ISSN 0104-4443 Licenciado sob uma Licença Creative Commons [T] El concepto de presión selectiva y la dicotomía próximo-remoto [I] The concept of selective pressure and the proximate-ultimate dichotomy [A] Gustavo Caponi Doctor en Lógica y Filosofía de la Ciencia, Universidade Estadual de Campinas (Unicamp), profesor del Departamento de Filosofía, Universidade Federal de Santa Catarina (UFSC), becario del CNPq, Florianópolis SC - Brasil, e-mail: gustavoandrescaponi@gmail.com [R] Resumen Las presiones selectivas no son factores ambientales que existan independientemente de los linajes por ellas afectados. Su existencia y su configuración siempre dependen de las alternativas de evolución que el propio linaje genera y acepta; y dicha configuración, necesariamente, se va alterando conforme el propio linaje evoluciona. Aclarar eso es muy importante para desestimar algunas reticencias generadas por la dicotomía próximo-remoto. Esta polaridad continúa siendo un recurso conceptual válido e insustituible para entender la especificidad de la Biología Evolucionaria. Enmendar algunos equívocos generados por Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 198 CAPONI, G. Mayr en su caracterización del concepto de causa remota es una alternativa mejor que la de pensar en abandonar o en relativizar esa distinción.[P] [#] [K] Palabras clave: Laland Kevin. Mayr Ernst. Causa remota. Causa próxima. Presión selectiva. [B] Abstract The selective pressures are not environmental factors that exist independently of the lineages affected by them. Their existence and their configuration always depend on the alternatives of evolution that the lineage itself generates and accepts; and that configuration will be necessarily altered as the lineage itself evolves To clarify this point is important to reject some reserves generated by the proximate-ultimate dichotomy. This conceptual polarity remains a valid and irreplaceable resource for understanding the specificity of Evolutionary Biology. To adjust some misunderstandings generated by Mayr’s characterization of the concept of ultimate cause, is a better alternative than thinking in to live or in to relativize the distinction. [#] [K] Keywords: Laland Kevin. Mayr Ernst. Ultimate cause. Proximate cause. Selective pressure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igo ‘consagrada’, y no ‘enunciada’, porque ella no fue una propuesta original de Ernst Mayr (BEATTY, 1994, p. 346). Fue él, de todos modos, quien, basándose en ella, delineó la distinción entre una Biología Funcional de causas próximas y una Biología Evolucionaria de causas remotas (MAYR, 1961, p. 1502; 1993, p. 93); y es a él que debemos atribuirle el hecho de que dicha dicotomía se haya transformado en una referencia siempre pasible de ser considerada, tanto por biólogos como filósofos de la Biología (Cf. ARIEW, 2003). Ver CALLEBAUT et al., 2007; FOLGUERA, 2011; GILBERT et al., 1996; MARTÍNEZ, 2009; MORANGE, 2011. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto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`'&-'&6&*%/'2!