Lei de Dollo

A lei de Dollo ou lei de Dollo da irreversibilidade evolutiva ou principio de Dollo é unha hipótese proposta en 1893[1] polo paleontólogo belga nado en Francia Louis Dollo, que di que a evolución non é reversible. A formulación desta hipótese tal como a fixo inicialmente Dollo era: "Un organismo non pode retornar, nin sequera parcialmente, a un estado previo polo que xa pasou na súa liña de antepasados."[2] Esta enunciación é a miúdo malinterpretada como unha hipótese menos esixente referida á posibilidade de recuperar estruturas ou órganos perdidos, que é improbable que reaparezan na "mesma forma" (definida con pouca precisión).[3][4] Segundo Richard Dawkins, esta lei é "realmente só unha afirmación sobre a improbabilidade estatística de seguir exactamente a mesma traxectoria evolutiva dúas veces (ou calquera traxectoria determinada), en calquera dirección."[5] Stephen Gould suxeriu que a irreversibilidade hipoteca certas vías evolutivas unha vez que emerxen as formas principais: "[Por exemplo], unha vez que se adopta o plan corporal ordinario dun réptil, centos de opcións quedan para sempre pechadas, e as posibilidades futuras deben desenvolverse dentro dos límites do deseño herdado."[6]

Un exemplo

Nun estudo de 2009 sobre a evolución da estrutura das proteínas propúxose un novo mecanismo para a lei de Dollo. No estudo examinouse un receptor hormonal que evolucionara desde dunha proteína ancestral que podía ligarse a dúas hormonas pero que cambiou a unha nova proteína que era específica para unha soa hormona. Este cambio fora causado por dúas substitucións de aminoácidos, que impiden que se produza a unión da segunda hormona. Porén, ocorreron seguidamente outros cambios, que eran selectivamente neutros, xa que non afectaban á unión a hormonas. Cando os autores trataron de reverter a proteína ao seu estado ancestral facendo mutar os dous "residuos de unión", encontraron que os outros cambios producidos desestabilizaran o estado ancestral da proteína. Concluíron que para que evolucionase esta proteína revertendo ao estado anterior e recuperando a súa capacidade de unirse ás dúas hormonas, terían que producirse varias mutacións neutras independentes simplemente por azar e sen presión de selección. Como isto é extremadamente improbable, pode explicar por que a evolución tende a ir nunha soa dirección.[7]

Excepcións propostas á lei de Dollo

Aínda que non está claro cal é o limiar exacto para a violación da lei de Dollo, hai varios estudos de casos cuxos resultados discuten a validez dalgunhas interpretacións. Por exemplo, moitos taxons de gasterópodos reduciron as súas cunchas, e algúns perderon o enrolamento en espiral.[8] Segundo a interpretación de S. J. Gould da lei de Dollo, non sería posible recuperar a cuncha enrolada en espiral despois de que se perdera o enrolamento. Non obstante, uns poucos xéneros da familia de caracois Calyptraeidae puideron ter cambiado o seu ritmo de desenvolvemento (heterocronía) e recuperado unha cuncha enrolada a partir dunha cuncha non espiralada de tipo patela.[8][9] Outras "excepcións" propostas inclúen as ás dos insectos pau,[10] os estadios larvarios de píntegas,[11][12] perda de dedas en lagartos,[13][14] de clavículas en dinosauros terópodos non aviarios,[15] do pescozo, a rexión pectoral, e de musculatura das extremidades superiores en primates, incluíndo na liñaxe que deu lugar aos humanos.[16]

Notas

  1. Dollo, L (1893). Les lois de l'évolution. Arquivado 03 de maio de 2015 en Wayback Machine. Bull. Soc. Belge Geol. Pal. Hydr, VII:164-166.
  2. Dollo, quoted in "Evolution: Ammonites Indicate Reversal," in Nature, March 21, 1970
  3. Goldberg, Emma E; Boris Igić (2008). "On phylogenetic tests of irreversible evolution". Evolution 62 (11): 2727–2741. PMID 18764918. doi:10.1111/j.1558-5646.2008.00505.x. 
  4. Collin, Rachel; Maria Pia Miglietta (2008). "Reversing opinions on Dollo's Law". Trends in Ecology & Evolution 23 (11): 602–609. PMID 18814933. doi:10.1016/j.tree.2008.06.013. 
  5. Dawkins, Richard (1996) [1986]. The Blind Watchmaker. New York: W. W. Norton & Company, Inc. ISBN 0-393-31570-3. 
  6. Gould, Stephen J. [1993] (2007) "Eight little piggies," Vintage Books. ISBN 978-0-09-950744-4
  7. Bridgham, Jamie T.; Eric A. Ortlund; Joseph W. Thornton (2009). "An epistatic ratchet constrains the direction of glucocorticoid receptor evolution". Nature 461 (7263): 515–519. ISSN 0028-0836. PMID 19779450. doi:10.1038/nature08249. 
  8. 8,0 8,1 Collin, R.; Cipriani, R. (2003). "Dollo's law and the re-evolution of shell coiling". Proceedings of the Royal Society B 270 (1533): 2551–2555. PMC 1691546. PMID 14728776. doi:10.1098/rspb.2003.2517. 
  9. Pagel, M. (2004). "Limpets break Dollo's Law". Trends in Ecology & Evolution 19: 278. doi:10.1016/j.tree.2004.03.020. 
  10. Whiting, MF.,S. Bradler & T. Maxwell (2003). Loss and recovery of wings in stick insects. Nature 421 264-267.
  11. Chippindale, P. T. and J. J. Wiens. (2005). Re-evolution of the larval stage in the Plethodontid salamander genus Desmognathus. Herpetological Review 36(2) 113.
  12. Marshall, C. R., et al. (1994). Dollo's law and the death and resurrection of genes. Arquivado 25 de xullo de 2014 en Wayback Machine. Proc Natl Acad Sci USA 91 12283.
  13. Galis, F., et al. (2010). Dollo's law and the irreversibility of digit loss in Bachia. Evolution 64(8) 2466.
  14. Davies, E. Frogs re-evolved lost lower teeth. BBC News. January 31, 2011. Retrieved February 9, 2011.
  15. Paul, Gregory S (2002). Dinosaurs of the Air: the evolution and loss of flight in dinosaurs and birds. CJHU Press. p. 10. ISBN 0-8018-6763-0. 
  16. Diogo, R.; Wood, B. (2012). "Violation of Dollo's Law: Evidence of Muscle Reversions in Primate Phylogeny and Their Implications for the Understanding of the Ontogeny, Evolution, and Anatomical Variations of Modern Humans". Evolution 66 (10): 3267. doi:10.1111/j.1558-5646.2012.01621.x. 

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