REPÚBLICA BOLIVARIANA DE VENEZUELA UNIVERSIDAD PEDAGÓGICA EXPERIMENTAL LIBERTADOR INSTITUTO PEDAGÓGICO DE MATURÍN MATURÍN “ANTONIO LIRA ALCALÁ” ALCALÁ” DEPARTAMENTO DE CIENCIAS NATURALES CÁTEDRA: FISIOLOGÍA VEGETAL
FOTORRESPIRACIÓN
PROFESOR: NINO OMOGROSSO ESTUDIANTE: LUZ CAROLINA HIDROGO CI: 20!20"## MATURÍN MA$O DEL A%O 20&'
FOTORRESPIRACIÓ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
MEDIDA DE LA FOTORRESPIRACION C9.+-* ( /,- 5 )*+(9/* - O2 5. 31*-9)),7+ - CO2 3*1 =*>.( - 9+. 35.. - C#? /.++,-. + .9(+),. - CO2? ( +)9+1. +)9+1. 89 5. *()91,-.*()91,-.- 5. 5*),-.5*),-.1(3,1.*1,. ( (.91. <. . 6.>.( )*+)+1.),*+( - O2 &0 .5 &" 3*1 &00 E+ )./6, )./6,** + 31( 31(+) +),. ,. - 59@? 59@? .5 .9/+ .9/+.1 .1 5. )*+) )*+)+ +1. 1.), ),7+ 7+ - O2? )*+,+. .9/+.+-* .9/+.+-* 5. 5*),-.5*),-.- 1(3,1.*1,. 1(3,1.*1,.
./*(1. +. - CO2? +* ,+ 3*189 ( ,9.5 . 5. 89 *)911 + 5.( )*+-,),*+( =.6,9.5( - **(;+(,( U+ 31*)(* 1(3,1.*1,* ( 5 ,+,1+* - 5. **(;+(,(? 31*-9) CO2 < )*+(9/ O2 C9.589,1 /-,-. - 5. 31*-9)),7+ * )*+(9/* - CO2 < O2 (? + 1.5,-.- 5. /-,-. - 5. -,1+),. +1 5. **(;+(,( < **11(3,1.),4+ L. /-,-. ,+-,,-9.5 - 5. 5*),-.- - ).-. 9+* - (*( 31*)(*( +* ( 3*(,65 3*1 5*( /*-*( =.6,9.5( - (,/.),7+ - )./6,*( - CO2 < O2 L.( /-,-.( - **11(3,1.),4+ + .9(+),. - CO2 (*5* 39-+ (1 ,+-,).,.( - (9 1-.-1* .5*1 + )*+-,),*+( +.91.5( S 39-+ =.)1 /-,-.( /.( 31),(.( < )*/35>.( - 5.( /.+,9-( - 5. **(;+(,( *.5 < - 5. **11(3,1.),4+ 5. -,1+),. +1 ./6.( ( 5. **(;+(,( +.? )*+ 5 9(* - ,(**3*( 1.-,.),*( S,? 3*1 >/35*? 5.( =*>.( ( /.+,++ -91.+ 9+ ,/3* 31*5*+.-* + 9+. ./*(1. )*+
&!
CO2?
( /.1).+
1.-,.),./+ .5/,-7+? ,+1/-,.1,*( -5 ),)5* - ).5,+ < *1*( )*/39(*( S, 59* (.( =*>.( ( 3.(.+ . 9+. ./*(1. )*+ &2 CO2? +* 1.-,.),*? ( 5,/* ( .(,/,5.1. 3*1 **(;+(,(? 31* 5.( =*>.( ,+,),.5/+ 31*-9),1+ CO2? /.1).-* )*+ &! C 31*)-+ -5 )*+(9/* - **11(3,1.),4+ - (9( 1(1.( /.1).-.( )*+ &! C 1(95.+( - 5. 3*(,),7+ .+1,*1 A9+89 ./6,+ -+1* - .59+.( 1(1.( )1,,).(? 5. /-,-. - 5. 5*),-.,+,),.5 - 5. **11(3,1.),4+ C*+ 5.( <. -()1,.( < )*+ *1.( .31*,/.),*+( 31,/+.5(? ( (,/. 9+ .5*1 )*+(,-1.65 - 5. **11(3,1.),4+ 89? + /9)=.( 35.+.( C#? 39- (93*+1 9+ '0 3*1 &00 .5*1 - 5. **(;+(,( +. E+ )./6,* + 35.+.( C!? 5. **11(3,1.),4+ ,+ 9+ .5*1 -(31),.65
MECANISMOS DE LA FOTORRESPIRACIÓN E+ 5. ,+(,.),7+ - 5*( /).+,(/*( /*5)95.1( 1(3*+(.65( - 5. **11(3,1.),4+? 31*+* 1(957 ,-+ 89 5 ),-* 5,)45,)* CH2 OHCOOH (.6. ,/35,).-* + .59+.( .3.( - 5. **11(3,1.),4+ M9)=.( .5.( 31*-9)+ + **(;+(,( 1.+-( ).+,-.-( - ),-* 5,)45,)* 89 395(. .5 1,*1 E+ *1*( *1.+,(/*( )*+ **(;+(,( C#? 5 5,)45,)* (,1 )*/* (9(1.* 3.1. 5. 31*-9)),7+ **11(3,1.*1,. - CO2 D91.+ 6.(.+ ,/3* 9 9+ +,/. /.675,)* -5 ),-* 5,)45,)*? .9+89 ( (.6;. 89 +;. 89 11 .5* )*+ 5. **(;+(,( E+ &&? O1+ < B*( (9,1,1*+ 89 5 ),-* 5,)45,)* ( *1/.6. . 3.1,1 - 1,695*(. &?' -,*(.* )*+ *,+* E+ &#? L*1,/1 < A+-1( 35,).1*+ 5 *1,+ - 5. **11(3,1.),4+ 3*1 9+. 1.)),7+ +1 1,695*(. &?' -,*(.* < *,+* 5 )+1* .),* - 5. R9DP).16*,5.(. E5 5,)45,)* 31*-9),-* ( -(39( /.6*5,@.-* 3*1 -,1(*( 31*)(*( E+1 5.( ;.( 31,+),3.5( -5 /.6*5,(/* 3*(1,*1 -5 ),-* 5,)45,)* +/*( L. R9DP).16*,5.(.? .-/( - )..5,@.1 5. *1/.),7+ - -*( /*5)95.( - # *(*5,)1.* . 3.1,1 - 9+. - 1,695*(.-,*(.* < 9+. - CO2? 39- .-/,,1 O2 )*/* (9(1.* .51+.,* .5 CO2 S, ( O2 5 89 ..). . 5. 1,695*(.-,*(.* + 5 )+1* .),* -5 +@,/. 5. /*5)95. - 1,695*(.-,*(.* ( 1*/3 -.+-* 9+. /*5)95. - # *(*5,)1.* < *1. - 5,)*5.**(.*? 31*6.65/+ (+ 5 /).+,(/* S 1.. - 9+. .),,-.- *,+.-. .-,),*+.5 . 5. .),,-.- ).16*,5.(. -5 +@,/. P*1 ( /*,*? /9)=*( .9*1( 31,1+ 55./.1 . 5. .),,-.- )*/* 1,695*(. &?' -,*(.*).16*,5.(.*,+.(. A5 5,)*5.* *1/.-* 3.(.+ 5*( ).16*+*( & < 2 - 5. /*5)95. - 1,695*(. -,*(.* C*/* + 5. 1.)),7+ (*5* ( *1/. 9+. /*5)95. - # *(*5,)1.*? (
31*)(* (93*+ 9+. -,(/,+9),7+ - /*5)95.( - ,+1/-,.1,*( -5 ),)5* - C.5,+ 9+*? 5. *31.),7+ );)5,). - 5.( .3.( 3 y 5 (93*+ 5*6.5/+ 5 31*)(*:
O2 Q NADPH QHQ
H2O Q NADPQ
9 +* ( (,+* 5 31*)(* ,+1(* .5 89 ( 31*-9) + 5 1.+(3*1 5)17+,)*
**(,+,)* E+*+)(? ( 31*)(* **11(3,1.*1,* (93*+ 9+.
,+9,5,@.),7+ -5 NADPH 31*-9),-* + 5 1.+(3*1 5)17+,)* **(,+,)*
5,)45,)* ,+=1+ .5 31*)(*? 9+ .9/+* - 5. )*+)+1.),7+ -
,+1/-,.1,*( (93*+ 9+ .9/+* - 5. 5*),-.- -5 31*)(* P1),(./+? .5 .9/+.1 5. )*+)+1.),7+ - O2 -6 .9/+.1 5. *1/.),7+ -5 ,+1/-,.1,* 5,)45,,)* < )*+ 55* 5. 5*),-.- -5 31*)(* P*1 *1. 3.1? 5. 59@ ( 1891,-. 3.1. 5. *1/.),7+ - NADPH 89 ( 9,5,@.-* + 5 9+),*+./,+* -5 ),)5* E5 *1* 31*)(* **11(3,1.*1,* 189,1 9+ /.6*5,(/* 951,*1 -5 5,*.5.* ,/35,). 5. 3.1,),3.),7+? .-/( - )5*1*35.(*( < 31*,(*/.(? - /,*)*+-1,.( E( /.6*5,(/* -5 5,*.5.*? (+ 5.( .3.( 6, 7, 8, 9 y 10 =.(. 5. *1/.),7+ - #*(*5,)1.*? (93*+ 9+. 31*-9)),7+ - CO2 < NH# + 5. .3. 7. S,+* 5. +,).? 5. .3. 3.1) 5 /).+,(/* /.<*1,.1,* - *1/.),7+ - CO2 + **11(3,1.),4+ L*( -*( 31*)(*( **11(3,1.*1,*( ,++ .1,.( .3.( )*/9+( +1 55.? 5. )..5,@.-. 3*1 5,)*5.* *,-.(. 3 ( ,+=,6,-. 3*1 )*/39(*( .+5**( (19)91.5( -5 .),-* 5,)45,)*? )*/* 5*( ),-*( =,-1*,(95+,)*(? 89 (*+ )*+*),-*( ,+=,6,-*1( - 5. **11(3,1.),4+
E5 (9+-* 31*)(* **11(3,1.*1,* )*+(,-1.-*? ;. 5,),+. < (1,+.? 3.1) /( ,/3*1.+ )9.+,.,./+? ,+)59(* ( -9-. - 5. *31.,,-.- in vivo -5 31,/1* S =. (91,-* 89 5 (9+-* 31*)(* 39- (1 9+. ;. - *1/.),7+ **(,+,). - ./,+*),-*(? 31* + 5. 31),). (93*+ 9+ 1.+ -(.( - +1;. < ).16*+.-*( - 5. **(;+(,(
ASIMILACIÓ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
PLANTAS C3 L.( 1.)),*+( - ,>.),7+ - ).16*+* 89 *)911 + 5 (*/.? 5 NADPH < 5 ATP? 31*-9),-* + 5.( 1.)),*+( - ).391. - 5. +1;.? ( 9(. 3.1. 1-9),1 9+ )*/39(* - 1( ).16*+*? 5 5,)1.5-=,-* *(.* E+ (. ;. -*+- 5 ).16*+* ( ,>. 3*1 /-,* - 9+ 5,)1.5-=,-* *(.* ( 5. -+*/,+. ;. - 1( ).16*+*( * C# E( ,3* - 35.+. 3*( 5 /).+,(/* **(,+,)* /( 6(,)* L. /.<*1;. - 5.( (3),(? + (3),.5 . .8955.( 89 31+)+ . ./6,+( =/-*( < /35.-*(? (*+ - ( 193* L.( 35.+.( C# ( )*/3*1.+ /( ,).@/+ + )*+-,),*+( - /31.91. +* /9< .5. < .5. =9/-.- 15.,. S*+ C# 5. /.<*1 3.1 - 5.( 35.+.( )95,.-.(? )*/* 5 1,*? 5 ,1.(*5 * 5.( )*5( E+ ( ).(* 5. ,>.),7+ - ).16*+* ( 55. . ).6* 3*1 /-,* -5 ),)5* - C.5,+? + -*+- 5. +@,/. 1,695*(. -,*(.* ).16*,5.(. R9BPCO )*/6,+. 9+. /*5)95. - ).16*+* )*+ 5 /.1,.5 - 3.1,-.