&'-#+'*/$(.*./$&+'(&'A.(#'(&' -/+'+&"&+'A.A/+'%&$("1#$'&$'+!'&A/-!*.@$E D/$%"#".#0&$%&'#'-/'2!&'/*!""&'*/$'&-'&+%!(./'(&-'.08#*%/'&A/-!%.A/' (&' -#+' &V.7&$*.#+' /"7#$.O#*./$#-&+' (&' -#' /$%/7&$.#,' 2!&' &$' -/+' N-%.0/+'#5/+'6!&'8!&+%/'&$'-#'8#-&+%"#'8/"'-#'C./-/71#'IA/-!*./$#".#'(&-' &+#""/--/':FGa\bG,'KLLX,'8E';;c'++6>P'&+#+'/%"#+'*!&+%./$&+'2!&'R#-#$(' &%'#-E':KL;;>'%#0?.)$'8-#$%&#$,'8#"&*&$'A.$*!-#"+&'(&'!$'0/(/'0B+'.$0&(.#%/'*/$'&+/+'%@8.*/+'4#'%&0#%.O#(/+'8/"'-#'a&/"1#'(&'-#'Y&-&**.@$' S#%!"#-':FGa\bG,'KLLX,'8E'M=>'4'2!&'(.&"/$'-!7#"'#'-/'2!&'+&'(./'&$' 3 Los autores del artículo son reconocidos biólogos y filósofos de la Biología que cabe mencionar: Kevin Laland; Kim Sterelny; John Oding-Smee; William Hoppitt; y Tobias Uller. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 199 200 CAPONI, G. --#0#"'d8"/7"#0#'#(#8%#*./$.+%#e':RIJ]SaHS,';<c<,'8E';XT>E'W/"'&+/,' 8#"#'&A#-!#"'3#+%#'2!)'8!$%/'-#'*/08"&+.@$'(&'-/+'&6&*%/+'&A/-!%.A/+' (&'-#'8-#+%.*.(#('6&$/%18.*#'4'-#'*/$+%"!**.@$'(&'$.*3/+'"&#-0&$%&'&V.7&'!$#'$!&A#'4'0&U/"'0#$&"#'(&'&$%&$(&"'-#'#"%.*!-#*.@$'&$%"&'6#*%/"&+' 8"@V.0/+' 4' "&0/%/+,' +&"B' $&*&+#"./' "&&V#0.$#"' !$#' $/*.@$' *-#A&' &$' -#' #"%.*!-#*.@$' (&' &+#' %&/"1#`' &-' 8"/8./' */$*&8%/' (&' !(.$",/ .(0(+)$' vaE'W#"#(.70#')+%&,'#!$2!&'$/'+.$@$.0/'/'&2!.A#-&$%&,'(&'%/(/'-/'2!&' *-B+.*#0&$%&'+&'3#'&$%&$(.(/'8/"'f*#!+#'"&0/%#g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h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omprender es mejor que ajustar I$'!$#'8".0&"#,'8&"/'$/'%/%#-0&$%&'*/""&*%#,'#8"/V.0#*.@$,'-#'(.+%.$*.@$'&$%"&'!$#'C./-/71#'h!$*./$#-'(&'*#!+#+'8"@V.0#+'4'!$#'C./-/71#' 4 5 6 Cf. CAPONI, 2008, p. 138. Cf. CAPONI, 2008, p. 129 ss. Cf. CAPONI, 2008, p. 128. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto IA/-!*./$#".#' (&' *#!+#+' "&0/%#+,' 8!&(&' 8"&+&$%#"+&' &$' A."%!(' (&' -#' (.+%.$*.@$' &$%"&' 6&$@0&$/+' /"7#$1+0.*/+' 4' 6&$@0&$/+' 8/?-#*./$#-&+E' R#'C./-/71#'h!$*./$#-'8!&(&'+&"'#+1'&$%&$(.(#'*/0/'#?#"*#$(/'%/(/'&-' #08-./'*/$U!$%/'(&'(.+*.8-.$#+'?./-@7.*#+'/*!8#(#+'&$'&-'&+%!(./'(&'-#+' *#!+#+'8"@V.0#+'2!&,'#*%!#$(/'#'$.A&-'(&'-/+'/"7#$.+0/+'.$(.A.(!#-&+,' $/+'&V8-.*#$,'$/'+@-/'*@0/'-/+'6&$@0&$/+'A.%#-&+'+&'&$*#(&$#$'&'.$%&7"#$'&$'-#'*/$+%.%!*.@$'i(&+#""/--/j'4'&-'6!$*./$#0.&$%/'iQ+./-/71#j'(&' &+#+'&+%"!*%!"#+P'+.$/'2!&'%#0?.)$'$/+'8&"0.%&$'&$%&$(&"'*@0/'&+/+'/"7#$.+0/+'.$(.A.(!#-&+'.$%&"#*%N#$'*/$'+!'0&(./'?.@%.*/'4'#?.@%.*/E'R#' C./-/71#'IA/-!*./$#".#,'0.&$%"#+'%#$%/,'+&"1#'&+#'/%"#'C./-/71#'/*!8#(#' &$'.$(#7#"'-#+'*#!+#+'"&0/%#+'2!&,'#*%!#$(/'#'$.A&-'(&'-#+'8/?-#*./$&+,' $/+'&V8-.*#"1#$'8/"'2!)'-/+'(.6&"&$%&+'-.$#U&+'(&'+&"&+'A.A/+'8"&+&$%#$'-/+' *#"#*%&"&+'2!&'&6&*%.A#0&$%&'#3/"#'8"&+&$%#$'/'#-7!$#'A&O'8"&+&$%#"/$c. I+%/4'+!8/$.&$(/,'&$%/$*&+,'2!&'-#'C./-/71#'h!$*./$#-'.$*-!4&,' $/'+@-/'#'-#'h.+./-/71#'4'#'-#'C./-/71#'(&-' &+#""/--/':9FkG,';<^K,'8E'=^P' ;<<^,'8E';MM>P'+.$/'2!&'&--#'%#0?.)$'.$*/"8/"#'#'-#'F!%/&*/-/71#E'k'&+%/4' +!8/$.&$(/' %#0?.)$,' 2!&' &-' */$*&8%/' (&' +-4.-/ !(%&)-' 8!