On the Origin of Species
On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life is a work of scientific literature by Charles Darwin that is considered to be the foundation of evolutionary biology. It was published on 24 November 1859. Darwin's book introduced the scientific theory that populations evolve over the course of generations through a process of natural selection, although Lamarckism was also included as a mechanism of lesser importance. The book presented a body of evidence that the diversity of life arose by common descent through a branching pattern of evolution. Darwin included evidence that he had collected on the Beagle expedition in the 1830s and his subsequent findings from research, correspondence, and experimentation.
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 other 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. Darwin was already highly regarded as a scientist, so 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, and it has now become 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:Background
Developments before Darwin's theory
In later editions of the book, Darwin traced evolutionary ideas as far back as Aristotle; the text he cites is a summary by Aristotle of the ideas of the earlier Greek philosopher Empedocles. Early Christian Church Fathers and Medieval European scholars interpreted the Genesis creation narrative allegorically rather than as a literal historical account; organisms were described by their mythological and heraldic significance as well as by their physical form. Nature was widely believed to be unstable and capricious, with monstrous births from union between species, and spontaneous generation of life.The Protestant Reformation inspired a literal interpretation of the Bible, with concepts of creation that conflicted with the findings of an emerging science seeking explanations congruent with the mechanical philosophy of René Descartes and the empiricism of the Baconian method. After the turmoil of the English Civil War, the Royal Society wanted to show that science did not threaten religious and political stability. John Ray developed an influential natural theology of rational order; in his taxonomy, species were static and fixed, their adaptation and complexity designed by God, and varieties showed minor differences caused by local conditions. In God's benevolent design, carnivores caused mercifully swift death, but the suffering caused by parasitism was a puzzling problem. The biological classification introduced by Carl Linnaeus in 1735 also viewed species as fixed according to the divine plan, but did recognize the hierarchical nature of different taxa. In 1766, Georges Buffon suggested that some similar species, such as horses and asses, or lions, tigers, and leopards, might be varieties descended from a common ancestor. The Ussher chronology of the 1650s had calculated creation at 4004 BC, but by the 1780s geologists assumed a much older world. Wernerians thought strata were deposits from shrinking seas, but James Hutton proposed a self-maintaining infinite cycle, anticipating uniformitarianism.
Charles Darwin's grandfather Erasmus Darwin outlined a hypothesis of transmutation of species in the 1790s, and French naturalist Jean-Baptiste Lamarck published a more developed theory in 1809. Both envisaged that spontaneous generation produced simple forms of life that progressively developed greater complexity, adapting to the environment by inheriting changes in adults caused by use or disuse. This process was later called Lamarckism. Lamarck thought there was an inherent progressive tendency driving organisms continuously towards greater complexity, in parallel but separate lineages with no perceptible extinction. Geoffroy contended that embryonic development recapitulated transformations of organisms in past eras when the environment acted on embryos, and that animal structures were determined by a constant plan as demonstrated by homologies. Georges Cuvier strongly disputed such ideas, holding that unrelated, fixed species showed similarities that reflected a design for functional needs. His palæontological work in the 1790s had established the reality of extinction, which he explained by local catastrophes, followed by repopulation of the affected areas by other species.
In Britain, William Paley's Natural Theology saw adaptation as evidence of beneficial "design" by the Creator acting through natural laws. All naturalists in the two English universities were Church of England clergymen, and science became a search for these laws. Geologists adapted catastrophism to show repeated worldwide annihilation and creation of new fixed species adapted to a changed environment, initially identifying the most recent catastrophe as the biblical flood. Some anatomists such as Robert Grant were influenced by Lamarck and Geoffroy, but most naturalists regarded their ideas of transmutation as a threat to divinely appointed social order.
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 helping Robert Grant's research into marine invertebrates. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it. Starting in 1827, at Cambridge University, 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 December 1831, he joined the Beagle expedition as a gentleman naturalist and geologist. 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, 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 argument that human populations, if unrestrained, 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 basic framework of his theory of natural selection, but he was fully occupied with his career as a geologist and held back from compiling it until his book on The Structure and Distribution of Coral Reefs was completed. As he recalled in his autobiography, he had "at last got a theory by which to work", but it was only in June 1842 that he allowed himself "the satisfaction of writing a very brief abstract of my theory in pencil".