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ra. Susan Hess - Instituto De Química - Facultad De Ciencias )*+ ,-($. / 0 !! " # $2 ! %&' () " " !* ! + 1 )*+ ,-($. / "! "! A mis Padres, hermanos, hijo y familia. )*+ ,-($. / 2 !! " # $2 ! %&' () " " !* ! + 3 )*+ ,-($. / /!" /!" En forma muy especial quiero agradecer a mi profesora patrocinante Dra. Miren Alberdi Lag, quien me permitió desarrollar este trabajo en el laboratorio que ella dirige y de quien siempre recibí constante ayuda tanto teórica como práctica. Quien con su constante apoyo y dedicación contribuyó al desarrollo de esta tesis como también a la formación de un potencial hombre de ciencia. A las profesoras integrantes de mi comisión de tesis Dra. Susan Hess F. que con sus consejos y enseñanzas aportó a este trabajo y con ello permitió que éste llegara a buen término y Dra. Ilona Concha Grabinger por aceptar formar parte de esta comisión. A Nancy Ulloa (Magíster en Ciencias, mención Botánica), quien formaba parte del grupo de trabajo y de quien aprendí mucho, con respecto a la forma de trabajar en el laboratorio y abordar los diferentes problemas. También merecen un reconocimiento importante, los profesores Magdalena Romero del Instituto de Botánica, Juan Carlos Paredes y Joel Pardo del Instituto de Química de esta Universidad, de quienes recibí tanto críticas constructivas como voces de apoyo. Este trabajo fue realizado en el laboratorio de Ecofisiología Vegetal del Instituto de Botánica e Instituto de Química de la Universidad Austral de Chile y fue financiado por los proyectos: DID-UACH S-20002-88 y FONDECYT No 1000610. )*+ ,-($. / 4 !! " # $2 ! %&' () " " !* ! + 5 )*+ ,-($. / 7& 8 7& 8 + 6 7 +. 8 6 . 6 9 6 : ';. * < - #( ". $ ;$ 9 $= :. ! 8 9':. $= $ && 9!&&: 9#: 9>#: . 8 6. ;( ; 8 3?. !&& ; (6 9;@: ; 9A!&&: !&& # >#. ;. '. ;7 ( ># ' . <$8 # $ ' < 9>: ; 9/:. < 9:. -6. ; %6 8 ;6. < 8. ( 8 ; 6 8 $ # (. 8 )*+ ,-($. / B !! " # $2 ! %&' () " " !* ! + ;$ ; + <$ - . -8 $;; ;$ $ ( 889 < $ < C< 6 < << < <; < < < < 6 $$ $ . $ < < C< < 6<< 96 : < << < 9': < << < && 9!&&: 9#: % 9>#: ';. C< < 6<< . 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" # $2 ! %&' () " " !