&(&' #?#"*#"' %/(/+'-/+'6#*%/"&+'(&'*#0?./'&A/-!%.A/'8"&A.+%/+'8/"'-#'A&"+.@$'$&/+.$%)%.*#'(&'-#'a&/"1#'(&'-#'Y&-&**.@$'S#%!"#-`'(&".A#'7&$)%.*#,'0!%#*.@$,' 0.7"#*.@$,'+&-&**.@$'+&V!#-'4'-#'8"/8.#'+&-&**.@$'$#%!"#-^E'R#+'8"&+./$&+'+&-&*%.A#+'+&"1#$,'&$%/$*&+,'+@-/'!$'%.8/'(&'*#!+#'"&0/%#P'4'"&*/$/*&"'&+%/,'$/'+@-/'.08-.*#'."'!$'8/*/'0B+'#--B'(&'-/'(.*3/'#-'"&+8&*%/'8/"' &-'8"/8./'9#4",'+.$/'2!&'%#0?.)$'.08-.*#'."'!$'8/*/'0B+'#--B'(&'-/'2!&' R#-#$('&%'#-E':KL;;>'%&$1#$'&$'0&$%&'*!#$(/'&+*".?.&"/$'+!' - (!`'&--/+' %#0?.)$'8"/8&$(.&"/$'#'.(&$%.Q*#"'&-'*/$*&8%/'(&'+-4.-/!(%&)- con el de !(.$",/ .(0(+)$8-E' I+%/,' (&' %/(/+' 0/(/+,' &+' #2!1' 0&$/+' .08/"%#$%&' 2!&'+!?"#4#"'-#'.$*/""&**.@$'2!&'*/$--&A#'8"&+&$%#"'#'-#+'*#!+#+'"&0/%#+'(.*.&$(/,'*/0/'-/'3#*&$'R#-#$('&%'#-E':KL;;,'8E';T;K>,'2!&'&--#+'&V8-.*#$'f8/"'2!)'!$'/"7#$.+0/'%.&$&'!$'"#+7/'4'$/'/%"/gE'I+/,'&$'"&#-.(#(,' +&"1#' #+!$%/' (&' *#81%!-/+' (&' -#' C./-/71#' h!$*./$#-' */0/' -#' l&$)%.*#< 7 8 9 Cf. SOBER, 1993, p. 8; WEBER, 2004, p. 3. Cf. CAPONI, 2003, p. 82. Aludo a lo que Mayr (1982, p.631-2) llama Genética de la transmisión: esa genética de Morgan hoy cooptada por la Genética Molecular (Cf. MAYR, 1982, p. 808 ss.). La Genética de Poblaciones sí es, claro, una ciencia de causas remotas: ella es un capítulo de la Biología Evolucionaria (Cf. MAYR, 1982, p. 632; CAPONI, 2003, p. 81). Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun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a&/"1#' (&'-#'Y&-&**.@$'S#%!"#-'+/$'-#+'8/?-#*./$&+,'4'$/'-/+'/"7#$.+0/+,'2!&' &A/-!*./$#$P'&+#'0.+0#'.(&#'8!&(&'+&"'0&U/"'&V8"&+#(#'(.*.&$(/'2!&' -/+' 2!&' &A/-!*./$#$' +/$' -/+' -.$#U&+E' &' 3&*3/,' *!#$(/' &$' -#' a&/"1#' (&'-#'Y&-&**.@$'S#%!"#-'+&'#-!(&'#'8/?-#*./$&+,')+%#+'+/$'&$%&$(.(#+,' $/' +.08-&0&$%&' */0/' &$%.(#(&+' &*/-@7.*#+' :&+' (&*."`' */0/' +!?+.+%&0#+' (&$%"/' (&' !$' &*/+.+%&0#>P' +.$/' */0/' &$%.(#(&+' 7&$&#-@7.*#+' :*/0/'+!?-.$#U&+'(&'!$'-.$#U&'4m/'*/0/'-.$#U&+'2!&'.$*/"8/"#$'/%"/+' +!?-.$#U&+>;LE'F+1,'8#"#'+&"'0B+'&V#*%/+,'&$'-!7#"'(&'(&Q$."'!$#'*#!+#' "&0/%#'*/0/'#2!&--#'*!4/'&6&*%/'+@-/'+&'"&n&U#'&$'&-'8-#$/'8/?-#*./$#-,'(&?&"1#0/+'(&Q$."-#'*/0/'#2!&--#'*!4/'&6&*%/'+@-/'+&'"&n&U#'&$' &-'(&A&$."'(&'!$'-.$#U&E W&"/,'.$%"/(!*."'&+#'8"&*.+.@$'&$'&-'*/$*&8%/'(&'+-4.-/!(%&)-,' -&U/+' (&' /?-.7#"$/+' #' -.0.%#"' +!' #8-.*#*.@$,' 8&"0.%&' #08-.#"-#' 8#"#' */$%&08-#"'6#*%/"&+'&A/-!%.A/+'$/'8"&A.+%/+,'/'$/'(&0#+.#(/'%&$.(/+' &$'*!&$%#,'8/"'-#'A&"+.@$'$&/+.$%)%.*#'(&'-#'a&/"1#'(&'-#'IA/-!*.@$E' h#*%/"&+'2!&,'(.6&"&$%&0&$%&'(&'-#'(&".A#'7&$)%.*#,'-#'+&-&**.@$'$#%!"#-,'-#'+&-&**.@$'+&V!#-'4'&-'#.+-#0.&$%/'7&/7"BQ*/,'8!&(&$'%&$&"' &6&*%/+'A&".Q*#?-&+'0B+'#--B'(&-'-.$#U&'*/08!&+%/'8/"'!$#'N$.*#'8/?-#*.@$'/'&+8&*.&E'a#-'&-'*#+/'(&'-/+'*/$+%"&5.0.&$%/+'/$%/7&$)%.*/+' 3/4' .$A/*#(/+' 8/"' -#' C./-/71#' IA/-!*./$#".#' (&-' &+#""/--/;; ellos 8!&(&$'#*%!#"'+.0!-%B$&#0&$%&'+/?"&'(.+%.$%/+'-.$#U&+,'#!$'*!#$(/' )+%/+'+&'&$*!&$%"&$'"&8"/(!*%.A#0&$%&'#.+-#(/+'4,'8/"'&+/,'.08/+.?.-.%#(/+'(&'A&"+&'#6&*%#(/+'8/"'!$#'0.+0#'8"&+.