQHK8I)[:4OH.PH.,

'!L/R/JM8K/PH3./HG41GHP4N3PFGK1234

U-$V'+C+'+E!WI-'+C+'+E!

C! #$fX%&'(!ET(E+, f127/.1K3=>:?;%5@A (+CD! (+\E' '+(BB '+'(* +,+++

1.KM3N4P3MH2G3.G3I4G300F01N417H./41GHP473K1O/0HG4JN/G3224IG300F01N41N/71KHG4G/7J/F.P473K1O/0HG4JN/G322I4G/3._873473K1O/0HG4JN/G322I4P3]3.234N32J/.234K/4O1GK3NHF7Y4H.G/7J1KHO034H.K3N1GKH/.I4,N/aKMI4`127/.HG41GHP4OH/28.KM3KHG4JN/G3224I01K3N104N//K4]/N71KH/.4I0HJHP4OH/28.KM3KHG4JN/G322I40HJHP4/RHP1KH/.I40/G10H_1KH/.I4

737ON1.34PH212237O08I4.3,1KHQ34N3,F01KH/.4/]4P3]3.234N32J/.234K/4H.23GKI4/N,1.HG4G8G0HG4G/7J/F.P473K1O/0HG4JN/G322I4N32J/.234K/41O2GH2HG41GHP42KH7F0F2I4N32J/.234K/4]F.,F2IN32J/.234K/4M3NOHQ/N3I4N32J/.234K/4`127/.HG41GHP42KH7F0F2I4N32J/.234K/4

/_/.3I4N32J/.234K/4a1K3N4P3JNHQ1KH/.4IN32J/.234K/4a/F.PH.,

GM0/N/J012K43.Q30/J3IGM0/N/J012K42KN/71IGM0/N/J012K4KM801Z/HP4737ON1.3IJ012714737ON1.3

HN/.4H/.4OH.PH.,I0H./031K34'C#L0HJ/R8,3.12341GKHQHK8I0H./031K34B#L0HJ/R8,3.12341GKHQHK8IJN/K3H.4

OH.PH.,

0H./031K34B#L0HJ/R8,3.1234

U-$V'+'C+''+"\WIX5h'e"

CC #$$$$XE('CS(B+, `127/.HG41GHP=>:?;%5@A (+BCD '+(\( '+!'( (+BCB (+"DB

1O2GH2HG41GHP473PH1K3P42H,.10H.,4J1KMa18I4M8J3N/27/KHG4210H.HK84N32J/.234IH.P/031G3KHG41GHP4OH/28.KM3KHG4JN/G322I4`127/.HG41GHP4OH/28.KM3KHG4JN/G3224I`127/.HG41GHP473PH1K3P42H,.10H.,4J1KMa18I4031]423.32G3.G3I4/RHP1KH/.LN3PFGKH/.4JN/G322I4/R80HJH.473K1O/0HG4JN/G322I4N32J/.234K/41FRH.42KH7F0F2I4N32J/.234K/4G1P7HF74H/.I4N32J/.234K/43KM803.342KH7F0F2I4N32J/.234K/4]F.,F2IN32J/.234K/4/_/.3I4N32J/.234K/4

210HG80HG41GHP42KH7F0F2IN32J/.234K/4a1K3N4P3JNHQ1KH/.I4N32J/.234K/4a/F.PH.,I4KN8JK/JM1.4G1K1O/0HG4JN/G322

G8K/2/0IJ3N/RH2/73'!L/R/JM8K/PH3./1K34

N3PFGK12341GKHQHK8I)[:4OH.PH.,

'!L/R/JM8K/PH3./HG41GHP4N3PFGK1234

U-$V'+C+'+E!WI-'+C+'+E!

CE #$$$%@'(('-('+, `127/.HG41GHP=>:?;%5@A (+D"* '+'(D '+E'C '+!E* +,+++

1.KM3N4P3MH2G3.G3I4G300F01N417H./41GHP473K1O/0HG4JN/G3224IG300F01N41N/71KHG4G/7J/F.P473K1O/0HG4JN/G322I4G/3._873473K1O/0HG4JN/G322I4P3]3.234N32J/.234K/4O1GK3NHF7Y4H.G/7J1KHO034H.K3N1GKH/.I4,N/aKMI4`127/.HG41GHP4OH/28.KM3KHG4JN/G3224I01K3N104N//K4]/N71KH/.4I0HJHP4OH/28.KM3KHG4JN/G322I40HJHP4/RHP1KH/.I40/G10H_1KH/.I4

737ON1.34PH212237O08I4.3,1KHQ34N3,F01KH/.4/]4P3]3.234N32J/.234K/4H.23GKI4/N,1.HG4G8G0HG4G/7J/F.P473K1O/0HG4JN/G322I4N32J/.234K/41O2GH2HG41GHP42KH7F0F2I4N32J/.234K/4]F.,F2IN32J/.234K/4M3NOHQ/N3I4N32J/.234K/4`127/.HG41GHP42KH7F0F2I4N32J/.234K/4

/_/.3I4N32J/.234K/4a1K3N4P3JNHQ1KH/.4IN32J/.234K/4a/F.PH.,

GM0/N/J012K43.Q30/J3IGM0/N/J012K42KN/71IGM0/N/J012K4KM801Z/HP4

737ON1.3

HN/.4H/.4OH.PH.,I0H./031K34'C#L0HJ/R8,3.12341GKHQHK8I0H./031K34B#L0HJ/R8,3.12341GKHQHK8IJN/K3H.4

OH.PH.,

0H./031K34B#L0HJ/R8,3.1234

U-$V'+'C+''+"\WIX5h'e"

GO função molecular -./012345"!$60$7"##898$:$8;<"!= >02"$63$?10="@.3A$"!?B<-"$CD3.<0$6"$6360!E F"6-3.3$63$G3HI0$60!$!"#2".=0! 9>JK9$!"#$%&'

LJ$?10<"!!0$D-0MN#-<0 LJ$<02?0.".="$<"M;M31

Page 109: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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2G2LH04G7851011G2LJOI74G.0E5TG7HG.KJ0E53473P78GPG41G7HG1M341G7E37541G6EI73H.045676O6J567632M3K4872GE0L3J567MH36G11I7HG1M341G7E370L1651567065871E52KJK1I7HG1M341G7E37PK4.K1IHG1M341G7E37FGHL5T3HGI7HG1M341G7E37/01234567065871E52KJK1I7HG1M341G7E37

3N34GI7HG1M341G7E37R0EGH78GMH5T0E5347IHG1M341G7E37R3K4854.