* ! + $ - ( ! ! 2.N ; - ( ;- 9#>GY#: . . = 8 J $ ; ( 0 0 :#9( ":O #& 88 2' $ # 0 9 ; P ( "!;8 0 )$78 $ Q ; L ' D ( ; P$ ;8 L9 / !$; 9 $ . . $ ))7 I@D.$ .$ (: +:R (: (BR . 4D:( 8$ 6 : -$8 % % (: +:R (: (++( : 4 R @0 "$- '$ 20 % % ! ; "%$% '$ 50 ")$% ' 23 )*+ ,-($. / 5& 8 9 8:: ( . 0$ 0 0$ :#9( $ Q A )*+ ,-($. / 24 !! " # $2 ! %&' () " " !* ! + 25 )*+ ,-($. / =& : =& : =&7& #!1! " ! " # !; 6. ; 9ZN.N4: ; 9:. (6 9: ;$ 9;: ># # 9 0N: ># %1 %0 6. 8 1NN 5NN b . 13 < %1 %0 %1 %0 0NNN 5NN b # 1NN 5NN b . 8 ; 9ZN.N4: . # . ;8 $ . -8 ; 01 13 < ; #. %1 %0 %1 %0 1NN b 0NNN b $ ; ; ( # ># (6 9;@: 9 00: 8 ; ># # 9hN.N4: ; $ N.D2%N.B0 ># N.D2%N.B4 # # ># ;@ )*+ ,-($. / 2B !! " # $2 ! %&' () " " !* ! + . 8 . ( . ;. ># # 9:. % 8 !&& %. ># ' . -( 7 7. ; 9 00: ; ; ; # ># 9N 5 <: . - - 9ZN.N4:. 01 <. ; 9hN.N4: # '. 8 ; ;@ 9 \ %N.DFI ZN.N4: 9>:. % 8 %. 8 ; $ ( # . 8 ># 9 00: ( ; !&& 9A!&&:. ( $ ( $= # ># 9 01: $ A!&& $; ' - # >#. A!&& 8 ; ;@. 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" /% ! + "!" "" & %! !" " " > ! "%" ! !* " M !!! " 1"!& 31 )*+ ,-($. / =& : "" " 0 63 67 @N A @2A 1NN N 5 01 0D 13 N 5 01 0D 13 N 5 01 0D 13 5NN 0NNN " 2"! @GA ># 5F.D2aN.F2 B2.4NaN.4N $j B2.10aN.5N #$j B0.4NaN.4N #$ BN.4NaN.4N $j 5F.D2aN.F2 BN.4NaN.4N j 5D.B4aN.NN 5F.NNa1.NN BN.B0aN.B0 5F.D2aN.F2 B1.NNaN.NN $j B3.NNaN.NN #$j B3.4NaN.4N j BN.4NaN.4N j # 5D.F3a0.05 BN.NNa0.NN B0.NNaN.NN # BN.4Na0.4N 5D.0DaN.54 5D.F3a0.05 5B.NNaN.NN 5B.4NaN.4N 5D.4NaN.4N 5D.4NaN.4N 5D.F3a0.05 $ 5F.4NaN.4N $ 5B.NNaN.NN B0.4NaN.4N 55.1Da0.05 # . 0NNN b %1 %0. ; 9hN.N4: 01. 0D 13 < ;. ; ; ( $= 9 02: ' ; 9ZN.N4: ># #. $; 5 01 < - %1 %0 %1 %0 5NN b . 13 < <$ 5NN b $;. # ># ( = 8 - < @< . 8 ; 9hN.N4: >#. 9$ 2: # -8 ; ; 9ZN.N4: 5 %1 %0 0D <I 5 13 < 5NN b < @< . 8 8 $ ; 9ZN.N4: %1 %0 9# >#: 13 < 1NN b 01 < %1 %0 <$ (6 0NNN b 9$ 2: 8 9< Q< .:@ ># ; 9ZN.N4: #. ' - . 6( 0D < 0NNN %1 %0 b 9$ 2: ># . ; . 68 - <$ 5 13 < 9ZN.N4: $. # %1 %0 ; 1NN b N 01 01 13 < # N. 01 0D < )*+ ,-($. / 32 !! " # $2 ! %&' () " " !* ! + (6 ; 9ZN.N4: $; 9N 5: $7 ; $ 01 13 < 0D < =&=& " > %1 %0 ># - 95NN b :. ; -6 8. . 