Emergencia de las formas vivas: aspectos dinámicos de la biología

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Emergencia de las formas vivas: aspectos
dinámicos de la biología evolutiva
Lorena Caballero
Universidad Nacional Autónoma de México
-7&2
Ciudad de México
CopIt ArXives
2008
CopIt ArXives
Ciudad de México Copyright © 2008 por Lorena Caballero
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Publicado 2008 por CopIt ArXives
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7=^B 89 A9HZ:CF5G ;9B[H=75G D5F5 @CG A975B=GACG 89@ 89G5FFC@@C
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@C 7CAC BI9GHFC A[HC8C 9LD@=75H=JC 9B7CBHF5F9ACG 9B @5 ;9B[H=75
BI9GHF5 9LD@=757=^B D9FC BC 8969ACG 89 5GIA=F@C 5 DF=CF= B= H5A
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9B F95@=858 6I9B5G F5NCB9G D5F5 7F99F EI9 @5 ;9B[H=75 G9 F98I79 5@
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=7 A9H5D<CFG :CF 89J9@CDA9BH5@ A97<5B=GAG H<CI;< GCA9 D9F<5DG A5BM 89J9@CDA9BH5@
A97<5B=GAG K=@@ =B8998 69 ;9B9H=7 A97<5B=GAG CH<9FG K=@@ 69 =FF98I7=6@9 HC ;9B9H=7 GI6
GHF5H9G !H A5M K9@@ HIFB CIH 9J9B =: K9 H5?9 89J9@CDA9BH 5G CIF 9LD@5B5B8IA H<5H
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" F57=5G 5 @5 =BPI9B7=5 89 D9BG58CF9G 7F\H=7CG 7CAC CI@8 M $9KCBH=B
M 9GD97=5@A9BH9 GI DFCDI9GH5 GC6F9 9@ 9EI=@=6F=C DIBHI58C 9B
8CB89 F97CBC79B @5 DCG=6=@=858 89 IB5 9JC@I7=^B BC @=B95@ 9AD=9N5 5 F9
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6=C@C;\5 89@ 89G5FFC@@C D5F5 7CADF9B89F DFC79GCG BC F9GI9@HCG DCF @5 ;9B[H=75
M @5 9JC@I7=^B M 89 5<\ @5 B979G=858 89 =B7CFDCF5F 5 @5 ;9B[H=75 5@ 89G5FFC@@C
M 5 @5 9JC@I7=^B 7CAC IB A=GAC DFC6@9A5 IB DFC6@9A5 89 HF9G 7I9FDCG 9G
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9A6F=^@C;CG 7CAD5F5H=JCG 89@ G=;@C 0!0 YGH5 BC 7CAC H9CF\5 :CFA5@ H=9B9
5BH97989BH9G 9B @5G =895G 89 F=GH^H9@9G M DCGH9F=CFA9BH9 9B 9@ G=;@C 0!0 9F5
D5FH9 =ADCFH5BH9 89 @5G =895G 89 J5F=CG =BJ9GH=;58CF9G 7CAC #5F@ FBGH .CB
59F FBGH 597?9@ %9?9@ +9FF9G 5@:CIF M 8IF5BH9 9@ G=;@C 00 %CF;5B
89 99F /588=B;HCB C@87<A=8H 'G6CFB CBB9F M 5FGH5B; 9BHF9 CHFCG
&C 9G <5GH5 7CB 9@ 89G7I6F=A=9BHC 89 ;FIDCG 89 ;9B9G 5GC7=58CG 7CB
G97I9B7=5G 89 65G9G G=A=@5F9G @5G @@5A585G 75>5G <CA9^H=75G CA9C6CL C
89 89G5FFC@@C M DF=B7=D5@A9BH9 9B F9@57=^B 5 @CG ;9B9G 89 @5 :5A=@=5 CL EI9
9GD97=O75B @5 G9;A9BH57=^B 9B =BG97HCG M D@5B9G 7CFDCF5@9G 9B J9FH96F58CG