6FJ3H3MJ01E7G4TGJ3MGI6FJ3H3MJ01E71EH320I6FJ3H3MJ01E7EFOJ0U35872G2LH04GIMJ012072G2LH04G

5H3475347L54854.IJ543JG0EG7)!#VJ5M3SO.G401G706E5T5EOIJ543JG0EG7@#VJ5M3SO.G401G706E5T5EOIMH3EG547

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60E0L3J567MH36G11I7HG1M341G7E3760825K27534I7HG1M341G7E37G483MJ012567HGE56KJK271EHG11I7HG1M341G7E37FG0EI7HG1M341G7E3710JE71EHG11IKL5cK5E54V8GMG48G4E7MH3EG547

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Page 112: PAPEL DAS ENZIMAS DE DEGRADAÇÃO DA PAREDE CELULAR

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211

Abstract Cell wall hydrolysis is one of the key processes needed for develop-ment of the technology for second-generation (2G) bioethanol production. Thus, finding and characterizing enzymes that can deal with the complexity of the walls has been the main focus of research. As a result, data on pretreatments of many kinds and performances of enzyme cocktails containing glycosyl hydrolases from microorganisms are becoming quickly available. Here we propose that the effi-ciency of the 2G process could be increased even further by acquiring control of mechanisms that plants themselves use to degrade their own walls, so that wall loosening provoked by such processes would decrease the energy demand for pre-treatments and facilitate hydrolysis. The examined in this chapter are plant-micro-organism interaction, cell wall storage mobilization, fruit ripening, abscission, and aerenchyma formation. These systems are seen as having in common the use of modules that are coupled sequentially in order to lead to cell wall modification, including hydrolysis, for performance of different biological functions. These modules are (1) target cells perception of a message from the hormonal balance, (2) cell separation, (3) cell expansion, (4) programmed cell death, (5) hemicellu-lose-cellulose relaxation/hydrolysis and (6) cellulose hydrolysis. We propose that the use of synthetic biology to transform bioenergy feedstocks could be a route to increase the efficiency of 2G processes.

13.1 Introduction

The main bioethanol producers in the world are the US and Brazil, the former pro-ducing it from maize starch and the second from sucrose from sugarcane culms. In order to increase bioethanol production without increasing acreage, second

Chapter 13Using Natural Plant Cell Wall Degradation Mechanisms to Improve Second Generation Bioethanol

Adriana Grandis, Amanda P. de Souza, Eveline Q. P. Tavares and Marcos S. Buckeridge

M. C. McCann et al. (eds.), Plants and BioEnergy, Advances in Plant Biology 4, DOI: 10.1007/978-1-4614-9329-7_13, © Springer Science+Business Media New York 2014

A. Grandis · A. P. de Souza · E. Q. P. Tavares · M. S. Buckeridge (!) Laboratory of Plant Physiological Ecology, Department of Botany, Institute of Biosciences, University of São Paulo, São Paulo, Brazile-mail: [email protected]

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212 A. Grandis et al.

generation (2G) bioethanol technologies could be used. These technologies con-sist in the production of free fermentable sugars from cell walls. To access these sugars it is of great importance the production of basic scientific information about what the cell walls are like and how enzymes attack each polymer as well as the entire composite.

The production of bioethanol by means of 2G technologies involves the use of strategies that should modify the organization of cell wall architecture. The so-called 2G route involves biomass pretreatments in which polysaccharides would become available for enzymatic hydrolysis by application of physical and/or chemical treatments (Soccol et al. 2010; Dos Santos et al. 2011).

Moreover, due to the complex nature of the plant cell wall, degradation of lig-nocellulosic biomass requires the use of different classes of enzymes that have to be used in high proportions in order to produce enough free sugars for efficient fermentation (Verma et al. 2010). The production of 2G bioethanol requires ca. 11 millions of filter paper units (FPU) (i.e. 19 kg) to produce 84 L of bioethanol (Himmel et al. 1997, 1999) or 15–25 kg of cellulase per ton of biomass (Carroll and Somerville 2009; Taylor et al. 2008). Furthermore, due to the different compo-sitions of the cell wall polymers, it would still be necessary to prospect for individ-ually different classes of enzymes for characterization and subsequently combine them into cocktails that should be specific for every type of biomass, including the pretreated ones. Thus, one of the first challenges to be faced in order to turn 2G bioethanol viable commercially is to develop efficient systems of enzymes for biomass degradation (Verma et al. 2010). Due to the high costs and limited capac-ity for enzyme production, new ways to make biomass available for fermentation could be helpful, since they could hopefully lead to the use of lower proportions of enzymes in the process.

The proposed way to couple pretreatment and hydrolysis has been the use of enzyme cocktails on cell walls of pretreated materials (Fig. 13.1). Such enzyme cocktails are usually artificial assemblies of extracellular proteins produced by microorganisms (Balat 2011). In fact, many initiatives exist to use microorgan-isms as producers of heterologous glycosyl hydrolases so that enzyme engineering could be used to improve their action.

The sources of enzymes chosen by the scientific community varies, but with the advent of metagenomic techniques, microorganisms and animal genes that encode for glycosyl hydrolases became the main focus for enzyme search in the hope to find new targets that could be somehow more efficient to hydrolyze plant cell walls. This strategy relies mainly on the idea that microorganisms that per-form composting of plant residues in the environment and some animal digestive systems are capable to hydrolyze the plant cell walls (Gómez et al. 2008).

