Monday, July 25, 2011

On the Origin of Species

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Various evolutionary ideas had already been proposed to explain new findings in biology. There was growing support for such ideas among dissident anatomists and the general public, but during the first half of the 19th century the English scientific establishment was closely tied to the Church of England, while science was part of natural theology. Ideas about the transmutation of species were controversial as they conflicted with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique, unrelated to animals. The political and theological implications were intensely debated, but transmutation was not accepted by the scientific mainstream.

The book was written for non-specialist readers and attracted widespread interest upon its publication. As Darwin was an eminent scientist, his findings were taken seriously and the evidence he presented generated scientific, philosophical, and religious discussion. The debate over the book contributed to the campaign by T.H. Huxley and his fellow members of the X Club to secularise science by promoting scientific naturalism. Within two decades there was widespread scientific agreement that evolution, with a branching pattern of common descent, had occurred, but scientists were slow to give natural selection the significance that Darwin thought appropriate. During the "eclipse of Darwinism" from the 1880s to the 1930s, various other mechanisms of evolution were given more credit. With the development of the modern evolutionary synthesis in the 1930s and 1940s, Darwin's concept of evolutionary adaptation through natural selection became central to modern evolutionary theory, now the unifying concept of the life sciences.

Summary of Darwin's theory

Darwin's theory of evolution is based on key facts and the inferences drawn from them, which biologist Ernst Mayr summarised as follows:

- Every species is fertile enough that if all offspring survived to reproduce the population would grow (fact).

- Despite periodic fluctuations, populations remain roughly the same size (fact).

- Resources such as food are limited and are relatively stable over time (fact).

- A struggle for survival ensues (inference).

- Individuals in a population vary significantly from one another (fact).

- Much of this variation is inheritable (fact).

- Individuals less suited to the environment are less likely to survive and less likely to reproduce; individuals more suited to the environment are more likely to survive and more likely to reproduce and leave their inheritable traits to future generations, which produces the process of natural selection (inference).

- This slowly effected process results in populations changing to adapt to their environments, and ultimately, these variations accumulate over time to form new species (inference).




Inception of Darwin's theory

Darwin went to Edinburgh University in 1825 to study medicine. In his second year he neglected his medical studies for natural history and spent four months assisting Robert Grant's research into marine invertebrates. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it. At Cambridge University starting in 1827, Darwin learnt science as natural theology from botanist John Stevens Henslow, and read Paley, John Herschel and Alexander von Humboldt. Filled with zeal for science, he studied catastrophist geology with Adam Sedgwick. In mid-July 1837 Darwin started his "B" notebook on Transmutation of Species, and on page 36 wrote "I think" above his first evolutionary tree.

In December 1831, he joined the Beagle expedition as a geologist and naturalist. He read Charles Lyell's Principles of Geology and from the first stop ashore, at St. Jago, found Lyell's uniformitarianism a key to the geological history of landscapes. Darwin discovered fossils resembling huge armadillos, and noted the geographical distribution of modern species in hope of finding their "centre of creation". The three Fuegian missionaries the expedition returned to Tierra del Fuego were friendly and civilised, yet to Darwin their relatives on the island seemed "miserable, degraded savages", and he no longer saw an unbridgeable gap between humans and animals. As the Beagle neared England in 1836, he noted that species might not be fixed.

Richard Owen showed that fossils of extinct species Darwin found in South America were allied to living species on the same continent. In March 1837, ornithologist John Gould announced that Darwin's Rhea was a separate species from the previously described rhea (though their territories overlapped), that mockingbirds collected on the Galápagos Islands represented three separate species each unique to a particular island, and that several distinct birds from those islands were all classified as finches. Darwin began speculating, in a series of notebooks, on the possibility that "one species does change into another" to explain these findings, and around July sketched a genealogical branching of a single evolutionary tree, discarding Lamarck's independent lineages progressing to higher forms. Unconventionally, Darwin asked questions of fancy pigeon and animal breeders as well as established scientists. At the zoo he had his first sight of an ape, and was profoundly impressed by how human the orangutan seemed.

In late September 1838, he started reading Thomas Malthus's An Essay on the Principle of Population with its statistical proof that human populations breed beyond their means and struggle to survive. Darwin related this to the struggle for existence among wildlife and botanist de Candolle's "warring of the species" in plants; he immediately envisioned "a force like a hundred thousand wedges" pushing well-adapted variations into "gaps in the economy of nature", so that the survivors would pass on their form and abilities, and unfavourable variations would be destroyed. By December 1838, he had noted a similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected".

Darwin now had the framework of his theory of natural selection "by which to work", but he was fully occupied with his career as a geologist and held off writing a sketch of his theory until his book on The Structure and Distribution of Coral Reefs was completed in May 1842.

Time taken to publish

Darwin had his basic theory of natural selection "by which to work" by December 1838, yet almost twenty years later, when Wallace's letter arrived on 18 June 1858, Darwin was still not ready to publish his theory. It was long thought that Darwin avoided or delayed making his ideas public for personal reasons. Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed, and concern about upsetting his clergymen naturalist friends or his pious wife Emma. Charles Darwin's illness caused repeated delays. His paper on Glen Roy had proved embarrassingly wrong, and he may have wanted to be sure he was correct. David Quammen has suggested all these factors may have contributed, and notes Darwin's large output of books and busy family life during that time.

A more recent study by science historian John van Wyhe has determined that the idea that Darwin delayed publication only dates back to the 1940s, and Darwin's contemporaries thought the time he took was reasonable. Darwin always finished one book before starting another. While he was researching, he told many people about his interest in transmutation without causing outrage. He firmly intended to publish, but it was not until September 1854 that he could work on it full time. His estimate that writing his "big book" would take five years was optimistic.

Publication and subsequent editions

On the Origin of Species was first published on Thursday 24 November 1859, priced at fifteen shillings. The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November, and all available copies had been taken up immediately. In total, 1,250 copies were printed but after deducting presentation and review copies, and five for Stationers' Hall copyright, around 1,170 copies were available for sale. Significantly, 500 were taken by Mudie's Library, ensuring that the book promptly reached a large number of subscribers to the library. The second edition of 3,000 copies was quickly brought out on 7 January 1860, and incorporated numerous corrections as well as a response to religious objections by the addition of a new epigraph on page ii, a quotation from Charles Kingsley, and the phrase "by the Creator" amended to the closing sentence. During Darwin's lifetime the book went through six editions, with cumulative changes and revisions to deal with counter-arguments raised. The third edition came out in 1861, with a number of sentences rewritten or added and an introductory appendix, An Historical Sketch of the Recent Progress of Opinion on the Origin of Species, while the fourth in 1866 had further revisions. The fifth edition, published on 10 February 1869, incorporated more changes and for the first time included the phrase "survival of the fittest", which had been coined by the philosopher Herbert Spencer in his Principles of Biology (1864).

In January 1871, George Jackson Mivart's On the Genesis of Species listed detailed arguments against natural selection, and claimed it included false metaphysics. Darwin made extensive revisions to the sixth edition of the Origin (this was the first edition in which he used the word "evolution"), and added a new chapter VII, Miscellaneous objections, to address Mivart's arguments. The sixth edition was published by Murray on 19 February 1872 with "On" dropped from the title. Darwin had told Murray of working men in Lancashire clubbing together to buy the 5th edition at fifteen shillings and wanted it made more widely available; the price was halved to 7s 6d by printing in a smaller font. It includes a glossary compiled by W.S. Dallas. Book sales increased from 60 to 250 per month.

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