5 < 13 <. %1 %0 90NNN b :. $-8 %1 %0 01 < 8 9 03: # 1NN b 01 < 8 8 ; -6. . ,$ ; 9ZN.N4: -6 $ 13 01 <. %1 %0 . ' 0NNN 5NN b ' <$. ;6 >#. . ; . * < # ; ( -6 ># 9 03: # ; -6. . $; # . 5 < 7. ; ( . 7 < 01 <. < 13 < 9ZN.N4:. $; $7 8 ; 9ZN.N4: -6 < 90D %1 %0 13 <: 1NN b . 8 ( 3& !< " ! @2 A + ! $ @2 $A ! " @A + " @A ! " # & ! - "! . "! " /% ! " ! "" " 0& @KA L ! " /% ! + "!" "" & %! !" " " > ! "%" ! !* " M !!! " 1"!& 33 )*+ ,-($. / =& : "" " 0 63 67 @N A @2A 1NN 5NN 0NNN N 5 01 0D 13 N 5 01 0D 13 N 5 01 0D 13 2 67 @/ / # A 4.DNaN.D5 B.D5a0.01 $j D.BNaN.03 $j 5.F4aN.4B $ B.35aN.0F $ 4.DNaN.D5 4.FFaN.B4 4.F3aN.FB B.0Da0.03 D.20aN.00 j 4.DNaN.D5 5.BFa0.BB B.B2a0.01 5.FFa0.1F B.1BaN.3D 67 @/ / # A 4.50aN.3B 4.B3aN.32 5.53aN.23 5.01aN.23 4.FBaN.3D 4.50aN.3B 3.1DaN.01 3.DBaN.24 4.42aN.3B 4.N5aN.5B 4.50aN.3B 5.2FaN.2N 3.B3aN.13 D.FFaN.F1 3.20aN.4N )*+ ,-($. / 2 $ 67 @/ / # A 1.13aN.23 2.01aN.30 $j 2.3FaN.N5 $j 1.D2aN.15 $ 1.F5aN.00 $ 1.13aN.23 1.42aN.1F $ 1.2BaN.31 $ 1.BDaN.31 $ 2.2DaN.N4 $j 1.13aN.23 1.42aN.4F 1.DBaN.2D 1.BFaN.40 1.D0aN.0D j 67 @/ / # A 1.1BaN.03 1.3NaN.01 1.D2aN.02 1.51aN.02 1.44aN.0B 1.1BaN.03 0.F2aN.NF 0.FBaN.03 1.NBaN.0F 0.F4aN.14 1.1BaN.03 1.44aN.0B $ 0.F0aN.N5 2.B2aN.23 $ 0.52aN.0D 34 !! " # $2 ! %&' () " " !* ! + ( . / ""' "' #F 9 0 M ( 5& < ! ! $ 2 O2 $ + !< ! !" 2 P2 $O! ! " @A + " @A ! " # & ! +- " " "! " /% @#H4I4?A ! "" " !I J ! - "! . "! " /% ! " ! "" " 0& @KA L ! " /% ! + "!" "" & %! !" " " > ! "%" ! !* " M !!! " 1"!& 35 )*+ ,-($. / =& : "" " 0 63 67 @N A @2A 2 O2 $ 1NN 1.5NaN.N0 1.41aN.N3 1.3FaN.N1 1.35aN.N1 1.41aN.ND j 1.5NaN.N0 1.25aN.0N 1.42aN.N4 1.4DaN.N2 1.35aN.N0 1.5NaN.N0 1.53aN.00 1.5DaN.N4 j 1.40aN.N1 1.4FaN.N1 1.34aN.NF 1.2FaN.ND 1.23aN.N3 1.22aN.N5 1.23aN.N3 1.34aN.NF $ 1.12aN.N3 $ 1.3DaN.05 $ 1.5BaN.N1 1.4FaN.N4 1.34aN.NF 1.41aN.N4 1.3BaN.N4 1.30aN.N2 1.52aN.N3 3.F1aN.NB j 4.10aN.N5 j 4.0DaN.02 j 3.DDaN.03 j 3.DFaN.N3 j 3.F1aN.NB j 3.F0aN.ND j 3.D3aN.0B j 3.B3aN.N1 j 3.FFaN.N3 j 3.F1aN.NB $j 4.N1aN.N2 j 3.F3aN.N4 $j 3.DDaN.01 $ 3.5NaN.N3 $j 3.32aN.NB 3.5NaN.N5 $ 3.BFaN.N2 #$ 3.3DaN.N2 #$ 3.34aN.NB # 3.32aN.NB 3.1NaN.11 $ 2.DBaN.N3 $ 2.DBaN.03 $ 3.N4aN.10 $ 3.32aN.NB 3.2DaN.ND 2.FDaN.N4 $ 4.N4aN.0B 2.D2aN.NB $ 5NN 0NNN N 5 01 0D 13 N 5 01 0D 13 N 5 01 0D 13 2 P2 $ O! $ -6 # . . ; ; ;6 9 \ %N.5I ZN.N4: ># 8 ; ; 9 \ N.BI ZN.N4: ; 6. . # ># 9 03: 8 6 $ 8 %1 %0 ># 1NN b . 