Some initiatives exist in which enzymes coding genes from microorganisms have been cloned into plants in order to try to loosen walls and turn them more amenable to hydrolysis (Kaida et al. 2009; Xin et al. 2011). Although expres-sion of the enzymes in vivo have been demonstrated, there is no evidence that these biomasses would be more easily hydrolysed and/or pretreated than biomass derived from untransformed plants.

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Thus, even with the large effort of the scientific community world wide to improve pretreatment and hydrolysis processes, cell wall recalcitrance to hydroly-sis continues to be a barrier to turn the 2G processes economically viable (Himmel et al. 2007). A common mistake has been to attribute recalcitrance solely to the presence of lignin. However, this recalcitrance is also due to the enormous com-plexity of the architecture of the cell wall with its several polymers integrated in a composite that has been selected in a way to avoid enzyme attack and degradation.

One additional strategy to be used to improve the knowledge about 2G pro-cesses that has not been consistently thought of, or experimented with the same intensity, is the use of hydrolytic systems that plants themselves possess. Some of them can be quite efficient to degrade and/or change the architecture of cell walls and might lead to important additions to 2G processes, especially with the advent of the synthetic biology era, in which the capacity of changing biological systems (e.g. turning on/off entire biochemical pathways) is expect to be developed.

Similarly to the strategies currently being designed in which biological engi-neering is going to be used within microorganisms to produce more efficient enzymes, plant biotechnology could help not only by the introduction of genes of animals and/or microorganisms, but also by finding mechanisms of cell wall hydrolysis that already exist in nature. The main mechanisms of this kind known are: (1) plant-microorganism interaction; (2) seed cell wall storage mobilization; (3) fruit ripening, (4) abscission and (5) aerenchyma formation.

In this chapter, we will provide a review of aspects of some representative mechanisms that could be relevant to improve plants so that they would be better

Fig. 13.1 Schematic representation of the second-generation process of bioethanol production, highlighting the different types of pretreatments and hydrolysis

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prepared for pretreatment and hydrolysis. Here, this process will be named bio-logical pretreatment. The idea is that in the future, plants could have some of their hydrolytic processes redesigned to alter some features of the cell walls in the way of turning them more suitable for use in industry, without however loosing the perspective of the technologies current under development for 2G bioethanol production.

In order to understand how walls can be hydrolysed, some aspects of cell wall composition, structure and architecture will be introduced in the following section. The understanding of the cell wall complexity will allow the reader to appreciate how some natural systems in which walls are modified could be used for biotech-nology applications in bioethanol production.

13.2 Composition, Structure and Architecture of the Plant Cell Wall

The cell walls are composed of a mixture of polymers, mainly carbohydrates, pro-teins and secondary metabolites. These polymers interact through a mixture of covalent and non-covalent linkages to form a “supramolecular complex” that is thought to be a multifunctional structure responsible for controlling the mechani-cal properties of the cell.

In order to elucidate plant cell wall structure and functions, wall scientists have followed some key steps mainly during the second half of the 20th Century. The first step was to set up methods for extraction, determination of composition and structure of wall components. This step was followed by the development of meth-ods to determine the structure (linkage types) and physico-chemical properties of these polymers. All this work culminated in the proposition of a succession of models that pictured a general assembly for the polymers in the wall (Keegstra et al. 1973; McCann and Roberts 1991; Carpita and Gibeaut 1993). Likewise, many aspects of the biochemistry of the wall were developed, culminating in the characterization of several glycosyl hydrolases (e.g. Fry 2004) that correlate with physiological events in plants, which are controlled by plant hormones and envi-ronmental cues. With the advance of cellular and molecular biological techniques, part of the wall research community invested in finding and correlating wall struc-ture with genes associated with its metabolism (e.g. Holland et al. 2000; Lima et al. 2001) and also succeeded to find probes (enzymes and monoclonal antibodies) capable to localise wall components in the cell (e.g. Willats et al. 2001; Pattathil et al. 2010).

McCann and Roberts (1991) coined the term “architecture” of the cell wall in a reference to the existence of three different independent domains in the wall: pec-tin, cellulose-hemicellulose and proteins. This view implies that the three domains might interact and this sole interaction would have emergent properties. Carpita and Gibeaut (1993) showed that at the level of higher plants, the wall structure has different polysaccharides (e.g. arabinoxylan, !-glucan, xyloglucan and mannan)

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that may play similar functions. Thus, the “architectural” model of the wall points out that cell wall structure is degenerated and that the domains are chang-ing during evolution, i.e. monocots and ferns changed their main hemicelluloses from xyloglucan to arabinoxylan and mannans respectively, with a concomitant decreased pectin domain (Buckeridge 2006; Sarkar et al. 2009).

In higher plants, cellulose, (1 ! 4)-!-D-glucan, is the most abundant biologi-cal material on earth and in dry wood correspond around of 40–50 % of dry mass. Hemicellulose is the second polysaccharide representing 25–35 % of dry mass and their composition is a mixture of various polymerised monosaccharides, such as xylose and arabinose or glucose, mannose and galactose whose proportions change according to the plant tissue and the species.

The principal hemicelluloses found in nature are xyloglucan, galactomannan or galactoglucomannan, !-glucan and glucuronoarabinoxylan (GAX). Xyloglucan is a major hemicellulose in the primary walls of flowering plants, all non-mono-cots and about one-half of monocot species. Some seeds store this polysaccha-ride in large amounts as a resource for the embryo after germination (Buckeridge et al. 2000a). Xyloglucans typically comprise (1 ! 4)-!-D-glucan back-bones, with three of every four glucose residues substituted with (1 ! 6)-"-D-xylopyranosides, and some of the xylose residues are further substituted with (1 ! 2)-!-D-galactopyranosides. The most widespread technique used for stud-ies on xyloglucan has been the digestion with a Trichoderma endo-!-glucanase (Anderson and Stone 1975; Tiné et al. 2006). This enzyme hydrolyses xyloglucan polymers only on unbranched glucosyl residues.

