On the Origin of Species

• 1. Introduction

  ›

  On the Origin of Species arrived in bookshops on the 24th of November 1859, priced at fifteen shillings, and every available copy was gone before the end of the day. The book had been offered to booksellers at Murray's autumn sale on the 22nd of November, and all copies were taken up immediately. The first printing ran to 1,250 copies, but after presentation and review copies were set aside, roughly 1,170 were actually available for sale. Five hundred of those went straight to Mudie's Library, putting the book into the hands of a vast network of subscribers almost overnight.

  The man behind it was Charles Darwin, an English naturalist already held in high regard by the scientific community. He had spent more than two decades quietly gathering evidence for a theory that would unsettle not just biology but philosophy, theology, and the way human beings understood their own place in nature. The question this documentary will follow is: how did a cautious, methodical naturalist who had once studied barnacles for eight consecutive years arrive at a manuscript that would reshape the life sciences? And why did it take him so long to publish it?

• 2. An Idea Decades in the Making

  ›

  Darwin went to Edinburgh University in 1825 to study medicine. In his second year he began neglecting medicine in favour of natural history, spending four months assisting Robert Grant's research into marine invertebrates. Grant shared his enthusiasm for the transmutation of species, but Darwin rejected the idea. The intellectual environment Darwin moved into at Cambridge, beginning in 1827, reinforced that skepticism. Botanist John Stevens Henslow taught him science as natural theology, and he read William Paley alongside John Herschel and Alexander von Humboldt.

  In December 1831 Darwin joined the Beagle expedition as a gentleman naturalist and geologist. He read Charles Lyell's Principles of Geology on the voyage, and from the very first stop ashore, at St. Jago, found Lyell's uniformitarianism a compelling key to reading geological history. He discovered fossils resembling giant armadillos and began tracking the geographical distribution of modern species. The three Fuegian missionaries the expedition returned to Tierra del Fuego were friendly and, in Darwin's view, civilised; yet he found their relatives on the island to be, in his words, "miserable, degraded savages". That encounter eroded his sense of an unbridgeable gulf between humans and other animals.

  As the Beagle neared England in 1836, Darwin noted that species might not be fixed at all. In March 1837, ornithologist John Gould announced that Darwin's rhea was a separate species from the previously described rhea, that mockingbirds from the Galapagos Islands represented three distinct species each unique to a single island, and that several clearly different birds from those islands all belonged to the finch family. Darwin began filling notebooks with speculation about whether one species could change into another, and around July of that year he sketched a genealogical branching diagram - a single tree of life, discarding Lamarck's notion of independent lineages progressing independently upward.

  In late September 1838, reading Thomas Malthus's An Essay on the Principle of Population, Darwin connected the statistical argument about human populations outstripping their resources to the struggle for existence he had observed among wildlife. He immediately envisioned what he described as "a force like a hundred thousand wedges" driving well-adapted variations into gaps in the natural order. By December 1838 he had noted the parallel between breeders selecting traits and a Malthusian nature selecting among variants thrown up by chance. He had the core framework. It was only in June 1842, however, that he allowed himself what he later called "the satisfaction of writing a very brief abstract of my theory in pencil".

• 3. The Long Delay

  ›

  Darwin expanded his pencil sketch to a 35-page "Pencil Sketch" in June 1842, then wrote to botanist Joseph Dalton Hooker in January 1844, and by July had rounded the sketch into a 230-page "Essay". He left instructions that the Essay should be published if he died prematurely. Yet he did not rush it into print.

  Biographers have proposed several explanations for the gap between conception and publication. Some have pointed to fear of religious persecution, concern about upsetting his clergymen friends or his pious wife Emma, and the embarrassment of an earlier paper on Glen Roy that had proved wrong. Darwin himself wrote in his autobiography that he had "gained much by my delay in publishing from about 1839, when the theory was clearly conceived, to 1859; and I lost nothing by it." A more recent study by science historian John van Wyhe found that the belief Darwin delayed for personal reasons only dates back to the 1940s, and that his contemporaries considered the time he took entirely reasonable. Darwin always finished one book before starting another; his barnacle studies alone took eight years and made him the world's leading expert on their classification.

  In November 1844, the anonymously published Vestiges of the Natural History of Creation, written by Scottish journalist Robert Chambers, widened public interest in transmutation of species. Darwin read it soon after publication and scorned its amateurish geology and zoology, but he carefully reviewed his own arguments when leading scientists attacked Vestiges on scientific and moral grounds. The public debate the book sparked did, however, move evolutionary speculation closer to the mainstream, preparing an audience for what Darwin was building.

