In the Light of Evolution III: Two Centuries of Darwin
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FROM THE ACADEMY: COLLOQUIUM PERSPECTIVE In the light of evolution III: Two centuries of Darwin John C. Avise1 and Francisco J. Ayala1 Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697 harles Darwin’s enthusiasm autobiography (13) that Paley’s logic able to itself, under the complex and and expertise in natural history ‘‘gave me as much delight as did Euclid’’ sometimes varying conditions of life, contributed hugely to his eluci- and that it was the ‘‘part of the Aca- will have a better chance of surviving, dation of evolution by natural demical Course [at the University of and thus be naturally selected. From the Cselection, which stands as one of the Cambridge] which...wasthemost use strong principle of inheritance, any se- grandest intellectual achievements in the to me in the education of my mind.’’ lected variety will tend to propagate its history of science. Darwin was a lifelong Darwin was still a natural theologian new and modified form.’’ Darwin’s clear observer of nature, stating in correspon- when he boarded the Beagle in 1831 on elucidation of natural selection launched dence that some of his happiest times in what would become a fateful voyage, for a revolutionary new paradigm in biology youth were spent fishing on rainy days Darwin and for humanity, into un- wherein organismal traits could be stud- and ‘‘entomologizing’’ when England’s charted philosophical (as well as scien- ied and interpreted as products of natu- weather was nice. At the age of 22, he tific) waters. ral (rather than supernatural) forces boarded the HSM Beagle for a 5-year In the articles of this Colloquium, amenable to rational scientific inquiry. stint as Captain Fitzroy’s traveling com- leading evolutionary biologists and sci- Scientific studies of natural selection are panion and the ship’s naturalist, an ence historians reflect on and commem- now more popular and powerful than appointment that introduced him to bio- orate the Darwinian Revolution. The ever, and they have revealed the evolu- diversity on a global geographic scale. authors of these Proceedings canvass tionary origins and trajectories of nu- Darwin’s breadth and depth of natural- modern research approaches and cur- merous biological features and taxa. history experience would later be on full rent scientific thought on each of the A major limitation in Darwin’s char- display in his most defining scientific 3 main categories of selection (natural, acterization of evolution concerned he- works (3–5) in his detailed treatises on artificial, and sexual) that Darwin ad- reditary mechanisms, a difficulty that orchids, insectivorous plants, coral reefs, dressed during his career. Although his the field began to rectify early in the barnacles, and earthworms (6–10). legacy is associated primarily with the 20th century by incorporating Mende- The year 2009 marks the 200th anni- illumination of natural selection in The lian genetics and population genetics versary of Charles Darwin’s birth and Origin, Darwin also contemplated and into the emerging evolutionary synthesis the 150th anniversary of his most influ- wrote extensively about what we would (14). Today, in the genomics era, scien- ential publication (3). Darwin trans- now term artificial selection and sexual tists routinely extend studies of natural formed the biological sciences in much selection, as reflected for example in selection and trait evolution to the level the same way that Nicolaus Copernicus, two books titled, respectively, The Varia- of DNA itself, as several papers in these Galileo Galilei, and Isaac Newton, cen- tion of Animals and Plants Under Domes- Proceedings will attest. Genomic dissec- turies earlier, transformed the physical tication (1869) and The Descent of Man tions are also providing fresh insights sciences—by demonstrating that the uni- and Selection in Relation to Sex (1871). into the ancient mystery alluded to in verse operates according to natural laws In a concluding section of these Pro- the title of Darwin’s seminal work: how that fall within the purview of rational species originate. Ironically, The Origin scientific inquiry. In 1543, Copernicus ceedings, several science historians com- published De revolutionibus orbium ment on Darwin’s seminal contributions. says relatively little about the evolution celestium (‘‘On the Revolutions of the Thus, these Proceedings are organized of reproductive isolating barriers, which Celestial Spheres’’) that challenged con- in 4 parts: Natural Selection, or Adapta- under the modern biological species ventional wisdom that the Earth was the tion to Nature; Artificial Selection, or concept are key to understanding clado- center of Creation, and instead pro- Adaptation to Human Demands; Sexual genetic (speciational) processes. moted the idea that natural laws govern Selection, or Adaptation to Mating De- In the opening presentation of these the motion of physical objects in the mands; and The Darwinian Legacy, 150 Proceedings, Via (15) takes a fresh per- universe. In 1859, in