The cell walls from grasses (Poales), which are the main plants used in bio-energy, have a distinct architecture when compared with non-monocots and non-Commelinid monocots (Carpita and Gibeaut 1993). Besides the fact that low levels of pectin are present in their walls, grasses contain as their main hemicel-luloses the mixed-linked (1 ! 3), (1 ! 4)-!-d-glucan (!-glucan) and glucurono-arabinoxylan (GAX).

!-Glucan interlaced by GAX of low degree of substitution and glucomannans, is thought to be tightly associated with the cellulose microfibrils. GAX of higher degree of arabinosyl substitution and some glucomannan constitutes the major pore-determining interstitial material between the microfibrils (see below).

One of the principal enzymes involved in the depolymerization of the endosperm !-glucan is the (1 ! 3), (1 ! 4)-!-D-glucan endohydrolase, whose activity parallels that of the B. subtilis enzyme in that hydrolysis is restricted to (1 ! 4)-!-linked glucosyl residues adjacent to (1 ! 3)-!-linked residues on the non-reducing side (Parrish et al. 1960). Like the B. subtilis endohydrolase, hydrolysis of !-glucan with these enzymes yields mostly cellobiosyl- and cello-triosyl-(1 ! 3)-glucosyl oligosaccharides and smaller amounts of larger cellodex-trin-(1 ! 3)-glucose oligomers (Anderson and Stone 1975).

In GAX, the arabinose residues are attached at the O-3 positions along the (1 ! 4)-!-linked xylan backbone, and the glucuronic acids are attached to the O-2 positions (Carpita 1996). Arabinoxylans are widespread in the walls of all flower-ing plants, but in nongramineous species the polymer is of much lower abundance,

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and the arabinose residues are attached mostly at the O-2 rather than the O-3 of the xylosyl units.

A highly substituted GAX (HS-GAX), with six of seven xylosyl branched units is associated with the maximum growth rate of coleoptiles of the grass species Lolium multiflorum and a sequence-dependent xylanase was found that requires branching with GlcA to cleave the neighboring (1 ! 4)-!-D-xylosyl linkage (Nishitani and Nevins 1991). When maize GAXs are depleted of arabinosyl units by mild-acid hydrolysis, this endo-"-D-xylanase releases a homogeneous group of deca- or undecamers of glucuronoxylan. The xylosyl units are also substituted with acetyl groups at the positions O-2 and O-3 and some methyl groups have been detected (Carpita 1996).

Porosity of the GAX domain could be determined by the extent of removal of the appendant units. Some highly substituted GAX remain intercalated in the small amount of pectins that are also found in the primary wall. Because "-glucans are thought to be tightly associated with individual cellulose microfibrils (Carpita et al. 2001), GAX might be the main molecule in the interstitial space. However, Buckeridge and Crivellari (unpublished) have recently performed experiments with sugarcane walls that did not support this hypothesis. When isolated walls (alcohol insoluble residues) of leaves and culms of sugarcane were treated with lichenase (a glycosyl hydrolase that attacks specifically "-glucan), most of the pol-ymer present in the walls was hydrolyzed and, at the same time, its retrieval did not interfere in the action of endo "-xylanase on the wall. Also, De Souza et al. (2013) observed that "-glucans in sugarcane walls are lightly associated with cel-lulose and/or the cellulose-hemicellulose matrix in the wall. These data suggests that in sugarcane, although "-glucan is present in some association with other pol-ymers in the wall, they seem to be free for enzyme hydrolysis whereas GAXs are likely to be strongly bound to other polymers, probably cellulose.

Another distinctive feature of Poales and their relatives is the enrichment of aromatic substances in nonlignified walls (Carpita 1996). A large portion of the aromatic substances is esters of hydroxycinnamates (Harris and Hartley 1980). The GAXs are cross-linked in walls by both esterified and etherified hydroxycin-namates and by other phenolic substances, as ferulic acid (Iiyama et al. 1994).

On the basis of the knowledge of the structure of the cell wall polysaccharides, the mode of action of the glycosyl hydrolases and also the genes that encode for these enzymes, it is now possible to design routes that can deal with the complex hydrolysis mechanisms of either the less complex seed polysaccharide degrad-ing systems or even to access more complex wall systems such as the ones from grasses. The latter is not an easy task as relatively little knowledge exists about the composition, structure and architecture of the wall of most bioenergy feedstocks.

The cell walls of one of the most important bioethanol feedstocks, sugar-cane, has been recently analysed by De Souza et al. (2013), who demonstrated that hemicelluloses are quantitatively the main components of the cell walls. Regarding carbohydrates alone, whereas hemicelluloses account for ca. 60 % of the walls, cellulose makes around 30 %, and pectins about 10 %. Hemicelluloses are of three types: arabinoxylan, "-glucan and xyloglucan, the former being about

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50 % of the hemicelluloses with the other two types sharing the rest in equal proportion. These authors highlighted the elevated complexity of cell walls and proposed a model for enzyme hydrolysis in which phenylpropanoids in the wall would have to be retrieved first in order to open the way for carbohydrate specific enzymes, pectins being the first to be attacked, followed by the members of the cellulose-hemicellulose matrix.

In summary, cell wall hydrolysis for bioenergy production purposes can be thought of as a procedure capable to disassemble cell wall architecture so that individual polysaccharides would be available to hydrolases. The goal of the pro-cess is the production of fermentable monosaccharides. On the other hand, even if the cell wall architecture can be dismantled, the polysaccharides released would still offer the challenge related to their branching patterns, which are different for every polysaccharide, resulting in a rather large diversity of combinations that are named fine structures.

Although the structure of cell wall polymers have been studied for several spe-cies, less attention is given to fine structure studies. Since the fine structure of polysaccharides could involve a formation of a glycomic code and interfere in hydrolysis efficiency (De Souza and Buckeridge, unpublished), the understanding of branching patterns and its combinations could be an important issue for 2nd generation processes (see topic 13.4).

In order to illustrate how natural systems could be helpful in bioenergy sec-tor, in the following sections we will discuss five major systems existent in nature, which include cell walls modifications by enzymes produced for specific situa-tions. It must be highlighted here that these items do not intend to review every biological process exhaustively, but only give key information that are relevant for the mechanisms of cell wall modifications (including hydrolysis) that could be used to transform plants and make them more suitable as bioenergy crops.