PLANTAS C4 E( ,3* - 35.+.( 31(+. 9+. .+.*/;. *5,.1 - ,3* 1.+@ * - )*1*+.? 89 5 31/, ,+)1/+.1 + (9( =*>.( .1,.( )( 5. )*+)+1.),7+ - CO2 ./6,+.5 E+ 5.( 35.+.( C! 5.( 1.)),*+( 31,.( .5 ),)5* - C.5,+ )*+(,9<+ 5. 55./.-. ;. - !"#$ y S%!#& L. ,>.),7+ - CO2 3*1 5. +@,/. - R9B,(CO +* 31*)- - 5. ./*(1. ,+ 59.1? + 31,/1 1/,+* - *1/. 1.+(,*1,.? + 5 ),*(*5 - 5.( )595.( -5 /(*,5*? -*+- 5. +@,/. PEP ).16*,5.(. 5* 9+ .5 .),-* *(*+*53,19,)* PEP? - #C L. +@,/. PED ).16*,5.(. ,+ 1.+ .,+,-.- 3*1 5 CO2 < 3*1 *1. 3.1 +* ,+ .),,-.- *,+.(.? 3*1 5* 89 5 O2 +* ,+1,1 + 5. ,>.),7+ E5 CO2 ).3.-* + 5 /(4,5* < 5,61.-* + 5. .,+. 1(95. /9)=* /. )*+)+1.-* + (. 89 5 **(,+,)* - 5.( 35.+.( C# C*/3,,+-* />*1 89 )*+
5 O 2 < .*1),+-* 5. .,,-.- ).16*,5.(. - 5. R9B,(CO D (. ).16*,5.),7+ ( *6,+? )*/* 31,/1 31*-9)* - ,>.),7+ -5 CO2? 9+ .),-* -,).16*,5;)* - !C? 5 .),-* *.5.),)* E5 *.5.),)* ( 59* 1.+(*1/.-* + *1* .),-* - !C 89 (1 1.(3*1.-*? ;. 35.(/*-1/*(? . 5.( )595.( - 5. .,+. E+ .59+.( 35.+.( C! (. 1.+(*1/.),7+ ( 9+. 1-9)),7+? 89 ,+ 59.1 + 5* )5*1*35.(*( -5 /(4,5*? < - 5. 89 ( *6,+ .),-* /5,)* + *1.( ( 1.. - 9+. 1.+(./,+.),7+ 89 *)911 + 5 ),*(*5 < - 5. 89 ( *6,+ .),-* .(31,)* E5 /5,)* +1. . 5* )5*1*35.(* - 5. .,+. < .55; ( -().16*,5.-* . .),-* 3,1,)* E5 .(31,)* ( 1)*+1,-* . *.5.),)* + )595. - 5. .,+.? <. (. + /,*)*+-1,. + ),1.( (3),( * + 5 ),*(*5 + *1.( 35.+.( 5 *.5.),)* ( 1-9),-* . /5,)* < -().16*,5.-* . 3,1,)* E+ *-*( 5*( ).(*( 5. -().16*,5.),7+ 5,61. CO2 + 5. .,+.? 89 39- +1.1 .5 ),)5* - C.5,+ .5 (1 ,>.-* 3*1 5. +@,/. R9B,(CO 5 .),-* 3,1,)* 89 89-. 1.+(3*1.-* +9./+ .5 /(*,5*? -*+-? 31,. *(*1,5.),7+ 3*-1 1,+,),.1 5 ),)5* P*1 ).-. CO2 ,>.-*? 5 /.6*5,(/* C! ,+ 9+ /.<*1 )*( +1,)* 89 5.( C#? )*+ .(* - ' ATP < 2 NADPH + )*/3.1.),7+ - # ATP < NADPH - 5.( 35.+.( C# A .5.( /31.91.( #0WC ( .*1) 5. *,+.),7+ - 5. R9B,(CO < 3*1 5* .* 5. 31-,-. - 5. **11(3,1.),*+ <. 89 5 )* - 5. ,. !"#$ y S%!#& E( 1-9),1 * .+95.15. *,+.),7+ - 5. R9DP C*+ ).-. 6.>. =9/-.- 15.,.? 5*( (*/.( ,+-+ . )11.1(? ,/3,-,+-* 5 .-)9.-* (9/,+,(1* - CO2 .5 ,+1,*1 - 5.( =*>.( L.( (3),( C! 31+)+ . 9+.( ./,5,.( - .+,*(31/.(? +1 5.( 89 -(.).+ 5.( G1./;+.(? ./.1.+.).( S +)9+1.+ +1 55.( 5.( (3),( )95,.-.( - /.<*1 31*-9),,-.- .1;)*5.? )*/* 5 /.;@? 5 (*1* * 5. ).. - .@).1 < .59+.( - 5.( /.5.( =,16.( .1(,.( )*/*? C<+*-*+ -.)<5*1