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7CBC7=A=9BHC 7CAC D5FH9 =ADCFH5BH9 89BHFC 89 @5 7=9B7=5 AC89FB5 9B @5
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M 89G57H=J5F CHFCG 9B 9@ ACA9BHC 7CFF97HC += 6=9B 9G 7=9FHC EI9
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F9EI9F=ACG G=B 9A65F;C 5@;C AZG EI9 9GH5G 897@5F57=CB9G 56
GC@IH5A9BH9 J9F8589F5G 56GC@IH5A9BH9 J57\5GT +M8B9M F9B9F
9B *C69FH $5 =B79FH=8IA6F9 579F75 89 @CG A975B=GACG CBHC;[B=7CG M @5 A5B9F5 9B @5
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F9GI@H58C 89 @5 B979G=858 89 9B7CBHF5F D5HFCB9G ;9B9F5@9G 89BHFC 89 @5G F989G
7CAD@9>5G 89 @5 HF5M97HCF=5 CBHC;9B[H=75 M J9F 89BHFC 89 [GH5G @CG DFC79GCG
EI9 D9FA=H9B @5 9JC@I7=^B <57=5 CHF5G :CFA5G C =B7@IGC @5 7CBG9FJ57=^B 89
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89G5FFC@@C M @5 9JC@I7=^B G9 9GH56@979 7CB ;F5B :I9FN5 9B @CG D@5BH95A=9BHCG
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7=^B 89 BI9J5G 9GD97=9G C S&CJ98589G JC@IH=J5GT
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H5 D5F5 9BH9B89F 9@ AIB8C 89 @5G :CFA5G += 89BHFC 89 9GH9 DFC79GC 9B 9@
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75@5G 89 H=9ADC M 9GD57=C DCF H5BHC @CG DFC79GCG =BJC@I7F58CG 5D5F9BH9A9BH9
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89BHFC 89 9GH9 A5F7C @CG 8=:9F9BH9G 8=G7IFGCG 9A9F;9B 7CAC :CFA5G :57H=6@9G M
@CG DFC6@9A5G D5F5 9BH9B89F @5 8=J9FG=858 6=C@^;=75 F9GI@H5B H9B9F F9GDI9GH5G
M 5J5B79G 9BHFC 89 @CG D@5BH95A=9BHCG 89 9GH9 HF565>C D5FHC 89 @5 B979G=858
89 7CBG=89F5F 5 @5 H9CF\5 89 @5 7CAD@9>=858 M @CG G=GH9A5G 8=BZA=7CG M5 EI9
BCG 5MI85 5 HF565>5F 7CB @5 AI@H=D@=7=858 89 :9B^A9BCG EI9 A5BH=9B9B 5 @CG
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7CAD@9>=858 M @5 BC @=B95@=858 =B7@IGC 89 @CG DFC79GCG ;9B[H=7CG 9G J=H5@ 9B
9@ 5J5B79 89 BI9JCG 7CBC7=A=9BHCG 9B @5 6=C@C;\5
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" GI 9GHI8=C 89BHFC 89@ A5F7C 89 @5 H9CF\5 89 @5 7CAD@9>=858 M <5 ;9B9F58C
7F97=A=9BHC M 579DH57=^B 9B AI7<5G ZF95G 89 @5 6=C@C;\5 EI9 9G 9G9B7=5@
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@5 5D@=757=^B 89 @5 H9CF\5 89 @CG G=GH9A5G 8=BZA=7CG 89BHFC 89 @5 6=C@C;\5
M 7^AC 9GH56@97=^ @5 :CFA5 9B @5 EI9 8=7<C DFC6@9A5 DC8F\5 G9F HCA58C
7CAC IB DFC6@9A5 6=C@^;=7C (CF 9GH5 F5N^B 9G =ADCFH5BH9 J9F EI9 9@ 75A=BC