  Hooker was persuaded to take away a copy of Darwin's Essay in January 1847 and eventually sent back a page of notes offering Darwin much-needed feedback. Reminded of his lack of expertise in taxonomy, Darwin began the barnacle study that would occupy him until 1854. It was not until September 1854 that he could work on evolution full-time. His 1846 estimate that writing his "big book" would take five years proved optimistic. By 1856, his theory was far more sophisticated than it had been in 1844, with a mass of supporting evidence. Geologist Charles Lyell, recognising the implications of an 1855 paper by Alfred Russel Wallace on species distribution, urged Darwin in a letter of the 1st and the 2nd of May 1856 to publish and establish priority. Darwin began a "sketch" on the 14th of May 1856, and by July had decided to write a full technical treatise.

• 4. Wallace and the Race to Print

  ›

  On the 18th of June 1858, Darwin received a parcel from Wallace, who was staying on the Maluku Islands - Ternate and Gilolo. Inside were twenty pages describing an evolutionary mechanism, sent as a response to Darwin's recent encouragement, with a request to forward the pages to Lyell if Darwin thought them worthwhile. The mechanism was closely similar to Darwin's own theory of natural selection.

  Darwin wrote to Lyell that "your words have come true with a vengeance,... forestalled" and that he would "of course, at once write and offer to send it to any journal" that Wallace chose. He added that "all my originality, whatever it may amount to, will be smashed". Lyell and Hooker proposed a joint publication: Wallace's pages would appear alongside extracts from Darwin's 1844 Essay and his 1857 letter to Asa Gray. On the 1st of July 1858, papers by Wallace and Darwin under the title "On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection" were read at the Linnean Society. They drew little reaction at the time.

  Historians have noted meaningful differences between the two men's arguments. Darwin described natural selection as analogous to the artificial selection practised by animal breeders and emphasised competition between individuals. Wallace drew no comparison to selective breeding and focused instead on ecological pressures that kept varieties adapted to local conditions. Some historians have suggested Wallace was describing group selection rather than selection acting on individual variation. Soon after the Linnean meeting, Darwin decided to write an abstract of his whole work, beginning it on the 20th of July 1858 while on holiday at Sandown, composing parts from memory and sending manuscripts to friends for checking.

• 5. From Manuscript to Bookshop

  ›

  By mid-March 1859 Darwin's abstract had grown far beyond anything that could fit in a journal article. Lyell suggested the publisher John Murray, and met with him to gauge his interest. On the 28th of March Darwin wrote to Lyell offering to assure Murray "that my Book is not more un-orthodox, than the subject makes inevitable". Murray agreed to publish without even reading the manuscript, offering Darwin two-thirds of the profits on the same terms he extended to Lyell himself.

  Murray's reader Whitwell Elwin, after reviewing the first three chapters, advised that Darwin should write a book solely about pigeons, suggesting that "Every body is interested in pigeons". Darwin replied that this was impractical: he had only the last chapter still to write. The title went through several drafts. An early version read On the Mutability of Species. Another proposed An abstract of an Essay on the Origin of Species and Varieties Through natural selection. Darwin wanted to keep the phrase "natural selection" because it was "constantly used in all works on Breeding". Murray eventually persuaded Darwin to drop "abstract" and "varieties", and the book appeared as On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. Murray paid Darwin £180 for the first edition.

  The second edition of 3,000 copies appeared on the 7th of January 1860, incorporating corrections and, in response to religious objections, a new epigraph from Charles Kingsley and the phrase "by the Creator" added to the closing sentence. The book went through six editions in Darwin's lifetime. The fifth edition, published on the 10th of February 1869, included for the first time the phrase "survival of the fittest", coined by the philosopher Herbert Spencer in his Principles of Biology in 1864. The sixth edition, published on the 19th of February 1872 by Murray as The Origin of Species, halved the price to 7 shillings 6 pence after Darwin told Murray of working men in Lancashire clubbing together to buy the fifth edition at fifteen shillings. Book sales rose from 60 to 250 copies per month.

• 6. What the Book Actually Argued

  ›

  Biologist Ernst Mayr later summarised Darwin's core argument as a chain of facts and inferences. Every species produces more offspring than can survive. Populations stay roughly stable in size despite that fertility. Resources are limited. A struggle for survival follows. Individuals within a population vary from one another, and much of that variation is heritable. Those less suited to their environment are less likely to survive and reproduce; those better suited pass their traits on. Over generations, populations shift toward better adaptation, and given enough time those shifts accumulate into new species.

  Darwin built the argument in stages across twelve chapters. The opening chapters examined animal husbandry and plant breeding going back to ancient Egypt, using fancy pigeon breeding as a concrete illustration of what selection could accomplish. He noted that "the diversity of the breeds is something astonishing", yet all were descended from one species, the rock dove. He distinguished two kinds of variation: rare abrupt changes he called "sports" or "monstrosities", such as Ancon sheep with short legs, and the ubiquitous small differences, such as slightly shorter or longer pigeon bills. For Darwin, the small changes were the ones that mattered most in evolution.