PLANTAS CAM S ).1.)1,@.+ 3*1 +1 ,>.),7+ - CO2 +*)91+. < )*/35/+.1 5 31*)(* - **(;+(,( .5 -;. (,9,+ E( /).+,(/* **(,+,)* 5 31/, ,.1 5. 31-,-. - .9. L.( (,5.( CAM (,+,,). “METABOLISMO ACIDO DE LAS CRASULACEAS”? -6,-* . 89 (. .1,.+ **(,+,). ( -()1,6,7 ,+,),.5/+ + 35.+.( - (. ./,5,. S 1.. + +1.5 - 35.+.( *1,,+.1,.( -(1,).(? (*/,-.( . ,++(. ,59/,+.),7+? .5.( /31.91.( < 31*+9+),.-*( -,), =;-1,)*(? .-.3.-.( . )*+-,),*+( - .1,-@ 6.(.+( 1/.( E+ (.( 35.+.( 5 >,-* **(,+,)* ( =*/*+*? (,+ .,+. -,1+),.-. +, ./3*)* )5*1+89,/. /3.5,@.-. P1* (9( (*/.( /9(1.+ 9+ 3)95,.1 )*/3*1./,+* <. 89? .5 )*+1.1,* - 5.( -/( 35.+.(? ( .61+ - +*)= < ( ),11.+ - -;. E5 /.6*5,(/* - 5.( 35.+.( CAM 31(+.+ ./6,+ 9+.( 1.)),*+( 31,.( .5 ),)5* - C.5,+? (,/,5.1( . 5. - 5.( 35.+.( C! < ( 1,,). .(,/,(/* 9+. )*/3.1,/+.5,@.),*+? 31* +* (3.),.5 <. 89 5 )5*1+89,/. ( 9+,*1/ (,+* /3*1.5: 5.( 1.)),*+( -5 ),)5* - C.5,+ *)911+ - -;.? )*+ 5*( (*/.( )11.-*(? /,+1.( 89 5.( 1.)),*+( 31,.( ,++ 59.1 - +*)= )*+ 5*( (*/.( .6,1*( D91.+ 5. +*)=: 5*( (*/.( .6,1*( 31/,+ 5. ,>.),7+ -5 CO2 ./*(1,)* 3*1 5 PEP ).16*,5.(. + 5 ),*(*5: 5 PEP (*61 5 89 .). (. +@,/. 31*)- - 5. -1.-.),7+ -5 .5/,-7+? .)9/95.-* + 5*( )5*1*35.(*( -91.+ 5 -;. D 5. ).16*,5.),7+ -5 PEP ( *6,+ .),-* *.5.),)*? 89 59* ( 1-9),-* . /5,)* E5 .),-* /5,)* +* ( 1.(3*1. . *1.( )595.( (,+* 89 ( .)9/95. + 5. .)9*5. - 5. /,(/. )595. D91.+ 5 -;.: )*+ 5*( (*/.( )11.-*( 5 /5,)* (.5 - 5. .)9*5. < ( -().16*,5. . 3,1,)* + (. 1.)),7+ ( 5,61.CO2? 89 +1. . 5*( )5*1*35.(*( 3.1. ,+,),.1 .55; ),)5* - C.5,+ E5 ),-* 3,1,)* ( 1.+(*1/.-. + PEP? 89
59* 3.(. . *(.* - 1,*(. 5.( 1,*(.( + 5*( )5*1*35.(*( -.+ 59.1 . 5. (;+(,( < .)9/95.),7+ - .5/,-7+? . 3.1,1 -5 )9.5 ( 1+1.1 5 PEP -91.+ 5. +*)=
OTROS MECANISMOS DE CONCENTRACIÓN DE CO2 A59+*( .5( .)9,)*( .(,/,5.+ CO2 -5 /-,* 3*1 (,/35 -,9(,7+ =.(. 5. 196,()*? -5 /*-* (,/,5.1 . )*/* 5* =.)+ 5.( 35.+.( 11(1( -5 ,3* C#? (*( (*+ (,+ /6.1*? 5. /,+*1;. E+ 5 ./6,+ .)9,)*? 5. 1.-,* - )*+)+1.),*+( O2CO2 ( .9+ /.<*1 89 5. ./*(1. M.( .9+? 5 )*,),+ - -,9(,7+ -5 CO2 + .9. ( )9.1* *1-+( - /.+,9- ,+1,*1 .5 )*,),+ + .,1 < -)1) .-/( 5. /31.91. L*( 31,+),3.5( 193*( - .5( .)9,)*( 39-+ )5.(,,).1( + ),+)* ).*1;. (+ (9 /).+,(/* - .-89,(,),7+ - ).16*+* ,+*1+,)*:
V.5( 89 .-89,1.+ 5 CO2 -,1)./+ -5 /-,* 3*1 -,9(,7+ ( ( 9+ /).+,(/* 3*)* )*/+ + 35..( .)9,).(? 31*3,* - ./6,+( (3),.5/+ +1,89),-*( + CO2 3*1 *1.+*1.;.? .5( )*/* 389.( )=.1).( /3*1.5( 917,).(
R,@*,.( 89 )1)+ (*61 5 (-,/+* +1,89),-*( + CO2 3*1 *1.+*1.;. E+ ( ).(*? 5 CO2 ( 1.+(3*1. -(- 5.( 1.;)( . 5*( .55*( . 1.( - (3.),*( .1*( ,+1)595.1( E( /).+,(/* ( 31*3,* - 35..( ()9+-.1,.( .)9,).(? )*/* 5. ,(*.5( T./6,+ ( +)9+1. + ,(*,-*( 11(1( 89 ).1)+ - (*/.(? )*/* stylites andicola.
P5.+.( < .5.( 89 9,5,@.+ HCO# 3.1. 5. .)9/95.),7+ -5 ).16*+* ,+*1+,)* E( 5 /).+,(/* /.( )*/+ + 5. /.<*1 3.1 - 5* .5( .)9,)*( elodea canadensis, patamogeton lucens, chlamydomonas) L*( /).+,(/* .)*( - 6*/6* - HCO# + (3),.5 (, ( 1.. - /).+,(/* 5)1*+,)*( * /-,.-*( 3*1 1.(3*1 - ,3* 9+,3*1? .,3*1 * (,/3*1? .),*( * 3.(,*( .(, )*/* (9 ,+11),. )*+ 5.