13.3 Cell Wall Degradation During Plant-Microorganism Interaction

When a pathogen invades a plant tissue, a biochemical conflict takes place, in which host and invader will produce a series of substances, the former to defend itself and the latter to penetrate in the cells (Garcia-Brugger et al. 2006). Among the strategies used by both organisms there are cell wall hydrolases (Walton 1994). In this way, the process of plant-microorganism interaction offers an opportunity to find ways to efficiently disassemble the plant cell wall. Both, saprophytic and pathogenic microorganisms produce extracellular enzymes that can degrade plant polysaccharides. These microorganisms, including bacteria and fungi, and also invertebrates as nematodes can digest cell wall polymers in order to obtain sug-ars and energy for their growth as well as to penetrate and colonize the cells in a plant tissue (Walton 1994; Annis and Goodwin 1997). In most cases, plants will respond with a reaction named hypersensitivity, which includes programmed cell

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death and with that can restrict the infectious region to a small area surrounding the infection. However, in these cases, pathogens can use this opportunity to feed on dead cells and gain nutrients and with that they may become even more inva-sive (Greenberg and Yao 2004).

The first class of cell wall hydrolases produced by the pathogen is the pecti-nases (Tomassini et al. 2009). They are thought to loosen the tissue through diges-tion of the middle lamella and consequently kill the plant cells that they attack (Cooper 1983). The pectinases known to be involved in fungal and bacterial inva-sion are endo- and exo-polygalacturonases, pectate-lyases, and pectin methylester-ases (Walton 1994; Lagaert et al. 2009). It is also believed that even saprophytes need to use cell wall degradation strategies in order to establish interaction with the plant and for that they also produce pectinases (e.g. Marques et al. 2006).

Pectins are polysaccharides that control cell wall porosity. Some studies dem-onstrated that the porosity of the substrate walls is the main limiting factor in the enzymatic hydrolysis of lignocellulosic biomass (Chandra et al. 2007). Thus, the knowledge about how microorganisms use the pectinases to invade the plant could be a way to improve the biomass hydrolysis to 2G bioethanol.

Futhermore, several other enzymes that are able to hydrolyse cell walls can be produced by microorganisms. Recently, the need for enzymes to be used in the production of bioethanol from biomass led many groups to produce a large amount of scientific information about glycosyl hydrolases from microorgan-isms. The isolation and characterization of enzymes as well as cloning their genes, open a way to engineering those derived from microorganisms, i.e. modify them to perform hydrolysis more efficiently (Serpa and Polikarpov 2011; Ward 2011). Then, new cocktails of enzymes could be designed that would efficiently produce free fermentable sugars for bioethanol production. A review of the enzymes from microorganisms used for bioenergy has been recently published (Polizeli et al. 2011).

13.4 Seed Cell Wall Storage Mobilization

Cell Wall Storage Polysaccharides (CWSP) are found as the principal storage compound in seeds of many taxonomically important groups of plants. These groups developed extremely efficient biochemical mechanisms to disassemble cell walls and use the products of hydrolysis for growth. During evolution, several dif-ferent groups of polymers have been selected as reserves (e.g. mannans, galacto-mannans, galactoglucomannans, arabinogalactans and xyloglucans) (Buckeridge et al. 2000a, 2000b; Buckeridge 2010). The selective pressures that ended up pro-ducing CWSP were directed towards an increase their proportion by introducing a storage cell wall between already existent primary walls (Tiné, Braga, Pattathil, Hahn and Buckeridge, unpublished).

The CWSP have different chemical structures that are related with different biological functions. Mannans and galacto- and glucomannans that are present in palm and coffee seeds and lettuce and tomato respectively are insoluble in water

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and display strong intermolecular interaction. Because of these characteristics, their biological function is usually associated with conferring hardness to plant tis-sues. Endo-!-mannanase is the principal enzyme involved in mannan hydrolysis in all species studied and several genes have been cloned whose expression is spe-cific to the seed (Bewley et al. 1997; Lisboa et al. 2006; Gong and Bewley 2007).

In legumes, the main function of the endospermic cell walls appear to be stor-age, with the yield of galactomannan reaching more than 30 % of the seed dry weight in many species (Buckeridge et al. 2000a). These walls are thickened with galactomannan and in certain cases (e.g. Trigonella foenum-graecum and Schyzolobium parayba) the protoplasm disappears, giving place to the storage wall. In these cases, the endosperm is non-living and degradation is performed by three enzymes ("-galactosidase, endo-!-mannanase and exo-!-mannosidase) made in the aleurone layer (Reid 1971; Buckeridge and Dietrich 1996).

Differently from being stored in the endosperm, arabinogalactan and xyloglu-cans are stored in cotyledons. This is extremely relevant because the cotyledon is, evolutionarily speaking, an adapted leaf and the integration of the metabolism of these polysaccharides with the entire plant is likely to preserve several features of the leaf cell walls.

Pectin polymers are also found as cell wall storage polysaccharides, notably arabinogalactan (Buckeridge et al. 2000b). The cotyledons of lupins (especially Lupinus angustifolius) have been used as a model to study mobilization of this polymer.

Lupinus angustifoluis seeds accumulate high proportions of galactan (Buckeridge and Reid 1994). These authors showed that an exo-!-(1,4)-galactanase purified from L. angustifolius seed acts specifically on this type of galactan. This enzyme is analogous to galactanases found in fruits (TBG4 in tomato for example—see below) where it is thought to play a major function in the changes of porosity in cell walls (Brummell 2006).

Of the CWSP known, xyloglucan seems to be the more complex in most senses. Its basic structure is similar to the primary wall xyloglucans. They have a backbone composed of (1 ! 4)-!-linked glucan (like cellulose) with regular branching with (1 ! 6)-"-linked xylosyl residues that can be branched further with (1 ! 2)-!-linked galactosyl residues. There is no case reported where fucose was detected in storage xyloglucan. Four enzymes responsible for storage xylo-glucan degradation have been detected, purified and characterized ("-xylosidase, !-galactosidase, !-glucosidase and xyloglucan-endo-!-glucanase, latter on renamed xyloglucan transglucosylase/hydrolase (XTH) (Buckeridge et al. 2000a; Buckeridge 2010).