@$'+&-&*%.A#,'/'8/"' 10 11 Cf. CAPONI: 2008, p. 128; 2011b, p. 98. Cf. ARTHUR, 2011, p. 339. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto !$' 0.+0/' 8"/*&+/' (&' (&".A#' 7&$)%.*#' /' (&' #.+-#0.&$%/' 7&/7"BQ*/' :DFW]SH,' KLL<,' 8E' XX>E' b$' 0.+0/' */$+%"&5.0.&$%/' /$%/7&$)%.*/,' (.6&"&$%&0&$%&' (&' -/+' 6#*%/"&+' &A/-!%.A/+' 8"&A.+%/+' 8/"' &-' $&/(#"o.$.+0/,' 8!&(&' 3#*&"+&' +&$%."' +.0!-%B$&#0&$%&,' &$' (.6&"&$%&+' -.$#U&+'+!8"#8/?-#*./$#-&+P'4'&+'#+1'2!&,'&$%"&'/%"#+'*/+#+,'&--/+'8!&(&$'8"&+&"A#"'*#"#*%&"&+'3/0@-/7/+'&$'(/+'&+8&*.&+'(.6&"&$%&+'*!4/' #$*&+%"#-'*/0N$'+&'&$*!&$%"#'Q-/7&$)%.*#0&$%&'0!4'-&U/+'(&'#0?#+' :F9bS Y]S,'KLLT,'8E'KMc>E W&"/' &$%&$(&"' -#' (.*/%/01#' 8"@V.0/["&0/%/' &$' %)"0.$/+' (&' !$#'(.+%.$*.@$'&$%"&'6#*%/"&+'*#!+#-&+'2!&'#*%N#$'&$'4'+/?"&'-/+'/"7#$.+0/+'4'6#*%/"&+'*#!+#-&+'2!&'#*%N#$'&$'4'+/?"&'-/+'-.$#U&+'%#0?.)$' 8"&+&$%#'!$#'A&$%#U#'&8.+%&0/-@7.*#'#(.*./$#-`'-#'(&'&A.(&$*.#"'-#'&+%"&*3#'"&-#*.@$'2!&'.$&A.%#?-&0&$%&'(&?&'&V.+%."'&$%"&'#0?/+'@"(&$&+' *#!+#-&+E'D/$6/"0&'-/'(&*1#'S.*/-#.'\#"%0#$$':;<=X,'8E'T<>,'f-#'A.(#' (&'-#'&+8&*.&'$/'&+'A.(#'al lado'(&'-#'(&'-/+'.$(.A.(!/+'/'1()!9.'(&')+%/+,'+.$/'&V*-!+.A#0&$%&'!$#'A.(#'en'&--/+,'4'."0&'&$'&--/+gP'4'&+'8/"'-#' 0&(.#*.@$'(&'&+/+'.$(.A.(!/+,'2!&'-&+'(#$'&V.+%&$*.#'4'0#%&".#-.(#(,' 2!&'-/+'-.$#U&+'+&'+/0&%&$,'%#$%/'#'-#+'&V.7&$*.#+'/"7#$.O#*./$#-&+'(&' -#'/$%/7&$.#,'2!&'+/$'"&+8/$+#?-&+'(&'-/+'*/$+%"&5.0.&$%/+'/$%/7&$)%.*/+':&+%!(.#(/+'8/"'-#'C./-/71#'IA/-!*./$#".#'(&-' &+#""/--/>,'*/0/' #'-#+'&V.7&$*.#+'&*/-@7.*#+,'2!&'+/$'"&+8/$+#?-&+'(&'-#+'8"&+./$&+'+&-&*%.A#+':(&+%#*#(#+'8/"'-#'a&/"1#'(&'-#'Y&-&**.@$'S#%!"#->E'F+1,'4'*#+.' 8#"#6"#+&#$(/' #' \#"%0#$$,' 8/("1#0/+' %#0?.)$' (&*."' 2!&' &-' /"(&$' (&' -#+' *#!+#+' "&0/%#+' $/' &+' !$' /"(&$' *#!+#-' 8#"#-&-/' /' #$%&"./"' #-' /"(&$'(&'-#+'*#!+#+'8"@V.0#+`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ev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 203 204 CAPONI, G. 8!&(&$' A&".Q*#"+&' *!#$(/' */$+.(&"#0/+' #' -#+' 8/?-#*./$&+' &$' %#$%/' 2!&'-.$#U&+P'4'#-7/'#$B-/7/'%#0?.)$'/*!""&'*/$'-/+'*/$+%"&5.0.&$%/+' /$%/7&$)%.*/+`'"&6&"."+&'#'!$'.(.:&/(,/0-/&2(!)-/1(/8-!$-,)(./-/.(!/.(0(++$&' nadas':9FkSFG 'Y9Ha\'&%'#-E,';<^T,'8E'K==>'.08-.*#'"&6&"."+&'#'!$' 6&$@0&$/' 2!&' &V*&(&' &-' $.A&-' (&-' /"7#$.+0/' .$(.A.(!#-' :DFW]SH,' KLL^,' 8E' ;Mc>E'F!$' *!#$(/,' .$+.+%/,' &+/+' +&+7/+' 4' &+#+' 8"&+./$&+' +&-&*%.A#+' +@-/' /*!""#$' 8/"2!&' -#' 8"/8.#' &V.+%&$*.#' (&' -/+' /"7#$.+0/+' .$(.A.(!#-&+'&+%B'+/0&%.(#'#'&V.7&$*.#+'/"7#$.O#*./$#-&+'4'&*/-@7.*#+' *!4/+'&6&*%/+'+&'3#*&$'+&$%."'&$'&-'(/0.$./'(&'-#+'*#!+#+'8"@V.0#+,' .08#*%#$(/'-!&7/'&$'&-'8-#$/'(&'-/+'-.$#U&+':DFW]SH,'KLL^,'8E';M^>P'4' &+%/'N-%.0/,'-/'+!?"#4/,'&+'0!4'.08/"%#$%&'8#"#'(.+*!%."'#-7!$#+'(&' -#+'*!&+%./$&+'8-#$%&#(#+'8/"'R#-#$('&%'#-E':KL;;>E a#-'*/0/'&-'8"/8./'9#4"':;<=;,'8E';TLK[;TLM>,'R#-#$('&%'#-E':KL;;,' 8E';T;K>'$/'+@-/'%.&$(&$'#'.(&$%.Q*#"'&-'*/$*&8%/'(&'+-4.-/!(%&)- con el (&'8"&+.@$'+&-&*%.A#,'+.$/'2!&'#(&0B+'%#0?.)$'%.&$(&$'#'*/$6!$(."'#' &+#+'8"&+./$&+'+&-&*%.A#+'*/$'-#+'.$%&"#**./$&+'/'&V.7&$*.