  Chapter VI confronted the hardest objections directly. Darwin quoted himself asking: if species descended from other species by gradual change, "why do we not everywhere see innumerable transitional forms? Why is not all nature in confusion?" His answer was that intermediate forms, being less numerous than the forms they connected, were most vulnerable to extinction. He also addressed the eye, asking whether natural selection could have produced an organ of such apparent perfection. His answer was to point to invertebrates with simple eyes consisting of nothing more than an optic nerve coated with pigment, arguing these represented functional intermediate stages. He concluded: "If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case."

  Chapter XIII discussed morphology and pressed a striking observation: the hand of a man formed for grasping, the mole's paw for digging, the horse's leg, the porpoise's paddle, and the bat's wing are all constructed on the same underlying pattern with the same bones in the same relative positions. This made no sense if each species had been created independently, but was exactly what common descent predicted. The book closed with a passage that became one of the most quoted in science: "There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved."

• 7. Reception and a Slow Revolution

  ›

  The Examiner reviewed the book on the 3rd of December 1859, noting that much of Darwin's volume was "tough reading" requiring concentrated attention, but that many parts abounded in information that was easy to comprehend and both instructive and entertaining. Richard Owen complained in the Quarterly Review that the style was too easy for a serious work of science. David Quammen later described the book as written in everyday language for a wide audience, but noted Darwin's literary style was uneven: some passages featured convoluted sentences, while others were almost lyrical.

  Darwin was already a respected scientist, so his findings were taken seriously. The debate the book generated contributed to the campaign by T. H. Huxley and his fellow members of the X Club to drive scientific naturalism into the mainstream and separate science from clerical authority. Within two decades there was widespread scientific agreement that evolution had occurred and that life shared a pattern of common descent. But scientists were slow to grant natural selection the central importance Darwin believed it deserved.

  From the 1880s through the 1930s, a period historians called "the eclipse of Darwinism", other proposed mechanisms of evolution received more credit. It was only with the development of the modern evolutionary synthesis in the 1930s and 1940s - integrating Darwin's mechanism with Gregor Mendel's theories of genetic inheritance, a combination dubbed Neo-Darwinian Evolution - that natural selection became the unifying concept of the life sciences. George Jackson Mivart's On the Genesis of Species, published in January 1871, had listed detailed arguments against natural selection and accused it of involving false metaphysics. Darwin responded in the sixth edition of Origin by adding a new chapter, Miscellaneous Objections, and proposing scenarios for the incremental evolution of features Mivart had cited as inexplicable, including baleen filters in whales and the camouflage of stick insects.

• 8. Beyond Britain

  ›

  Botanist Asa Gray, based in America, negotiated with a Boston publisher for an authorised American edition, but learned that two New York firms were already planning to exploit the absence of international copyright to print Origin without permission. Gray managed to secure a 5% royalty with Appleton's of New York, and in a May letter Darwin mentioned a print run of 2,500 copies for the American market, though it was unclear whether this figure covered only the first printing.

  Translation proved thornier. Darwin distributed presentation copies in France and Germany hoping suitable translators would come forward, since translators were expected to make their own arrangements with local publishers. The German translation published in 1860, by the distinguished elderly naturalist Heinrich Georg Bronn, imposed Bronn's own ideas, adding controversial themes Darwin had deliberately left out. Bronn translated "favoured races" as "perfected races" and added essays on the origin of life and on religious implications, partly shaped by his adherence to Naturphilosophie. Bronn produced a second German edition in 1862 but then died of a heart attack. Darwin subsequently worked closely with Julius Victor Carus, who published an improved German translation in 1867.

  In France, Darwin's attempts to find a translator fell through. The translation by Clemence Royer published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating social Darwinism and eugenics, along with numerous explanatory notes answering doubts that Darwin himself had merely flagged. Darwin had difficulty persuading Royer to remove her notes and remained dissatisfied until a translation by Edmond Barbier appeared in 1876. A Dutch translation by Tiberius Cornelis Winkler was published in 1860. In Darwin's lifetime the book also appeared in Italian and Russian by 1864, Swedish in 1871, Danish in 1872, Polish in 1873, Hungarian in 1873-1874, Spanish in 1877, and Serbian in 1878. As of 1977, Origin had appeared in an additional eighteen languages beyond those published in Darwin's lifetime, including a Chinese translation by Ma Chun-wu who published the preliminaries and chapters one through five in 1902-1904, completing his translation in 1920.