195.),7+ -5 3H ,+1.)595.1 (*+ 9+* - 5*( )./3*( - (9-,* - 5. **(;+(,( .)9,). - /.<*1 .)9.5,@.),7+
P5.+.( )*+ )*/3*1./,+* CAM * C! L*( /).+,(/*( .+1,*1(? .3.1 -5 6*/6* .),* - ).16*+*? 39-+ )*+(,-1.1( ;3,)./+ 31*3,* - 5.( 35.+.( C# E,(+ *1*( .5( .)9,)*(? (,+ /6.1*? 5.89 5. (1.,. )*+(,( + 9+. 31,/1. ,>.),7+ 1.5,@.-. 3*1 PED ).16*,5.(. * PEDE).16*,89,+.(. *1,,++-*( 9+ )*/39(* - )9.1* ).16*+* 89 3*(1,*1/+ ( -().16*,5.-* < 5. ()9+-.1,. 3*1 196,()* ( - ,3* /3*1.5? 3*1 5* 89 ( )*+(,-1. 89 (.( 35.+.( ,++ /).+,(/* **(,+,)* - ,3* CAM? .5 ( 5 ).(* - .59+*( ,(5*( .)9,)*( * - 5.( )1.(95).( .)9,).( S )*+*) ./6,+ .59+*( ).(*(? *-.;. -,()9,-*( - (3.),*+ (3),.5 - ,3* C! )*/* 5. )5*1*;). /.1,+.? )*+ /.6*5,(/* **(,+,)* (,/,5.1 .5 - 5.( 35.+.( C! 31* (,+ 5. .+.*/;. ,3* 1.+@
'ALANCE DEL META'OLISMO FOTORRESPIRATORIO C*+(,-1.+-* 5 31*)(* **11(3,1.*1,* -(- 5. 1,695*(.-,*(.* =.(. # *(*5,)1.* .3.( (9)(,.(: &? 2? #? J? ? "? < &0? 3*1 -*( /*5( - 1,695*(. -,*(.* 9,5,@.-*( ( )*+(9/ 1( /*5( - O2 .3.( &< # )*11,,+-* (. 95,/. 3*1 5. 31*-9)),7+ - /-,* /*5 - *,+* + ! < 9+ /*5 - ATP 31*-9),+-*( 9+ /*5 - CO2 < 9+ /*5 - NH#? )*+ 5 3.(* .-,),*+.5 - 9+ /*5 - 59./.* . 9+* - 2**59.1.*- 5*( -*( 59./.*( 9,5,@.-*( + J? 9+* ( 1)931. + "? ( -),1? 5*6.5/+: 2 R9DPQ # O2 Q &ATP Q & G59./.* **59.1.*
## PGA Q & CO2 Q & ADP Q 2 P , Q &2
E( ),)5* **11(3,1.*1,* ( )*/35. )*+ 5. )*+1(,7+ - 5*( 1( /*5( - #*(*5,)1.* 31*-9),-*(? + 9+* < /-,* /*5( - 1,695*(.-,*(.* ,. ),)5* - C.5,+? 5* 89 189,1 1( /*5( - NADPH < )9.1* < /-,* /*5( - ATP E+ )*+>9+*? 5. *31.),7+ -5 31*)(* **11(3,1.*1,* -5 /.6*5,(/* -5 5,*.5.* ,. 5,),+. < (1,+. =.(. #*(*5,)1.* < 1,695*(.-,*(.*? (93*+ 9+ 5.-* )*+(9/* - ATP .-/( - 5. 31*-9)),7+ - CO2 < 5 )*+(9/* - O2 E5 6.5.+) *.5 =.(. (. .3. ( .+ ,+),1*? 39( -3+- - 5. ,. 3*1 5. 89 5 NH # 31*-9),-* ( ,+)*13*1. + 5 2**59.1.* 3.1. 1+1.1 59./.*? 89 )*+(9/ 89,.5+( - 1-9)),7+
OTROS PROCESOS META'ÓLICOS RELACIONADOS CON LA FOTORREPIRACIÓN E( 31*6.65 89 ,(.+ *1*( 31*)(*( /.675,)*( 15.),*+.-*( )*+ 5. **113,1.),7+ .9+ 3*)* )*+*),-*( E( 6,+ (.6,-* 89 5 ),-* 5,)75,)* )*+ .9. *,+.-.? H2 O2? + 1.)),7+ )..5,@.-. 3*1 5. /,(/. 5,)*5.* *,-.(.? ( *,-.-* 1.)),7+ 31*,-(,). . ),-* *5,)* CO2H CO2H 89 ( 39- -().16*,5.1 .
),-* 71/,)* HCO2H E+ 5. /-,-. + 89 5 H 2O2 31*-9),-. + #? +* (. -()*/39(. 3*1 )..5.(.? !? 5. *1/.),7+ - ),-*( 71/,)*( < *5,)* 39- (1 9+ 31*)(* .-,),*+.5 - 5. **113,1.),7+ E( 3*)* )*+*),-* 5 /.6*5,(/* 3*(1,*1 -5 ),-* 71/,)* 3.1. 5 89 ,(+ .1,.( 3*(,6,5,-.-( P*1 *1. 3.1? .59+.( 35.+.( 39-+ .)9/95.1 1.+-( ).+,-.-( - ),-* *5,)*? < 9+. 3*(,6,5,-.- ( 89 (9 *1,+ ( + 5. **113,1.),7+ + =*>.( )*+ 6.>.( .),,-.-( )..5.(. < -().16*,5.+ - ),-* *5,)* P.1) (91* ./6,+ 89? ;. **11(3,1.),7+? ( *1/.+ 389.( ).+,-.-( - /*+7,-* - ).16*+*? CO? -(- ),-*( 5,*;5,)* < 71/,)* 3*1 -().16*,5.),7+ <* -(=,-1..),7+ F,+.5/+? ( =. (91,-* *1* *1,+ /.675,)* 3.1. 5 ),-* 5,)75,)* . 3.1,1 -5 )*/35>* 5,)*5.5-=;-* 1.+()*5.(.? ,+1/-,* + 5. 1.)),7+ - 5. 1.+()*5.(.? -5 ),)5* - C5.,+? 31* +* 3.1) /9< ,/3*1.+ (. ;. - *1/.),7+ -5 5,)75,)*
CONSEC(ENCIAS DE LA FOTORREPIRACIÓ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
P(NTO DE COMPENSACIÓN DEL CAR'ONO E+ 9+. ./7(1. )11.-.? + .9(+),. -5 CO2? 5.( =*>.( 5* 31*-9)+ < )*+(9/+ *,+* 3*1 1(3,1.),7+ /,*)*+-1,.5 *(? +1 0 < &0 33/ - CO2 L7,)./+? 5 39+* - )*/3+(.),7+ -3+- - 5. )*+)+1.),7+ - O2? 5. /31.91. < 5. ,++(,-.- - ,59/,+.),7+