6 8 913 <: >#. ;6 8 8 %1 %0 5 < 1NN b < 0D <. 13 %1 %0 < # 5NN b . ># ;6 9ZN.N4: 01 < < 13 < 8. $; ; 8 8 . 8 01 0D <. ; 8 13 <. 8 8 $ 0D < # ;6 $ >#. ; 8 0D < - 13 < 5NN 0NNN b %1 %0 ' <$ - ( . $;8 8 ;6 0D <. $; ; ( . 8 913 <: 8 ># # 5 13 < # . $ . + ( $. 8 ; ;6 )*+ ,-($. / 3B !! " # $2 ! %&' () " " !* ! + -6 6 . -6 Q 6 Q ;6 9)Q#QY: ># ; 9hN.N4:. <$ . 8 . %1 %0 01. 0D 13 <. 1NN 0NNN b 0D 13 < 5NN %1 %0 b 9$ 3: # %1 %0 ; 9ZN.N4: 1NN b 13 < # . 6 # ; 0D <. ; %1 %0 $ < 13 < # 0NNN b . $;8 01 <. ; 8 $ 0D 13 9$ 3: ' ; 6 ># # 8 N ' 5 < 8 - -8 ;6 6 Y@9)Q#QY: ># 8 - %1 %0 " 1NN b . 8 %1 %0 ; . 0D < 9ZN.N4:. 5NN 0NNN b 8 ; 5 < 9ZN.N4: 9 04: # -8 8 8 . - . %1 %0 ; 9ZN.N4: 01 < 1NN b I %1 %0 5 < 5NN 0NNN b . ( ; ( $= <$ -6 8 ;6 < ; Y@9YQ#Q): # - ; %6 8 6 9)Q#:@9)Q#QY: 8 -6. 8 - ;6 %6 . ;6. 6 8 >#. ( ; 9ZN.N4: 5 <. $ 01 13 <. %1 %0 %1 %0 5NN 0NNN b # 1NN b ; 6 ; 9ZN.N4: 01 13 <. 8 0D <. 9 04: $ ( ; ; %6 8 $; # (6 . 5 < < 6 = ; $ Y@9YQ#Q): 9ZN.N4: # 1NN 5NN b %1 %0 ; %6 8 ; N 0D <. 13 < 9 04: $ 6 %6 8 ># # 3D )*+ ,-($. / =& : 9ZN.N4: - 7 <$8 >8 ; %6 8 5 < (6 -6 ;6 ( 0$ 0 0 ""' "' $ / !) 9 U "#V8$8;' $ = U Q "W' X B2 2 9 0 A M ( =& Q @,PPA " # " @A + " @A& )*+ ,-($. / 3F !! " # $2 ! %&' () " " !* ! + ! +- " " "! " /% @#H4I4?A ! "" " !I J ! - "! . "! " /% ! " ! "" " 0& @KA L ! " /% ! + "!" "" & %! !" " " > ! "%" ! !* " M !!! " 1"!& "" " 0 63 67 @N A @2A 1NN N 5 01 0D 13 N 5 01 0D 13 N 5 01 0D 13 5NN 0NNN 4N Q 67 @N/ / # A BN.4Fa0.B5 DN.50aN.2F $ 0NN.44aN.B2 $ 0ND.DNaN.4B $ 0NF.N4a0.5D $ BN.4Fa0.B5 5D.B3aN.3F F0.15aN.3F $ 00B.3FaN.B2 $ 00D.N3aN.4B $j BN.4Fa0.B5 F3.3Na0.04 $ 0N3.B4a0.41 $ 011.15aN.24 $j 00F.1BaN.4D $j 0NB.23a0.00 $j DF.0Ba0.N4 $ 0ND.31a0.N1 $ 011.11aN.52 010.32a3.14 # 0NB.23a0.00 j F2.BBaN.44 j 0N2.2Fa0.F0 0NF.F1a0.00 D2.15a2.B4 0NB.23a0.00 j 0N5.14a0.F1 002.20aN.DF F3.FFaN.00 $ BN.32a1.0F $ )*+ ,-($. / =& : B D0 "5Y2'G"5Y2Y@' 0 0 R @G"5Y2Y@' ""' "' $ A !) )*+ ,-($. / 40 !! " # $2 ! %&' () " " !* ! + 41 )*+ ,-($. / ?& : ?& : $ $ $= <$ 7. . ; $ 8 !&& #. $ $ 8 ; =$ !&& <$ 8 !&& ; 6 (6 9;@:. ( (6 !&&. # ># ; - . 