In Hymenaea, Buckeridge et al. (1997) discovered that storage xyloglucan con-tains two types of constitutive blocks in the main chain, i.e., XXXG and XXXXG. Later on, Tiné et al. (2006) showed that these blocks are not combined randomly in the polymer molecules, which led to the proposition that storage xyloglucans might have a combination of branching patterns that forms a code that have to be decrypted by the enzymatic system in order to be hydrolysed (Buckeridge 2010). Indeed, the fact that branching of cell wall polymers might not be random has been also observed early on for galactomannans (Reid and Edwards 1995).

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As this kind of information is not available in most cases, research should be urgently directed to that in order to complement the efforts in the search for new enzymes as well as enzyme engineering.

13.5 Fruit Ripening

During ripening, the cells of fruits display changes in cell walls so that the final result will afford the release of seeds for dispersion. Several species have been adapted to human consumption and became sources of food, being nowadays extremely important in agriculture. That is the case of tomato, apple, pear, melon, papaya, grape to name but a few. Due to the economic importance of fruits, development and ripening have been intensively studied, with tomato being one of the most studied fruit regarding studies of cell wall changes related to texture (Brummell and Harpster 2001; Brummell 2006). Changes in fruit texture that occur during ripening in many cases lead to softening, this process being slightly different for each species (Harker et al. 1997; Brummell 2006). Nonetheless, in all cases the texture is related to changes in cell walls that are associated with the production of several hydrolases. Due to the fact that enzymes can act to hydro-lyse fruit cell wall polysaccharides and/or to allow structural changes that modify the mechanical properties of plant tissues, fruit systems can be considered as an opportunity to find mechanisms of cell wall degradation in plants that could help developing biotechnology for bioenergy.

In fruit systems, the most studied degradation process is the activity of endopo-lygalacturonases, pectin methylesterases and !-galactosidases on pectic matrix of the walls. Together with the detection of enzyme activities, changes in cell walls of tomato are consistent with a large decrease in uronic acid as well as even larger decreases in galactose in wall fractions (Ahmed and Labavitch 1980; Rose et al. 1998; Brummell and Harpster 2001; Brummell 2006).

It has been demonstrated that pectinases are associated with ripening and sof-tening through degradation of endopolygalacturonases present in the middle lamella. The action of this enzyme is thought to lead to cell separation, a phenom-enon that is linked to the subsequent softening of the fruits and change in their texture and taste (Fray and Grierson 1993; Barsan et al. 2010).

The xyloglucan-cellulose network has also been pointed out as an important target in ripening (Rose and Bennett 1999; Brummell 2006). These authors high-light the role of expansions and XTHs in the process and the former authors even compare the process of fruit ripening with the cell expansion system in hypoco-tyls. Brummell (2006) reviewed fruit ripening from the viewpoint of the wall changes, putting forward the idea that whereas pectin degradation varies among different fruit species (Brummell 2006), the xyloglucan-cellulose network seems to undertake changes that are much more consistent among species. However, the author also highlights the fact that, contrarily to what has been observed for pectins, xyloglucan small fragments have not been detected. Thus, the effects of expansin and enzymes on the xyloglucan-cellulose matrix seem to be more

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towards a relaxation of this matrix rather than hydrolysis. This seems to lead to cell wall swelling that happens concomitantly with pectin solubilization.

The increase in pore sizes, thought to be related to the attack of exo-!-galactanase in pectin matrix (Brummell 2006), probably opens the way for expan-sions, XTHs, cellulases, as well as other debranching enzymes, that will attack the cellulose-hemicellulose matrix of the wall, finally changing wall texture as a whole. Indeed, according to Carpita et al. (1979), pore sizes of the walls in plant tissues would be in the range of 35–40 Å, whereas the glycosyl hydrolases stoke radius [e.g. a 20 kDa protein would be approximately 2.5 nm according to Carroll and Somerville (2009)] is in general higher than the pore sizes, so that this is a clear limitation for penetration of enzymes in the cell wall matrix.

Although endo-!-glucanases have been found to increase during fruit ripen-ing, very little evidence has been produced that cellulose would be hydrolysed. However, it has been speculated that cellulases would be capable to act on the sur-face of microfibrils (Brummell 2006) without, however, any assignment of a func-tion for this event.

In spite of the fact of hydrolysis, in a broad sense, being the main process in course during fruit ripening, other steps such as cell separation and expansion, have been observed (Brummell 2006). On the other hand, differently from some other cell wall degradation processes such as aerenchyma formation and abscis-sion (see below), fruit ripening seems do not include programmed cell death.

From a general point of view, fruit ripening is a process in which walls are firstly attacked by pectinases that increases wall porosity, probably allowing the action of other enzymes like XTHs and endo-!-glucanases that will transform the cellulose-hemicellulose matrix, promoting swelling and increasing even more wall porosity. The extensins that bind to the cellulose-hemicellulose matrix of the wall also could participate of these processes, being probably the first proteins to interact with the wall matrix. Thus, the cell wall alteration plus a higher hydration capacity given by the presence of pectins with lower molecular weight modifies fruits softening.

All this knowledge is extremely useful for bioenergy purposes, as it makes pos-sible to understand details of how plants can manipulate their own walls. These events could be compared with the ones that occur during biomass pretreatment process used to achieve the 2G bioethanol. The possibility of applying such knowledge in bioenergy crops to produce similar effects in grass stems cell walls (e.g.) could lead to the establishment of a biological pretreatment that could be implemented using synthetic biology tools in the future.

13.6 Abscission

Abscission is a general term applied to the processes that takes place during the detachment of leaves, fruits, seeds and also cotyledons, lateral branches and many reproductive structures. In all cases studied, extremely precise biochemical path-ways lead to cell wall modification and breakdown in order to form a fracture line

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known as the abscission zone. The events that occur during abscission and dehis-cence processes are cleary related to cell wall modification, in a very similar way to the other ones reported in this chapter.