9GH56@97=8C 89BHFC 89 @5 J=B7I@57=^B 9BHF9 @5 9JC@I7=^B M 89G5FFC@@C BCG G=FJ9
89 AC89@C 9B @5 6`GEI985 89 F9GDI9GH5G 5 @CG DFC6@9A5G :IB85A9BH5@9G 89 @5
6=C@C;\5
+= @C;F5ACG 7CADF9B89F @CG J\B7I@CG 9BHF9 9@ 89G5FFC@@C M @5 9JC@I7=^B
DCG=6@9A9BH9 89G7I6F=F9ACG @5 A5B9F5 9B @5 EI9 @5G 9GD97=9G C7ID5B IB @I;5F
9B @5 8=J9FG=858 @5 :CFA5 9B @5 EI9 9A9F;9B 9JC@I7=CB5B M G9 9LH=B;I9B M
7^AC @CG A975B=GACG ACF:C;9B[H=7CG 7CBHFC@5B @CG 9GH58CG C 7CBO;IF57=CB9G
DCG=6@9G 89 @5G :CFA5G J=J5G 9 9GH9 AC8C @5 G=GH9AZH=75 DC8FZ 9@56CF5F
G=GH9A5G 89 7CBC7=A=9BHC M CF89B5A=9BHC 89 @5 8=J9FG=858 AZG 79F75BCG 5 @5G
75F57H9F\GH=75G 7CAD@9>5G 89 @CG G=GH9A5G J=JCG
B 9GH9 DFCA9H98CF 75A=BC @5 6=C@C;\5 G9 589BHF5 5 IB G=GH9A5 89 7CBC7=A=9B
HC AZG 79F75BC 5 @CG :9B^A9BCG 6=C@^;=7CG M DCG=6@9A9BH9 75A=B5B8C M F9
75A=B5B8C @C M5 5B858C DC8F9ACG 89 7=9FH5 A5B9F5 7CBH5F @5G A`@H=D@9G
<=GHCF=5G 89 @5 J=85DCFEI9 HC8C @C EI9 GIF;9 89BHFC 89 9@@5 BC DI989 G9F
7CBH58C 89 IB5 GC@5 A5B9F5 [GH5 9G G^@C IB5 89 9@@5G
=;IF5 5F57C@9G 7@5G=O758CG DCF
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34 F=GH^H9@9G ,<9 %9H5D<MG=7G ,F58I7=8C DCF "C<B
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34 X@J5F9N IM@@5 @9B5 S$5 8=J9FG=858 89 @5G :CFA5G J9;9H5@9GT =9B7=5G
*9J=GH5 89 8=:IG=^B 57I@H58 89 =9B7=5G -&% %[L=7C 6F=@>IB=C
34 @J5F9N IM@@5 @9B5 % 9B\H9N 9@@9N5 <5CG GD=BCN5+CHC
( (58=@@5$CB;CF=5 S9B9 F9;@5HCFM B9HKCF? AC89@G :CF D@5BH 89
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34 9@8589 (5HF=7=5 #99G #CCDG M (5I@ % F5?9GO9@8 S%C8I@5F=
HM =B8=J=8I5@=HM 5B8 9JC89JC =B 6IHH9FPM K=B;GT (&+ .C@ 34 9B\H9N #9=BF58 %5F=5B5 ,9G=G 89 A59GHF\5 *989G ;9B[H=75G M 89H9F
A=B57=^B 89 H=DC 79@I@5F 9B F56=8CDG=G H<5@=5B5 -&% 34 FCKB "5A9G 9CVF9M /9GH +75@=B; =B =C@C;M +5BH5 9
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9JC / / &CFHCB CAD5BM -+ 34 <5CG X@J5FC % @85B5 GD=BCN5+CHC 5F7\5 DCB79 89 $9^B 5F5M FFCMC @J5F9NIM@@5 SFCA 9B9 HC PCK9F D5HH9FBG
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B=N=B; DFC79GG D5FH !! ,<9 +DCBH5B9CIG CFA5H=CB C: (9F=C8=7=HM 5B8
H<9 C9L=GH9B79 C: GD=F5@ 5B8 /<CF@98 (5HH9FBGT " H<9CF =C@ 34 CI58M + M 1 CI89F S(<M@@CH5L=G 5G 5 8MB5A=75@ +9@: 'F;5
B=N=B; DFC79GG (5FH !!! ,<9 +=AI@5H=CB C: H<9 ,F5BG=9BH *9;=A9G C:
'BHC;9BMT " H<9CF =C@ 34 DD9F@9=B $_:69F; " S,<9 9J9@CDA9BH C: H<9 $5FJ5@ (=;