8 ; 8 8 9eN.D2: 9=MJ E '. 0FDBI V< . 0FF2I "6C E V<. 1NNN: 6 . 9eN.B0: 68 -. ( ># 901.3^ :. # 9B^ : ' ; ;@ . 8 $ <$8 ;$ ! ( ; N.D2. 7 <$ * 7. ! E H 90FDF:. ; ;@ $= N.B <$8. - . <$ $- . < 8 ># - $8. ;@ * )*+ ,-($. / 42 !! " # $2 ! %&' () " " !* ! + 9N.B0:. ( $ $-8 &&. ( ># # . . ;$ $ !&& - 9! E H. 0FDF. 0FFN. 0FFN$I , . 0FF2I . 0FF3:. < <= 9 : 6 . 6<$ . 8 <$8 $ 8 9; . 1NN0: $ 6<$ ; 9 : 8 9K . 0FF2:. = P 6 # $= $= 9, . 0FF5: ' +. . 8 <$8 8 7 (6 0NNN %1 %0 b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b :. T6U 8. 8 $ $= - ; -6. ;- T6U - ; <$8 < 8 6 . ; 8 -6 8 $ 9$= : ( $ 9#( : 9#$ . 1NN1: # ( $ 7 $ $= . ; . - 9` . 0FFFI ` . 1NNN: ; < . %1 %0 91.5 b G : 1 0N? . 8 %1 %0 7 ; 9#/: 4NN b . ; %1 %0 04NN b . h1N? #/ %1 %0 %1 %0 04NN b N.1 b G ;$ 1 . * . < . 8 9H. 1NNNI #$ . 1NN1: $7 8 ; - ; ; 9` . 0FFF: ! $. 8 7 ; $= . . . $ !&& 9 : 7 6 ; 7. $ . $= . . . $ < 6 %1 %0 90NNN b : 7 ; 45 )*+ ,-($. / ?& : $=. $ ; 6 ; -6 # $; > $; . 6 9> . 0FFB: G 6 6 . - >#. 8 ; * . . < ( ( 0 9 . 0FFF: ; . ;-. . $ - 6 %4%86 ( 0 6 ; 6 8 -6. ;- ; k% Y6 ( . 8 . ;6 6 =7 ;6 $7 %6 8 -6. - = 68 ( 0 8 $= ; %4%86 . ( ;6. -. ( %6 8 ;6 ; ( 9< 5N^ : ( ( 8 + < $ 8 9 . 0FFD . 0FFF: 6 ; 98 86 6:. . . . 8 ; 7 9 8 G : 6 9'%# E # . 1 0FF1: ;. +( ( $. . * . 1N? 9; (:. $ 8 9` . 0FFFI #$ . 1NN1: ! 6 ; 1N? 93? # 02? >#:. $ $ - - <$ ; 8 G . 1 <$ ; 8 7. $ ; 6 ; -8 ; ! $ (. <= 6<$ %6 8 6 8 9'%# E # . 0FF1: . <$ <$ 8 86 9/G!:. 7 <$ $ 6 ;. 7 < TU 6 97- . $: 8 ! H 90FFN:. +. )*+ ,-($. / 4B !! " # $2 ! %&' () " " !* ! + ; 68 -. ; 6 . < <= $ <$8 6 ; H 68. 7 . < ; + 7 6 - 9"7%< E #. 1NNN: $8 -. + 8 ; 6 > 8 7 . ( $ 8 8 7 < 8 $ 6 8 $ . ; - ;6 %6 . $ 8 8 4D )*+ ,-($. / ((:E; ((:E; Adams, W. W. III., Demmig-Adams, B., Barker, D. H., Kiley, S. 1996. Carotenoids and photosystem II characteristics of upper and lower halves of leaves acclimated to high light. Aust. J. Plant Physiol. 23:669-677. Adams, W. W. III., Demmig-Adams, B., Verhoeven, A. S., Barker, D. H. 1995a. ‘Photoinhibition’ during winter stress: Involvement of sustained xanthophyll cycle-dependent energy dissipation. Aust. J. Plant Physiol. 22:261-276. Adams, W. W. 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