E,( ./6,+ 9+ 39+* - )*/3+(.),7+ 59/,+*(.? (* (? ,++(,-.- - ,59/,+.),7+ 3.1. 5. 89 ( 89,5,61.+ **(;+(,( < 1(3,1.),*+( *5,.1( E+ ( ).(*? 5.( 35.+.( C! .5).+@.+ (9 39+* - )*/3+(.),7+ . 9+*( &00 59? /,+1.( 89 5.( 35.+.( C# 5* .5).+@.+ +1 &000 < '000 59 H.< 89 +1 + )9+. 89 5. 59@ ./6,+ .),. 5. **113,1.),7+ + 35.+.( C#? 3*1 5* 89 (.( 1891,1+ /( 59@ 3.1. 5 39+* - )*/3+(.),7+ L.( .5.(? .9+89 1.5,@.+ 9+. **(;+(,( ,3* C#? 31(+.+ 39+*( - )*/3+(.),7+ - CO2 6.>*( E+ /9)=*( ).(*(? + .5.(? /( 89 9+. 31*-9)),7+ **11(3,1.*1,. - CO2? 5* 89 *)911 ( 9+. )1),7+ .5 /-,* - 5,)*5.* < *1*( )*/39(*( *1+,)*( -1,.-*( - (9 /.6*5,(/* E+ .59+.( .5.(? ( =. -/*(1.-* 5. ,(+),. - 9+ 1.+(3*1 .),* - CO2 . 5. )595. P1*6.65/+ ( /).+,(/*? .5 .)9/95.1 CO2 + 5. )595.? 31/, 5*1.1 9+. .5. 15.),7+ YCO2 YO2 + 5 )+1* .),* - 5. R9DP ).16*,5.(.? 5* 89 -1/,+.1;. 6.>.( 5*),-.-( - **113,1.),7+
FOTOS)NTESIS APARENTE O INTERCAM'IO NETO DE AN)DRIDO CAR'ÓNICO T.+* 5. **(;+(,( )*/* 5. **113,1.),7+ ( 31*-9)+ + 5. =*>.? 5. .(. - .(,/,5.),7+ -5 CO2 **(;+(,( +. ( 31,+),3.5/+ 9+ 6.5.+) +1 5 CO2 ,+,),.5/+ ,>.-* **(;+(,( 619. < 5.( 31-,-.( **113,1.*1,.( C*/* (+ ,/35,).-*(? 5*. /,(/*( .(( + ./6*( 31*)(*(? 31* )*+ 59>*( + (+,-* )*+1.1,*? ( -,;),5 /-,1 3*1 (3.1.-* ./6*( 31*)(*( E+ 31,+),3,*? 3*1 ).-. ! ).16*+*( 89 +1. ( 3,1- 9+* )*/* CO2 + )*+-,),*+( ./*(1,).( .)9.5( 5* 89 ,+-,). 89 5. **(;+(,( 619. *(),5. .51--*1 - &2'[ E5 6.5.+) +1 5*( -*( ),)5*( (. -1/,+.-* 3*1 5.( 31*3,-.-( ),+,).( - 5. 196,()* < 5. )*+)+1.),7+ - (9(1.*( O2 < CO2 E(.( 31*3,-.-( (*+ -3+-,+( - 5. /31.91. 89 .5 ,+)1/+.1( .*1) 5. .),,-.- *,+.(.
-,(95*( + .9. 3*189 5. (*596,5,-.- -5 CO2 -)1) /( 89 5. -5 O2? 3*1 5* )9.5 .5 .9/+.1 /31.91.? ( .*1) 5. **11(3,1.),7+
FOTORRESPIRACIÓN * E+OL(CIÓN DE LOS DISTINTOS TIPOS DE FOTOS)NTESIS D,;),5/+ ).6 .1,69,1 9+. 9+),7+ + 5. 35.+. . 5. **11(3,1.),7+ A 3(.1 - 89 5.( 35.+.( 39-.+ =.)1 .5+ 9(* - 5. **11(3,1.),7+? (. + )*+>9+* 3.1) (1 +.,. 3.1. 5 .31*)=./,+* **(,+,)* - 5. +1;. 59/,+*(. 3*1 5. 35.+.? * .5 /+*( 5. **11(3,1.),7+ +* 3.1) 31*3*1),*+.1 +,+9+. +.>. 3.1. 5. 35.+. L. ,-. /.( .)3.-. =*< -;. ( 89 + 5. ./7(1. 31,/,,. + 89 .3.1),1*+ 5*( 31,/1*( *1.+,(/*( **(,+,)*(? +* =.6;. .3+.( *,+* .( .-.3.,.(? .5 /+*( + 5.( @*+.( - .5.( /31.91.( ./6,+.5(? + 5.( 89 5. **11(3,1.),7+ ( ,/3*1.+ < ( 1.+- 5. -,1+),. - 5.( 5*),-.-( - **(;+(,( +. +1 35.+.( C# < C ! F1)9+/+? )5,/.( ).591*(*( < ()*(? ,+)59(* ./6,+ ./6,+( (.5,+*(? .+ 9+,-*(? 5* 89 )*+,1,7 . 5.( 35.+.( C! +.>.( .-.3.,.(? .5 /+*( + (*( ./6,+(? 89 31/,,1*+ (9 -(.11*55* D *-.( *1/.(? + @*+.( /( /35.-.( *
1;.( < =/-.( +* ( -,1*+ /*,*( 3.1. 9+ -(35.@./,+* ,/3*1.+ - 35.+.( C# 3*1 35.+.( C! E+ )9.589,1 ).(*? 3.1) 89 5 .9/+* - 5. )*+)+1.),7+ - *,+* + 5. ./7(1. ( -,* /9< /31.+./+ + 5. *59),7+ - 5. ,-. (*61 5. ,11.? /,+1.( 89 ( (93*+ 89 5. .3.1,),7+ - **(;+(,( C! ( 1),+ P1*6.65/+? .5.( )*+)+1.),*+( - CO2 ./*(1,)* +* =,),1*+ +.>*(. 5. .3.1,),7+ - 5. **(;+(,( C! + 5 3.(.-* 1/** S+ .59+*( .9*1(? 5. (,/,5,9- /.675,). +1 35.+.( CAM < )595.( *)59(,.( - 5.( (*/.(? (9,1 89 .8955. (91,7 - 9+. +(,7+ .5 /(7,5* - 5.( ).1.)1;(,).( /.675,).( - 5.( )595.( *)59(,.( L,1.( /*-,,).),*+( -5 /.6*5,(/* **(,+,)* CAM? < )./6,*( /*1*57,)*( + 5.( =*>.(? (1;.+ 5 *1,+ - 5. **(;+(,( C! E+ ( (+,-*? 5 *1,+ - 5*( .)*+),/,+*( *59,*( + 5 /.6*5,(/* **(,+,)* 1.-,).1;. + 9+. 13,-. .-.3.),7+ . ./6,+( ()*( A9+89 3*(1,*1/+? .-.3.),*+( . 6.>.( )*+)+1.),*+( - CO2 55.1*+ . *1.( *1/.( - **(;+(,( L. +-+),. .)9.5 - .9/+* - 5. )*+)+1.),7+ - CO2 + 5. ./7(1. -6 .*1)1 5. **(;+(,( +. + 35.+.( C#? 89 3*-1;.+ .5)