#+'&*/-@7.*#+' 2!&'&A&$%!#-0&$%&'8!&(&$'+!+*.%#"'(.*3#+'8"&+./$&+;KE'I""/"')+%&'2!&' &2!.A#-&'#'*/$6!$(."'-#'-!*3#'8/"'-#'&V.+%&$*.#,'6&$@0&$/'8!"#0&$%&' &*/-@7.*/,'*/$'-#'8"/8.#'+&-&**.@$'$#%!"#-,'6&$@0&$/'&A/-!%.A/'8/"'#$%/$/0#+.#P'4'&""/"'2!&'#(&0B+'.08-.*#'8#+#"'8/"'#-%/'-#'(.6&"&$*.#'&V.+%&$%&'&$%"&'(/+'/8&"#*./$&+'*/7$.%.A#+'0!4'(.+%.$%#+`':;>'&-'#$B-.+.+'(&' -#+'.$%&"#**./$&+'&*/-@7.*#+'2!&'#6&*%#$'#'-/+'/"7#$.+0/+'.$(.A.(!#-&+' 4'#'-/+'7"!8/+'2!&'&--/+'*/$6/"0#$,'4'2!&'8/"'-/'%#$%/'+&'.$+*".?&$'&$' &-'(/0.$./'(&'-#+'*#!+#+'8"@V.0#+P'4':K>'#-'.(&$%.Q*#*.@$'(&'-/+'&6&*%/+' &A/-!%.A/+'2!&'&+#+'.$%&"#**./$&+'8!&(&$'%&$&"'&$'-/+'-.$#U&+'(&'&+/+' +&"&+'A.A/+,'4'2!&'+1'+&'.$+*".?&$'&$'&-'(/0.$./'(&'-#+'*#!+#+'"&0/%#+' :DFW]SH,'KL;L#,'8E'^^[^<>E I+'(&*."`'R#-#$('&%'#-E :KL;;,'8E';T;K>'*/$6!$(&$'&-'#$B-.+.+'6!$*./$#-'#!%/&*/-@7.*/,'2!&'8!&(&'+&"'3&*3/'&$'*#08/'3#+%#'8/"'$#%!ralista adepto a la teoría del diseño inteligente,' */$' -#' &V8-.*#*.@$' +&-&**./$#-'(#"o.$.#$#'2!&,'%/0#$(/'&$'*!&$%#'&+/+'#$B-.+.+'&*/-@7.*/+,' #8!$%#'#'&V8-.*#"'-#'3.+%/".#'(&'!$'-.$#U&'4'-/+'&+%#(/+'(&'-/+'*#"#*%&"&+' 12 Cf. CAPONI, 2008, p. 129 ss. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto 2!&' &+&' -.$#U&' 8"&+&$%#;ME' D-#"/,' 8#"#' &-' $#%!"#-.+%#' (#"o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resiones selectivas y causación recíproca R#' *#!+#*.@$' "&*18"/*#' +&"1#' -#' 0!%!#' (&%&"0.$#*.@$' &$%"&' #2!&--/' 2!&' +&' &$%.&$(&' */0/' 6#*%/"' "&0/%/' *!#$(/' +&' 3#?-#' (&' 13 14 Cf. FRANCIS, 1990, p. 410; CAPONI, 2010b, p. 58. Cf. LALAND et al., 2011, p. 1512. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 205 206 CAPONI, G. 8"&+.@$'+&-&*%.A#'4'-/+'*#"#*%&"&+'(&-'8"/8./'-.$#U&'(&'+&"&+'A.A/+'2!&' &+%B' +/0&%.(/' #' &--#E' b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f*!#$(/'!$' *#"B*%&"'&A/-!*./$#'8/"'+&-&**.@$'.$%&"+&V!#-gE'I$'&+%&'*#+/,'+&'$/+' (.*&,'f&-'6#*%/"'+&-&*%.A/'&+,')-'0.+0/,'!$'*#"B*%&"'&$'&A/-!*.@$g`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lectiva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f. BRANDON, 1990, p. 65. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto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`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ev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 207 208 CAPONI, G. %&08/"#(#+E'W&"/'&+/'(&8&$(&"B'0&$/+'(&'-#'*"!(&O#'(&'-#+'+&2!1#+' 2!&' (&-' +.08-&' 3&*3/' (&' 2!&' -/+' *#0?./+' /*!"".(/+' &$' &-' 8//-' 7&$)%.*/'(&'-#'8/?-#*.@$,'7&$&"&$,'/'$/,'-#'8/+.?.-.(#('(&'2!&'+!"U#$' A#".#$%&+'%/(#A1#'0B+'"&+.+%&$%&+E'a/(/'8"/*&+/'+&-&*%.A/,'&$'-#'0&(.(#'&$'2!&')-'#-%&"#'-#'8/?-#*.@$'&$'-#'2!&'/*!""&,'#-%&"#'%#0?.)$'-#+' "&7-#+'(&'-#'8"/8.#'+&-&**.@$E W&"/'3#4'0B+`'-#'.08/+.?.-.(#('(&'(&Q$."'8"&+./$&+'+&-&*%.A#+'*/$' .$(&8&$(&$*.#'(&-'-.$#U&'2!&'#'&--#+'+&'+/0&%&"1#,'$/'+@-/'%.&$&'2!&'A&"' */$'-#'&V.+%&$*.#'/'*/$'-#'8/+.?.-.(#('(&'8"/(!*."'&+%#(/+'A#".#$%&+'(&'!$' *#"B*%&"'2!&'(.*3/'-.$#U&'8"&+&$%&E'I-'+!"7.0.&$%/'(&'8"&+./$&+'+&-&*%.A#+' %.&$&'2!&'A&"'%#0?.)