On the Origin of Years Later. spective on the origin of species by char- Species, Darwin developed the equally acterizing genomic regions that appear Natural Selection, or Adaptation to revolutionary concept that a natural but to be diverging early in a speciation pro- Nature nonrandom process—natural selection— yields biological adaptations that other- The concept of natural selection—as the unconscious broker of adaptive evolu- This paper serves as an introduction to this PNAS supple- wise can give the superficial impression ment, which resulted from the Arthur M. Sackler Collo- of direct intelligent craftsmanship. tion—is Darwin’s seminal contribution. quium of the National Academy of Sciences, ‘‘In the Light of Darwin’s impacts have been felt far It provided a materialistic account of Evolution III: Two Centuries of Darwin,’’ held January 16– beyond science. Before Darwin, most nature’s operations that contrasted 17, 2009, at the Arnold and Mabel Beckman Center of the sharply with the traditional invocations National Academies of Sciences and Engineering in Irvine, scientists and theologians accepted what CA. It is the third in a series of colloquia under the general seemed obvious: that divine intervention of supernatural causation that predomi- title ‘‘In the Light of Evolution’’ (ILE; and see Box 1). The must have underlain nature’s design. nated before The Origin. The basic logic complete program and audio files of most presentations of natural selection is astonishingly sim- are available on the NAS web site at www.nasonline.org/ The traditional ‘‘argument from design’’ ple. As phrased by Darwin in The Ori- Sackler Darwin. Papers from the first two colloquia in the traces back at least to the classical ILE series, titled ‘‘Adaptation and Complex Design’’ and Greek philosopher Socrates in the 5th gin, ‘‘As many more individuals of each ‘‘Biodiversity and Extinction,’’ appeared in refs. 1 and 2, century B.C. (see ref. 11), and it was species are born than can possibly sur- respectively. expressed again in a thoughtful and ele- vive; and as, consequently, there is a Author contributions: J.C.A. and F.J.A. wrote the paper. gant treatise (Natural Theology) pub- frequently recurring struggle for exis- The authors declare no conflict of interest. lished in 1802 by the Reverend William tence, it follows that any being, if it vary 1To whom correspondence may be addressed. E-mail: Paley (12). Darwin later recalls in his however slightly in any manner profit- [email protected] or [email protected]. www.pnas.org͞cgi͞doi͞10.1073͞pnas.0903381106 PNAS ͉ June 16, 2009 ͉ vol. 106 ͉ suppl. 1 ͉ 9933–9938 Downloaded by guest on October 5, 2021 cess. She calls this the ‘‘magnifying and prezygotic isolation under gene and species in nature. In chapter 1 of glass’’ approach for speciation in action, flow, and the role of standing genetic The Origin, Darwin discussed several and contrasts it with the more tradi- variation in the process) and specifically domesticated plant and animal species, tional ‘‘spyglass’’ approach in which each (with reference to speciation in stickle- ranging from beans, melons, and plums completed speciation is characterized back fishes). For the sticklebacks, they to dogs, cattle, and horses. He devoted a retrospectively by scrutinizing genetic develop an interesting ‘‘transporter long section to how selective breeding differences between established sister model’’ of ecological speciation in which had altered the domestic pigeon, fancy taxa. Via develops and presents genetic ecological selection pressures in fresh- varieties of which were widely prized in evidence for a model in which incipient water streams consistently select for al- the Victorian era. Ten years later, he species become, in effect, genealogical leles different from those normally would expand greatly on these themes in mosaics in which ecologically important present in marine populations. However, The Variation of Animals and Plants Un- genomic regions (i.e., those under diver- occasional hybridization between fresh- der Domestication. gent ecological selection, sometimes water and marine forms ensures a con- In chapter 1 of The Origin, Darwin even in sympatry) become resistant to tinual supply of freshwater alleles in the lamented that ‘‘We hardly know any- genetic exchange, whereas gene flow sea, at low frequency and disassembled thing about the origin or history of our remains possible over most of the ge- by genetic recombination. When marine domestic breeds’’; and ‘‘The origin of nome. The key genomic regions under fish colonize a newly opened stream, most of our domestic animals will prob- divergent selection become focal points natural selection can act on this stand- ably for ever remain vague.’’ Darwin for ‘‘divergence hitchhiking’’ by linked ing pool of genetic variation to reconsti- would therefore be both pleased and loci, because they reduce the porosity of tute the freshwater genotype. The anal- surprised by recent scientific progress in the emerging species boundary to gene ogy in the title of their model is to a deciphering the evolutionary origins of exchange.