[email protected] 31*-9),,-.-( 317,/.( . 5.( - 35.+.( C! E5 31(9/,65 .9/+* .(*),.-* - 5. /31.91.? + )./6,*? -6 (1 .5* +.,* 3.1. 5. 31*-9),,-.- **(,+,). +. - 5.( 35.+.( C#
F(NCIÓN DE LA FOTORRESPIRACIÓN E+ *1-+ . /.+,,).1 5. 31*-9)),7+ ( =. ,++.-* 1-9),1 * 5,/,+.1 5. **11(3,1.),7+ + 35.+.( C# )*+(,-1.+-* 89 +* ( 9+ 31*)(* (+),.5 L. 31(9+),7+? +* ( )*11).? . 3(.1 - 89 ( =. 5*1.-* 69+.( .)9/95.),*+( - /.1,. (). )9.+-* 5.( 35.+.( =.+ )1),-* . .5. )*+)+1.),7+ - -,7,-* -
).16*+* * . 6.>. +(,7+ - *,+* S,+ /6.1*? . 6.>.( )*+)+1.),*+( - CO2? 3*1 .9(+),. - .)3*1 5)17+,)* ,(,*57,)*? ( 31*-9)+ ***,-.),*+( -(19),.( + 5 )5*1*35.(* E( -.* 39- 31*-9),1( + )*+-,),*+( + 89 5*( (*/.( (+ )11.-*( . 5. 59@ .( ,11.-,.+),.(? (,+ 31*65/.( - ***,-.),7+? 5. ,+=,6,),7+ **(,+,). 3*1 5 *,+* 9( 389. - ,+=,6,),7+ 91. -5 /,(/* . .5. 89 . 6.>. ,11.-,.+),. E55* )*+-9) . 5. ,-. 89 5. **11(3,1.),7+ 31* . 5. 35.+. - 5. ***,-.),7+ )9.+-* 5. )*+)+1.),7+ - CO2 ( 5,/,.+ 31,*-*( - +(,7+ =;-1,).? .5.( /31.91.(? ) 55* 89-. 1*
[email protected]* 3*1 5 =)=* - 89? ,59/,+.+-* =*>.(? . .5. ,11.-,.+),.? -91.+ .59+.( =*1.(? . 6.>. )*+)+1.),7+ - *,+* < )*+)+1.),7+ - -,7,-* - ).16*+* 3*1 -6.>* -5 39+* - )*/3+(.),7+? ( ).9(. 5. **,+=,6,),7+ - 5. **(;+(,( (96(,9,+ T./6,+ ( =. .(,+.-* . 5. **11(3,1.),7+ 9+. 9+),7+ + 5. (;+(,( - (.).1*(. < - ./,+*),-*(? )*/* 5. 5,),+. < 5. (1,+. S,+ /6.1*? 5. .(. - 59>* . 1.( - 5. 19. -5 5,)*5.* ( /9< (931,*1 . 5.( +)(,-.-( - ./,+*),-*( E+ )9.+* . 5. (;+(,( - (.).1*(.? 5. ;. . 1.( - 5.( 1,*(.( *(.* ( /9)=* /.( -,1). < /+*( -,(,3.-*1. - 5. +1;. P*1 5. 19. - C.5,+ ( )*+(9/ # ATP < 2! NAD P HQ HQ
.),,-.-( ).16*,5.(. < *,+.(. ( 39-+ .1,.1 -,1+),.5/+ (9(,9<+-* 5 M2Q? .1,.+-* 5. /31.91. < + (3),( -,(,+.(? 3*1 5* 89 5. ,+/9.6,5,-.- - 5. 1.)),7+ ).16*,5.(. *,+.(. +* ( *.5/+ ),1.
FL(O ANAPLERÓTICO A LA L( D(- 5 39+* - ,(. )5(,)*? ( =. )*+(,-1.-* 89 5. 59@ ,+=,6 5. 1(3,1.),7+ 3*1 .9/+* -5 )*),+ NADPH QHQ NADPQ? ).3.@ - ,+=,6,1 5 ),)5* *,-.,* 3*1 5.( 3+*(.( *(.*? < 3*1 5 ,+)1/+* -5 )*),+ ATP ADP? 89 ,+=,6,1;. 5. 59)*5,(,( < 5.( ).-+.( 1(3,1.*1,.? 1.),.( . 5*( 1.+(5*).-*1( )5*1*35(,)*( P, DHAP? DHAP PGA? MA OAA < MA A(3 OAA G59 R.5/+? 5. *31.),7+ - 5. **(;+(,( -1/,+.1;. /+*1( +)(,-.-( - )*/39(*( - .5. +1;. < - 3*+),.5 1-9)*1 .( ,59/,+.-.(? 9,5,@.+-* -,(,+.( )+,).( P1* *1*( (9-,*( (,/,5.1( (9,1+ 89 5. 1(3,1.),7+ - *()91,-.- =,6 .(.( /9< (,+,,).,.( . 5. 59@ L.( ,-+),.( 6,*89;/,).( ,+-,).+ 89 5*( )5*1*35.(*( 31(+.+? .+* 5. 59)75,(,(? )*/* 5 ),)5* *,-.,* - 5.( 3+*(.( *(.* 3.1),.5( ,+=,6,-*( . 5. 59@ P1* ( +)(.1,* /+),*+.1 89? + 5 ),*(*5? /9)=*( - 5*( ,(*+@,/.( ,+*59)1.-*( +* ( 195.+ -5 /,(/* /*-* 89 + 5 )5*1*35.(* E( .5./+ ,/31*6.65 89 ( 31*-9@). 5. (931(,7+ )*/35. - 5. 59)75,(,( < -5 ),)5* *,-.,* - 5.( 3+*(.( *(.*( 3*1 )*( - 5. ,59/,+.),7+? <. 89 (.( 19.( (*+ 5.( +,).( 9+( - -,1(*( )*/39(*( )*/* 5 .),5 C*A < 5*( \ )*),-*( 89 (*+ ,/31(),+-,65( 3.1. /95,9- - ;.( 6,*(,+,).( 89 ( 31*-9)+ ,++(./+ . 5. 59@? )*/* 5.( (;+(,( - 5;3,-*(? ./,+*),-*( < 31*;+.( E+ 5 ,+1,*1 - 5*( )5*1*35.(*( ,+ 59.1 5. (;+(,( - /9)=*( ./,+*),-*( < - 5.