$'*/$'&-'0/(/'&$'&-'2!&'&+&'-.$#U&'+&'"&-#*./$#,'&$' *#(#'0/0&$%/'(&'+!'3.+%/".#'&A/-!%.A#,'*/$'&-'#0?.&$%&'&$'&-'2!&'+!+' 8/?-#*./$&+'0&("#$E'b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k#'-/'(&*1#'9/$/(':;<c;,'8E';M<[;XL>` ]%"#'(.Q*!-%#('8#"#'-#'%&/"1#'+&-&*%.A#'8"/A.&$&'(&'3#?&"'+.(/'(&0#+.#(/' #' 0&$!(/' */08"&$(.(#,' /' 8"&+&$%#(#,' */0/' (&8&$(.&$%&' (&' -#+'+/-#+'*/$(.*./$&+'(&-'0&(./'&V%&"./"'*/0/'#7&$%&+'(&'+&-&**.@$E' p+%#'&+'!$#'*/$*&8*.@$'*/08-&%#0&$%&'&""@$&#E'W/"2!&'-#+'8"&+./$&+' (&'+&-&**.@$'2!&'&U&"*&$'+/?"&'-/+'/"7#$.+0/+'-#+'*/$(.*./$&+'&V%&"$#+,'$/'+/$'&$'$.$7N$'*#+/'.$(&8&$(.&$%&+'(&'-#+'8&"6/"0#$*&+'%&-&/$@0.*#+'*#"#*%&"1+%.*#+'(&'-#'&+8&*.&E']"7#$.+0/+'(.6&"&$%&+'A.A.&$(/' &$'&-'0.+0/',$+5&/(+&0":$+&,'%.&$&$'*/$'-#+'*/$(.*./$&+'&V%&"$#+':*/08"&$(.(/+'-/+'(&0B+'/"7#$.+0/+>,'.$%&"#**./$&+'0!4'(.6&"&$%&+'4'&+8&*1Q*#+E'Y/$'&+#+'.$%&"#**./$&+'&+8&*1Q*#+,'&$'8#"%&'escogidas por el Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto 0.+0/'/"7#$.+0/,'-#+'2!&' (&%&"0.$#$'-#'$#%!"#-&O#'4'-#'/".&$%#*.@$' (&' -#' 8"&+.@$' (&' +&-&**.@$' 2!&' )-' +!6"&E' .7#0/+' 2!&' -#+'condiciones iniciales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`' -#+' 8"&+./$&+' +&-&*%.A#+' +@-/' +&' */$Q7!"#$' #' 8#"%."' (&-' 8"/8./' -.$#U&' &$'&A/-!*.@$P'&$'$.$7N$'*#+/'&--#+'(&8&$(&$'&V*-!+.A#0&$%&'(&'*/$(.*./$&+'&V%&"$#+'#')-E'W/"'&-'*/$%"#"./,'&+#+'8"&+./$&+'(&8&$(&$'(&' -/'2!&'-#'3.+%/".#'&A/-!%.A#'8#+#(#'4#'3.O/'(&'&+&'-.$#U&P'4'%#08/*/' .08/"%#' #2!1' 2!&' 9/$/(' :;<c;>,' ?.@-/7/' (&' -#?/"#%/"./' #-' Q$,' 3#4#' !+#(/' (&' 0/(/' .$*/""&*%/' &-' %)"0.$/' f$.*3/' &*/-@7.*/g,' */$6!$(.)$(/-/'*/$'-/'2!&'+.08-&0&$%&'3!?.&+&'*/$A&$.(/'--#0#"'f#0?.&$%&' &V%&"$/g;=E'R/'2!&'.08/"%#'#2!1'&+'+!'*-#"#'A.+.@$'(&'2!&'&-'f#0?.&$%&' +&-&*%.A/g'$/'+&'.(&$%.Q*#'*/$'&+&'f#0?.&$%&'&V%&"$/g;c'4'(&8&$(&'(&' !$#'&+%"#%&7.#'#(#8%#%.A#'8"&A.#0&$%&'(&Q$.(#E Construcción de nichos y plasticidad fenotípica W&"/' *-#"/,' *!#$(/' R#-#$(' &%' #-E' :KL;;,' 8E' ;T;X>' +&' "&Q&"&$' #' -#' f*/$+%"!**.@$'(&'$.*3/+g,'&--/+'&+%B$'#-!(.&$(/'#'#-7/'(.+%.$%/'(&'&+#' (&%&"0.$#*.@$' &A/-!%.A#' (&' -#+' 8"&+./$&+' +&-&*%.A#+' #' -#' 2!&' #-!(1#' 9/$/(E'Y.7!.&$(/'&$'&+/'#'R&o/$%.$':;<c<>,'&--/+'+&'&+%B$'"&Q".&$(/' #'-/+'&6&*%/+'8"/(!*.(/+'&$'&-'#0?.&$%&'8/"'&-'8"/8./'#**./$#"'(&'-/+' /"7#$.+0/+'.$(.A.(!#-&+,'/'(&'+!+'8/?-#*./$&+':&$'+&$%.(/'&*/-@7.*/>E' I+/+'&6&*%/+,'+&'$/+'(.*&'*/$'%/(#'"#O@$,'0/(.Q*#$'&-'#0?.&$%&'(&'6/"0#' +.7$.Q*#%.A#' 4' &+/' 8!&(&' #*#""&#"' 0/(.Q*#*./$&+' .08/"%#$%&+' &$' 16 17 Cf. BRANDON, 1990, p. 67. Cf. BRANDON, 1990, p. 68. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun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f*#!+#*.@$'"&*18"/*#g'2!&'&+'.$3&"&$%&'#'&+#+'8"&+./$&+E I+'A&"(#(,'8/"'/%"#'8#"%&,'2!&'-/+'+&"&+'A.A/+'$/'8"&+&$%#$'6&$/%.8/+'"17.(/+'*!4/'(&+#""/--/'+&#'.$(&8&$(.&$%&'(&'-#+'*/$(.*./$&+'(&-' #0?.&$%&'&$'&-'2!&'&--/+'*"&*&$E'a/(/'6&$/%.8/'&+'0B+'/'0&$/+'8-B+%.*/' :JIYa[ICIG\FG ,'KLLM,'8E';L>,'0B+'/'0&$/+'+&$+.?-&'#'-/+'(.6&"&$%&+'6#*%/"&+'2!&'.$*.(&$'&$'+!'/$%/7&$.