A9BH (5HH9FBG =B ,F=HIFIG 5@D9GHF=G 5B8 A6MGHCA5 A9L=75BIAT 8
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" 34 G7CHC F=`;9B5 =J=G=^B 89 @5 B5HIF5@9N5 8=7=CB9G C@=C GD5]5
34 GD=BCG5+CHC 5F@CG(56@C (58=@@5$CB;CF=5 M @9B5 * @J5F9NIM@@5
S 9B9 *9;I@5HCFM &9HKCF? %C89@ :CF 9@@5H9 9H9FA=B5H=CB
8IF=B; F56=8CDG=G H<5@=5B5 @CK9F 9J9@CDA9BH H<5H =G *C6IGH 5B8
*97CJ9FG LD9F=A9BH5@ 9B9 LDF9GG=CB (FCO@9GT ,<9 (@5BH 9@@ .C@
34 9FF5H9F %CF5 "CG[ =77=CB5F=C 89 O@CGC:\5 @=5BN5 8=HCF=5@
%58F=8 34 CC8K=B F=5B CK H<9 @9CD5F8 7<5B;98 =HG GDCHG ,<9 9JC@IH=CB C:
7CAD@9L=HM (F=B79HCB -B=J9FG=HM (F9GG (F=B79HCB &9K "9FG9M 34 CI@8 +H9D<9B " ,<9 GHFI7HIF9 C: 9JC@IH=CB5FM ,<9CFM ,<9 9@?B5D
DF9GG C: 5FJ5F8 -B=J9FG=HM (F9GG 5A6F=8;9 %5GG57<IG9HHG 5B8 $CB
8CB B;@5B8 -+ 34 CI@8 +H9D<9B " ! 5J9 $5B898 +D@5G<9G 5B8 *9P97H=CBG =B &5HIF5@
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34
=7?9M %=7<59@ M @=J9 #=B; ,<9 5A6F=8;9 !@@IGHF5H98 @CGG5FM C:
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=CGG5MG "C<B /=@9M +CBG =B7 34 #5D@5B 5B=9@ M $9CB @5GG -B89FGH5B8=B; &CB@=B95F MB5A=7G
+DF=B;9F .9F@5; &9K 1CF? 34 #5IVA5B +HI5FH ,<9 'F=;=B C: 'F89F +9@: 'F;5B=N5H=CB 5B8 +9@97
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9 !BGH=HIH9 'L:CF8 -B=J9FG=HM DF9GG ! 34 #9@@9F %5?=B; +9BG9 C: $=:9 LD@5=B=B; =C@C;=75@ 9J9@CDA9BH
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-+ 34 #9FGN69F; %=7<9@ M $9K=G /C@D9FH S%97<5B=GAG :CF (CG=H=CB5@ +=;
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34 #CG?C 5FH &9IF5@ B9HKCF?G 5B8 :INNM GMGH9AG 8MB5A=75@ GMG
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(F9BH=79 5@@ !B7
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H<9 8=;=H5@ 5;9 ,<F99 *=J9FG (F9GG &9K 1CF? 34 $9J=+HF5IGG @5I89 @ D9BG5A=BHC G5@J5>9 F9J=5F=CG CB8C 89 7I@
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$ ;I=5FH9 98G $5 9JC@I7=^B 6=C@^;=75 -&% %[L=7C
34 %5F;I@=G $MBB 5B8 #5F@9B9 . +7<K5FHN =J9 ?=B;8CAG B =@@IGHF5H98
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CAD5BM &9K 1CF? 34 %5MF FBGH -B5 @5F;5 7CBHFCJ9FG=5 5FK=B M 9@ 85FK=B=GAC F\H=75
F=>5@JC 5F79@CB5 34 %5MF +D97=9G 7CB79DHG 5B8 H<9=F 5DD@=75H=CB 5D\HI@C 9B *=8@9M
% 98 JC@IH=CB 'L:CF8 *9589FG 'L:CF8 -B=J9FG=HM DF9GG
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34 %98F5BC CBNZ@9N $I=G %CF:C@C;\5 IB7=^B M =BZA=75 89 @CG 'F;5B