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
* .+.3517,)* ( *1,,+.+ ,+/-,../+ .+( 3*1 **(;+(,( E+ ( .(3)*? 3*1 )*+(,9,+? 5 59>* .+.3517,)* 1(3,1.*1,* - 5. 59@ ( .(/>.1;. /( . 5. **11(3,1.),7+ 89 . 5. 1(3,1.),7+ - *()91,-.- ;3,).
L. 3,19.*89,+.(. PA? ./6,+ 195.-. 3*1 5 (,(/. .-+,5.*(? )*/* 5. FF? 3*-1;. (*1.1( /-,.+ 9+ /).+,(/* 5.1.5 89 ,/35,). . 5. PEP C.(. < .5 NAD +@,/. /5,)* S,+ /6.1*? B.((=./ < )*5.6*1.-*1( &"& (9,1,1*+ 89 5 .9/+* - NH!Q 39- ,+)1/+.1 5 59>* .+.3517,)* 3.1. 5. (;+(,( - ./,+*),-*( . 1.( - 5. 31*3,. FF < - 5. PA L.( )*+-,),*+( 1-9)*1.( - 5. )595. -91.+ 5. **(;+(,( .*1)1;.+ 5. *31.),7+ 1-9),. -5 1+(,* - OAA . MA
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
CAR'OILACIÓN-OI/ENACIÓN S, 5. 196,()* *,+.1. /( 89 5. ).16*,5.(.? 5. 35.+. /*1,1;. < (. 15.),7+ ).16*,5.),7+ *,+.),7+ ( .).-. 3*1 9+. (1,. - .)*1(:
C.1.)1;(,).( ),+,).( -5 196,()*
C*+)+1.),7+ - CO2 < O2? /*5)95.( 89 )*/3,+ +1 (, 3*1 9+,1( .5 (,,* .),* - 5. 196,()*
T/31.91. .). . 5*( -*( .)*1( .+1,*1(
E5 .)*1 ,+1;+()* 15>. 5.( ).1.)1;(,).( ),+,).( - 5. 196,()* L. 5*),-.- ).16*,5.),7+ -3+- - 5. 5*),-.- /,/. - ).16*,5.),7+ .5. 5. 1-9+-.+),. - 5. .,+,-.- 3*1 5 CO2 < - 5. )*+)+1.),7+ - CO2 )*/* 5. *,+.),7+ -3+- - 5*( /,(/*( .)*1( (*5* 89 .35,).-*( . 5. )*+)+1.),7+ - O2 S, ( -,,-+ 5.( )9.),*+( - 5.( -*( 1.)),*+( ( *6,+ 5. 5*),-.- 15.,. - ).16*,5.),7+ )*+ 1(3)* . 5. *,+.),7+ + (. )9.),7+ ( 39-+ -,1+),.1 -*( 1/,+*( + 5 31,/1* (+ 15.),*+.-.( 5.( ).1.)1;(,).( ),+,).(
- 5. +@,/.? < ( . (* . 5* 89 ( -+*/,+. .)*1 ,+1;+()* _ E+ 5 *1* 1/,+* (+ 131(+.-.( 5.( )*+)+1.),*+( - CO2 < O2 S 39- -),1 89 5 .)*1 ,+1;+()* ( 5. 5*),-.- 15.,. - ).16*,5.),7+ )*+ 1(3)* . 5. *,+.),7+ . ,9.5( )*+)+1.),*+( - CO2 < O2 N* ( 9+ .5*1 ,-+,)* 3.1. *-.( 5. 196,()* C*+(,-1.+-* 89 5. 196,()* ,+ 9+ .5*1 ,+1;+()* - &00 < 89 + )*+-,),*+ +*1/.5(? ( -),1 . 2'WC 5. )*+)+1.),7+ - O2 ( - 2'0 < 5. - CO2 ,+ 9+ .5*1 - "? 5. 5*),-.- 15.,. - ).16*,5.),7+ *,+.),7+ - 5. 196,()* (1 # ).16*,5.),7+( 3*1 ).-. *,+.),7+ E,-+/+ (*( 1(95.-*( ( + .).-*( 3*1 )./6,*( + 5. /31.91.? <. 89 .5 .9/+.1 5. /31.91.? 5. .,+,-.- - 5. 196,()* 3*1 5 CO2 -+1* - 5. )595.? ( =)=* .*1) 5. *,+.),7+ E+ )*+-,),*+( +.91.5( 5. )*+)+1.),7+ - CO2 5. )*+1*5. 5. 35.+. 3*1 /-,* - (*/.( E5 (1( =;-1,)* 31**). 5 ),11 (*/,)* ,/3,-,+-* 5. +1.-. - CO2 < -,(/,+9<+-* 5. )*+)+1.),7+ - CO2 -+1* - 5. )595.? ( =)=* .*1) 5. *,+.),7+ L. /31.91. ./6,+ .). 5. )*+)+1.),7+ - CO2 < *,+* + -,(*59),7+ 5. )9.5 -3+- - 9+ )*,),+ 89 -,(/,+9< )*+ 5 .9/+* - 5. /31.91. E(* ,/35,). 89 .5 .9/+.1 5. /31.91. -,(/,+9< 5 )*),+ +1 CO2O2 S --9) +*+)( 89 5. *,+.),7+ (93*+ 9+ 31*65/. + )5,/.( )5,-*( < ()*( A5 )*+(,-1.1 5. **11(3,1.),4+ 5 1+-,/,+* -5 ),)5* - C.5,+ ). - 9+ 0 . 9+ '![
INTROD(CCIÓN
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
CONCL(SIÓN
E5 31*)(* **11(3,1.*1,* ( 1(9/ 5. 1.)),7+ - 9+. /*5)95. - R9DP )*+ O2 31*-9) 9+. - PGA5 < *1. - ácido fosfoglicólico? 89 13,-./+ ( =,-1*5,@.-* . ácido glicólico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
'I'LIO/RAFIA
ROBERT M DEVLIN &! FISIOLOGÍA VEGETAL SEGUNDA
EDICION EDITORIAL OMEGA SA BARCELO COLL? `$ OTROS &2 FIFOLOGÍA VEGETAL? MADRID
ED PIRAMIDE SA PAUL B ]EIS? MELVIN S? FULLER & TRATADO DE BOTANICA EDITORIAL CONTINENTAL