#':]kF9F,';<^T,'8E'KK>P'4'&+/,' */0/'-/'#8!$%#$'R#-#$('&%'#-E':KL;;,'8E';T;X>'8!&(&'%"#&"'*/$+&*!&$*.#+' &*/-@7.*#+'.08/"%#$%&+,'*#8#*&+'(&'#*#""&#"'*#0?./+'&A/-!%.A/+E'I-'%#0#5/'0&(./'(&'-/+'*/$&U/+'(&'!$#'8/?-#*.@$'*!4#+'%&08/"#(#+'(&'*"1#+,'#' -/'-#"7/'(&'A#".#+'7&$&"#*./$&+,'*/.$*.(#'*/$'-#'+/?"&#?!$(#$*.#'(&'!$' Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto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`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`'-/' 0.+0/'3#?1#'/*!"".(/'*/$'#2!&--#+'3.&"?#+'2!&,'8/"'8/(&"'#?+/"?&"'4' #-0#*&$#"'0B+'#7!#'(&-'+!&-/,'#7"#A#?#$'-/+'&6&*%/+'(&'-#'+&2!1#'2!&' &--#+'0.+0#+'+!6"1#$E 18 Cf. LALAND et al., 2011, p. 1514. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 211 212 CAPONI, G. I$'+1$%&+.+`'*!#$(/'#Q"0#$'2!&'f-/+'6#*%/"&+'"&0/%/+'$/'+/$'#%".?!%/+'&+%#?-&+'4'#!%@$/0/+'(&-'#0?.&$%&g,'4'2!&'f&--/+'.$*-!4&$'-B?.-&+' #%".?!%/+' (&' -/+' 8"/8./+' /"7#$.+0/+' 4' +!+' *#0?.#$%&+' &6&*%/+' +/?"&' &-' #0?.&$%&g,'R#-#$('&%'#-E':KL;;,'8E';T;X>'&""#$'(/+'A&*&+E'I""#$,'&$'8".0&"' -!7#",'8/"2!&,'.$%&"8"&%#$(/'.$*/""&*%#0&$%&'&-'*/$*&8%/'(&' !(.$",/.(0(+' tiva,'8-#$%&#$'!$#'(.Q*!-%#('.$&V.+%&$%&`'!$#'8"&+.@$'+&-&*%.A#,'*/0/'4#'-/' (.U&'4'-/'"&8&%1,'$/'&+'$!$*#,'&$'$.$7N$'*#+/,'!$'6#*%/"'#0?.&$%#-'&+%#?-&' 4'#!%@$/0/E'b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onsideraciones finales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ev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto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c,'8E'c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eferencias F9bS Y]S,'GE'The changing role of the embryo in evolutionary thought. D#0?".(7&`'D#0?".7&'b$.A&"+.%4'W"&++,'KLLTE FGHIJ,'FE'I"$+%'9#4"e+'d!-%.0#%&m8"/V.0#%&e'(.+%.$*%./$'"&*/$+.(&"&('#$('"&*/$+%"!*%&(E' !"#"$%&'&()!#"*"+)%,'AE;^,'8E'TTM[T=T,'KLLME FGa\bG,' JE !,*-.& -/01%"*& ,2.& -3"#45!"2E' D#0?".(7&`' D#0?".(7&' b$.A&"+.%4'W"&++,'KLLXE 19 Cf. WEST-EBERHARD, 2003, p. 11. Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 213 214 CAPONI, G. FGa\bG,'JE'Evolution`'#'(&A&-/80&$%#-'#88"/#*3E']V6/"(`'J.-&4[C-#*qo&--,' KL;;E CIFaak,' rE' W"/V.0#%&mb-%.0#%&' .+%.$*%./$' .$' %3&' 9!-%.8-&' D#"&&"+' /6' I"$+%' 9#4"E' !"#"$%&'&()!#"*"+)%,'AE'<,'8E'MMM[MT=,';<<XE CGFS ]S,'GE'Adaptation and environmentE'W".$*&%/$`'W".$*&%/$'b$.A&"+.%4' W"&++,';<<LE DFRRICFba,'JE'a3&'/"7#$.+0.*'+4+%&0+'#88"/#*3`'IA/[ &A/'#$('%3&'+%"&#0-.$.$7'/6'%3&'$#%!"#-.+%.*'#7&$(#E'H$`'YF9Y]9,'GEP'CGFS ]S,'GE':I(E>E'Integrating evolution and developmentE'D#0?".(7&`'9Ha'W"&++,'KLLcE'8E'KT[<KE DFW]SH,' lE' IV8&".0&$%/+' &$' ?./-/71#' &A/-!%.A#`' st!)' %.&$&$' &--/+' 2!&' -/+' /%"/+'$/'%&$7#$u'Episteme,'AE;=,'8E'=;[<c,'KLLME DFW]SH,' lE' R#' #08-.#' #7&$(#' (&' -#' h.-/+/61#' (&' -#' D.&$*.#E' 6!#"*"7,,' AE' ^, $E'K,'8E'cT[^K,'KLLcE DFW]SH,'lE'R#'C./-/71#'&A/-!*./$#".#'(&-'(&+#""/--/'*/0/'*.&$*.#'(&'*#!+#+' "&0/%#+E'8!$2"*&6!#"*97:"*,'AE';L,'$E'KL,'8E';K;[;XK,'KLL^E DFW]SH,' lE' &+#""/--/,' *#!+#+' "&0/%#+' &' 3.+%/".#' $#%!"#-E' Principia,' AE' ;M, $E';,'8E'K<[TL,'KLL<E' DFW]SH,' lE' h!