Roberts et al. (2002), highlight the facts that endopolygalacturonases and endo-!-glucanases are closely related to the processes of cell separation. In fruits like tomato, silencing of abscission-related polygalacturonases (e.g. TAPG1) led to a delay in abscission and increased the force needed to break the abscission zone in explants treated with ethylene (Jiang et al. 2008).

In the abscission zone, cortex cells of different plant tissues are thought to be positionally differentiated target cells that will receive the signal from hormones (ethylene, auxin, abscisic acid, gibberellin) in such a balance that the cross talk among these will lead to transdifferentiation and finally to abscission. McManus et al. (1998) demonstrated the existence of transdifferentiation in mature corti-cal cells of bean abscission tissues. According these authors, this means that cells from the cortex of some plant tissues have the flexibility to differentiate into func-tionally competent ethylene-responsive cells that exhibit a gene expression com-patible with an abscission cell.

Although not reported in most cases, cell wall degradation is thought to occur during abscission, since detection of activity of endo-!-glucanase has been reported for many abscission systems (Roberts et al. 2002). Also, accu-mulation of mRNA related with genes encoding Cel1 and Cel4 in tomato fruit and flower abscission zones have been reported (Gonzáles-Carranza et al. 1998).

The similarity of this process with the ones described above is remarkable. Abscission is widespread in plants, occurring in many different tissues of different organs in plants. It includes wall separation and expansion, which are associated with pectin degradation as well as modifications in the cellulose hemicellulose matrix. However, programmed cell death has not been reported for abscission pro-cesses, according to the literature revised for this chapter. Thus, there seems to be a great variety of combinations of action of hydrolases to perform cell separation and hydrolysis so that it could be used to design technologies associated to 2G bioethanol production.

13.7 Aerenchyma Formation

Aerenchyma comprises a series of interconnected gas chambers developing on parenchymatic tissue (cortex) of shoots and roots as a result of sequencial events, showing typical features of a programmed cell death process. Its formation is regarded as a response to oxygen shortage (Gunawardena et al. 2001a), exog-enous ethylene (Gunawardena et al. 2001b), nitrogen, phosphorus and/or sulphur starvation (He et al. 1994; Siyiannis et al. 2011), mechanical impedance (He et al. 1996b), oxidative stress (Steffens et al. 2011) and osmotic stress (Karahara et al. 2012).

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In opposition to the inducible aerenchyma, the constitutive formation is com-monly observed in aquatic species as Juncus effusus (Visser and Bögemann 2006) and Sagittaria lancifolia (Schussler and Longstreth 2000), and also in maize rela-tives (Mano et al. 2007) and in rice wetland species (Justin and Armstrong 1991), independently of environmental stimuli. In both cases, its development lies upon a cell separation process termed schizogeny (thought of as a cell separation process that does not include cell wall hydrolysis). Alternatively, for the lysogenic process (i.e. cell separation followed by cell wall hydrolysis) to take place, the cells tar-geted to form aerenchyma clearly undergo a programmed cell death. It is essential to mention that both processes rely on polysaccharide hydrolysis, whereas the cell separation in schizogenous aerenchyma formation depends on middle lamella deg-radation. As for the lysogenous process, cell wall modifications occur and for that a wide array of enzymes would be required.

Concerning cell wall modifications, most of the aerenchyma-related data is restricted to the lysigenic and inducible formation in maize roots. As reported for other cell degradation events, in this model the inductive role is played by ethylene (He et al. 1992, 1996a; Gunawardena et al. 2001a). One of the earliest signs of aerenchyma formation is the ethylene accumulation, followed by cell death and endo-!-glucanases activity (Kawase 1979; He et al. 1996a). Indeed, inhibitors of cell death or ethylene biosynthesis block the enhancement of endo-!-glucanases activity (He et al. 1994, 1996a), suggesting the coupling between cell wall degra-dation, ethylene signalling and programmed cell death.

Another potentially degradative enzyme associated with aerenchyma forma-tion is endo-!-xylanase. The rising activity levels of this enzyme were obtained in waterlogged maize roots, coinciding with the rising of ethylene levels (Bragina et al. 2001). Also concerning ethylene induction, the role of XTH, a putative cell wall loosening enzyme, was evaluated during aerenchyma formation. Transcripts of certain XTH accumulated in flooded maize roots forming aerenchyma, reaching its maximum within 12 h and remaining high for 144 h during flooding. On the other hand, the inhibition of ethylene synthesis blocked any rise of this transcript accumulation, while exogenous ethylene led to XTH transcript accumulation even under aerobic conditions (Saab and Sachs 1996). A transcriptional analysis cor-roborated the induction of XTH expression during the aerenchyma formation in maize roots, as well as for endo-!-glucanase (Rahji et al. 2011).

Even though the enzymatic or transcriptional approaches have been used to suggest the link between a set of glycosil hydrolases and aerenchyma formation, to our knowledge, data regarding cell wall modifications in situ have been reported only by Gunawardena et al. (2001b), who documented the distinctive levels of methyl esterification changing in the middle lamella in cross sections of the cortex of the maize root along aerenchyma formation. These authors observed that those changes initiated before ultrastructural modifications typical of cells undergoing programmed cell death that have been reported in a previous work (Gunawardena et al. 2001a). Indeed, rise in the expression of pectin methyl esterases, pectate lyase and an endopolygalacturonase have been observed during aerenchyma formation (Rahji et al. 2011). These data not only corroborate the existence of

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homogalacturonan modifications but also suggest that more severe changes on the cell wall structure and not only on the middle lamella can occur. We have observed a similar pattern of events in sugarcane roots, not only with the pectinases, but also with the observation of the rise in gene expression of several genes that encode hemicellulases (e.g. lichenase, endo-!-xylanase, XTH, !-glucosidase—unpub-lished results).