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$CIF89G GH9J5 M %5BI9@ 5@7CB= 7CAD=@58CF9G 34 %9=B<5F8H 5BG #@=B;@9F % S %C89@ :CF (5HH9FB CFA5H=CB CB
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=CH97<BC@ 34 %=F5ACBH9G (98FC S$5 ;9CA9HF\5 89 @5G :CFA5G J=J5GT =9B7=5G
*9J=GH5 89 8=:IG=^B 57I@H58 89 7=9B7=5G -&% %[L=7C 6F=@>IB=C
34 %=F5ACBH9G (98FC S(5=G5>9G 9A6F=C@^;=7CG M ;9B9GT =9B7=5G *9
J=GH5 89 8=:IG=^B 57I@H58 89 7=9B7=5G -&% %[L=7C B9FCA5FNC
34 %CF=B 8;5F @ A[HC8C $5 B5HIF5@9N5 89 @5 B5HIF5@9N5 ZH98F5
%58F=8 34 &5J5FFC + 8C@:C 1 @5B75 9FBZB89N 5]CG =@C;9B=5 M 7@5G=O
757=^B 89 @5G 5J9G IB5 BI9J5 J=G=^B 9B &`]9N5F:ZB " M $ ;I=5FH9
98G $5 9JC@I7=^B 6=C@^;=75 -&% %[L=7C
34 &=7C@=G !BHFC8I7H=CB HC BCB@=B95F G7=9B79 5A6F=8;9 -B=J9FG=HM
(F9GG -# 34 'R 5F5 *" S+MGH9A5H=7 9B9F5@=N5H=CB
+D97=9G (FC6@9AT +MGH =C@ =GHCF=75@ 5H9 5B8 H<9
34 (@5H^B =Z@C;CG 8=HCF=5@ (CFF`5 + %[L=7C 34 (=BHCCFF9=5 @5F5 ,<9 CJ5FM C: J9 ;; 5B8 +D9FA 5B8 (F9:CFA5
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34 *5V *I8C@: ,<9 +<5D9 C: $=:9 9B9G 9J9@CDA9BH 5B8 H<9 JC
@IH=CB C: B=A5@ CFAG ,<9 -B=J9FG=HM C: <=75;C (F9GG -+ 34 *C69FH "5GCB +7CHH A6FMC@C;M D=;9B9G=G 5B8 9JC@IH=CB ,5?=B; 9
J9@CDA9BH +9F=CIG@M 5A6F=8;9 +HI8=9G =B (<=@CGCD<M 5B8 =C@C;M
5A6F=8;9 -B=J9FG=HM (F9GG -# 34 +7CHH =@69FH 'D=HN 5B8 *5V S*9J=9K *9GMBH<9H=G=N=B; JC@I
H=CB5FM 5B8 9J9@CDA9BH5@ =C@C;MT 9J9@CDA9BH5@ =C@C;M 5FH=7@9 &C ! 34 +C@[ *=7<5F8 5B8 F=5B CC8K=B +=;BG C: $=:9
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34 +H9K5FH !5B $=:9RG 'H<9F +97F9H ,<9 &9K %5H<9A5H=7G C: H<9 $=J=B;
/CF@8 5AACB8GKCFH< B;@5B8 D9B;I=B 6CC?G 34 +H9K5FH !5B &5HIF9RG &IA69FG =G7CJ9F=B; 'F89F 5B8 (5HH9FBG =B H<9
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34 ,5M@CF +H99J9G M !5B % +IGG9L (5HH9FBG =B (@5BH 9J9@CDA9BH
5A6F=8;9 -B=J9FG=HM (F9GG -+ 34 ,<CADGCB R5F7M / 'B FCKH< 5B8 CFA &9K 1CF? CJ9F 34 .5@9B7=5 XJ5@CG +IG5B5 @ DFC6@9A5 89@ 7CB79DHC 89 9GD97=9 9B &`]9N
5F:ZB " M $ ;I=5FH9 98G $5 9JC@I7=^B 6=C@^;=75 -&%
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34 .M;CHG?M $9J (9BG5A=9BHC M $9B;I5>9 8=HCF=5@ (@[M589 I9BCG
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34 /588=B;HCB &9K (5HH9FBG =B 9B9H=7G 5B8 9J9@CDA9BH
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H=CB5FM 89J9@CDA9BH5@ 6=C@C;MT &5HIF9 JC@ 96FI5FM
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