$*.@$,' #(#8%#*.@$' 4' (.+&5/' &$' C./-/71#E& 8!$2"*& 6!#"*97:"*, AE';K,'$E'KX,'8E'c;[;L;,'KL;L#E DFW]SH,'lE'F$B-.+.+'6!$*./$#-&+'4'&V8-.*#*./$&+'+&-&**./$#-&+'&$'C./-/71#`'!$#' *"1%.*#'(&'-#'*/$*&8*.@$'&%./-@7.*#'(&-'*/$*&8%/'(&'6!$*.@$E'Ideas y Valores, $E';XM,'8E'T;[cK,'KL;L?E DFW]SH,'lE'F8"/V.0#vw/'&8.+%&0/-@7.*#'x'?./-/7.#'&A/-!%.A#'(/'(&+&$A/-A.0&$%/E'H$`'FCGFSaIY,'WE':I(E>E'6!#"*"7,&.,&0!"#"$!,E'W/"%/'F-&7"&`'F"%0&(,' KL;;#E'8E'K;;[KKME' DFW]SH,'lE']+'%BV/$+'*/0/'.$(.A1(!/+E'H$`'YaIhFS],'JEP'WID\RHkI,'9E' :I(E>E'6!#"*"7,&-&;!*591!,&.,& !"#"$!,E'Yw/'W#!-/`'b$.A&"+.(#(&'W"&+?.%&".#$#' 9#*q&$O.&,'KL;;?E'8E'c;[;;KE Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 El concepto de presión selectiva y la dicotomía próximo-remoto h]RlbIGF,'lE'I$6/2!&+'4'(&+&$6/2!&+'(&'-/+'8"/7"#0#+'(&'.$A&+%.7#*.@$'(&' -#'C./-/71#'IA/-!%.A#'(&-' &+#""/--/E'Ludus Vitalis,'AE';<,'$E'MT,'8E'MKT[MM;,'KL;;E' hGFSDHY,' GE' D#!+&+,' 8"/V.0#%&' #$(' !-%.0#%&E' !"#"$%& '& ()!#"*"+)%,' AE' T, 8E'XL;[X;T,';<<LE lHRCIGa,'YE'&%'#-E'G&+4$%3&+.O.$7'IA/-!%./$#"4'#$(' &A&-/80&$%#-'C./-/74E' <-3-#"+/-25,#& !"#"$%,'AE';cM,'8E'MTc[McK,';<<=E \FGa9FSS,'SE'OntologíaE'9)V.*/`'h/$(/'(&'D!-%!"#'I*/$@0.*#,';<=XE'AE'TE' RFRFS ,'ZE'&%'#-E'D#!+&'#$('&y&*%'.$'?./-/74'"&A.+.%&(`'H+'9#4"e+'W"/V.0#%&[ b-%.0#%&' .*3/%/04'Y%.--'b+&6!-u'Science,'AE'MMX,'8E';T;K[;T;=,'KL;;E RIJ]SaHS,' GE' R#' #(#8%#*.@$E' H$`' YDHISaHhHD'F0&".*#$' :I(E>E Evolución. C#"*&-/$#`'R/+'R.?"/+'(&'H$A&+%.7#*.@$'z'D.&$*.#,';<c<E'8E';M<[;TKE':R.?"/'8!?-.*#(/'&$'-#'&(.*.@$'&+8&*.#-'(&'-#'"&A.+%#'Y*.&$%.Q*'F0&".*#$>E' 9FGa{SI|,'9E'R/+'*/$+%"&5.0.&$%/+'(&-'(&+#""/--/'4'-#'.$%&7"#*.@$'&A/(&A/`'8"&*.+./$&+'4'(.+%.$*./$&+'&$'%/"$/'#-'%&0#gE'=:5,& !"#9$!:,&>"#"/0!,2,? AE';X,'8E';T;[;=^,'KLL<E 9FkSFG ' Y9Ha\,' rE' &%' #-E' &A&-/80&$%#-' */$+%"#.$%+' #$(' &A/-!%./$E' The @4,15-1#%&A-3!-B&"C& !"#"$%,'AE'=L,'$E'M,'8E'K=T[K^c,';<^TE 9FkG,'IE'D#!+&'#$('&y&*%'.$'C./-/74E'Science,'AE;MX,'8E';TL;[;TL=,';<=;E 9FkG,'IE'The growth of biological thoughtE'D#0?".(7&`'\#"A#"('b$.A&"+.%4' W"&++,';<^KE 9FkG,' IE' W"/V.0#%&' #$(' !-%.0#%&' *#!+#%./$+E' !"#"$%& '& ()!#"*"+)%,' AE' ^, 8E'<M[<X,';<<ME' 9FkG,'IE'Así es la biologíaE'9#(".(`' &?#%&,';<<^E 9]S] ,'rE'D#&,E,1&%&#,&2-:-*!.,.E'9/$%&'}A.-#`'D#"#*#+,';<c;E 9]GFSlI,'9E'J3#%'o.--'"&+!-%'6"/0'%3&'.$%&"#*%./$'?&%o&&$'6!$*%./$#-'#$(' &A/-!%./$#"4'?./-/74E'854.!-*&!2&;!*5"1%&,2.&()!#"*"+)%&"C& !"#"$!:,#&,2.& !"/-.!:,#&8:!-2:-*,'AE'XK,'8E'=<[cX,'KL;;E Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013 215 216 CAPONI, G. ]kF9F,'YE'The ontogeny of informationE'D#0?".(7&`'D#0?".(7&'b$.A&"+.%4' W"&++,';<^TE Y]CIG,'IE'Philosophy of biologyE']V6/"(`']V6/"('b$.A&"+.%4'W"&++,';<<ME JICIG,' 9E' Philosophy of experimental biologyE' D#0?".(7&`' D#0?".(7&' b$.A&"+.%4'W"&++,'KLLXE' JIYa[ICIG\FG ,' 9E' Developmental plasticity and evolutionE' ]V6/"(`' ]V6/"('b$.A&"+.%4'W"&++,'KLLME G&*.?.(/`'MLmLcmKL;K Received`'LcmMLmKL;K F8"/?#(/`';Km;KmKL;K < !&8(1`';Km;KmKL;K Rev. Filos., Aurora, Curitiba, v. 25, n. 36, p. 197-216, jan./jun. 2013