Gunawardena et al. (2001a) reported the formation of vesicles and vacuole rup-ture while the cell wall appeared intact. It is reasonable to hypothesize that the release of hydrolytic enzymes in this step could be responsible for the cell wall degradation (Bouranis et al. 2007).

The data produced that are related with aerenchyma formation to date indicates that this phenomenon is similar to the other processes discussed above in the sense that cell walls are modified, including cell separation, hemicellulose-cellulose matrix relaxation and perhaps some hydrolysis of cellulose. Thus, the knowlegde about aerenchyma formation can also be considered as a possible target for use in biotechnology that includes cell wall modifications for bioenergy purposes. Recently we have produced a consistent set of results describing events during the aerenchyma formation in sugarcane roots. We found that the events are essencially similar to maize (e.g.), with the exception that in sugarcane it is constitutively acti-vated. Our results (manuscripts in preparation) show very clearly that cell sepa-ration, programmed cell death, hemicellulose-cellulose changes and some attack to cellulose occurs in sugarcane. We are now searching for the transcription fac-tors responsible for regulating this processes aiming at possibly inducing them in culms so that this process might be used as a biological pretreatment in the future.

13.8 Cell Wall Modifications in Natural Systems in the Context of Bioenergy

As mentioned above, in order to establish 2G bioethanol production, the retrieval of energy from wall polymers would have to include cell wall modification. From the different systems discussed along this chapter, it is possible to think of com-bining some of them in order to help increasing the production of bioethanol from bioenergy feedstocks that are being studied for use in 2G processes.

In Table 13.1 the different modules that plants apparently use to transform their cell walls in order to perform different biological functions are shown in perspec-tive. Six events seem to be common to most of the natural processes that clearly involve modifications in cell walls. Some of these events have been investigated more deeply and others are implied, but not proven, as no direct experimental evi-dences have been produced for them yet. In Table 13.1, the events are placed in an order (from left to right) that they seem to occur in all systems. However, some steps are missing in some of the processes (e.g. cell separation in storage cell wall mobilization and programmed cell death in fruit ripening).

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22513 Using Natural Plant Cell Wall Degradation Mechanisms

The combination of events is consistent with a chain of processing steps that would follow a sequence of modifications in the walls that are apparently similar in the five processes reviewed in this chapter. The general idea is that in a par-enchymatic tissue, a given group of cells would be targeted to start the process (as seen in abscission according to Roberts et al. (2002) and also in aerenchyma formation). The following step would be cell separation that probably involves the action of endo-polygalacturonase(s) on the pectins of the middle lamella. This might be involved with the release of calcium that could be responsible for the signaling related to programmed cell death. The latter seems to occur in parallel with cell expansion and subsequent production and release glycosyl hydrolases that would attack hemicelluloses and cellulose. Every one of these events can be regarded as a module that can or not be used in the complete process that leads to partial or complete cell wall modification.

There are two possible ways to use this information for biotechnology pur-poses: (1) using them for production of transformed plants; or (2) express some of the hydrolase genes heterologously in order to use them as additions to enzyme cocktails already in use in industry.

For the first option, it should be possible to use the modern tools of synthetic biology in order to induce cell separation and expansion along with some modi-fication of the hemicellulose-cellulose matrix. In this case, plants would produce biomass with the characteristics expected to be analogous to a pretreated biomass material, i.e. materials that would be easily hydrolyzed when being processed in industry, especially during the hydrolysis process. For advancing the second option, some of the enzymes known to be present during hydrolysis processes in bioenergy feedstocks, such as sugarcane, miscanthus, maize, poplar, willow and others should be studied regarding their specificity. Some of these enzymes may be heterologously expressed for production and added to existent cocktails.

Table 13.1 Correlation among situations that include cell wall modifications and natural events in plant biology where these situations are key to biological function

YES = there is literature showing that it occurs in most (or all) systems studied; NO = not observed in the systems studied to date; YES/NO = observed only in some of the systems stud-ied to date; ? = not reported to occur to date

ProcessesTarget cells

Cell separation

Cell expansion

Programmed cell death

Hemicellulose hydrolysis

Cellulose hydrolysis

Plant microorganism interaction

? YES ? YES YES YES

Storage cell wall mobilization

? NO YES ? YES ?

Fruit ripening YES YES YES NO YES YES/NOAbscission YES YES YES ? YES YESAerenchyma

formationYES YES YES YES YES ?

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226 A. Grandis et al.

In both cases, there is a relatively long way to go in terms of production of the proof of concepts that will be necessary in order to develop the necessary tech-nologies. However, in some cases, such processes are already present in plants and could be improved using transformation, possibly of transcription factors that trig-ger the whole process.

It is important to note here that wood feedstocks would probably have to be treated differently regarding the strategies discussed in this chapter. Wood feed-stocks have large proportions of dead cells composed of high proportions of cellu-lose and also with a much higher degree of lignification. For this type of bioenergy feedstock, an addition to the strategy would be the modification of lignin com-position to facilitate access to the cell wall polysaccharides, which in fact is cur-rently under investigation (Vanholme et al. 2013). As grasses can have some lignin and phenypropanoids in their cell walls, as well as the cells of the vascular sys-tem, whose walls bear many features in common with the wood tissues, the use of the strategies proposed in this chapter for tissues containing large proportion or primary cell walls, along with the ones being developed for decreasing lignin interference, would probably have to be coupled in the future in order to improve bioenergy production even more.

Based on the examples discussed in this chapter, it is very likely that natural processes that involve cell wall modifications in plants will be useful as additions to 2G bioethanol technologies, via the use of modern molecular techniques associ-ated to systems and synthetic biology.

Acknowledgments This work is part of the production of the Instituto Nacional de Ciência e Tecnologia do Bioetanol-INCT do Bioetanol (FAPESP 2008/57908-6 and CNPq 574002/2008-1) and of the Centro de Processos Biológicos e Industriais para Biocombustíveis-CeProBIO (FAPESP 2009/52840-7 and CNPq 490022/2009-0). Financial support by FAPESP Projects 2010/17104-5, 2010/17070-3, 2011/07586-5 and 2011/02344-3.

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