(1809-82), British scientist, who laid the foundation of modern evolutionary theory with his concept of the development of all forms of life through the slow-working process of natural selection. His work was a major influence on the life and earth sciences and on modern thought in general.

Born in Shrewsbury, Shropshire, England, on Feb. 12, 1809, Darwin was the fifth child of a wealthy and sophisticated English family. His maternal grandfather was the successful china and pottery entrepreneur Josiah Wedgwood; his paternal grandfather was the well-known 18th-century physician and savant Erasmus Darwin. After graduating from the elite school at Shrewsbury in 1825, young Darwin went to the University of Edinburgh to study medicine. In 1827 he dropped out of medical school and entered the University of Cambridge, in preparation for becoming a clergyman of the Church of England. There he met two stellar figures: Adam Sedgwick, a geologist, and John Stevens Henslow (1795-1861), a naturalist. Henslow not only helped build Darwin's self-confidence but also taught his student to be a meticulous and painstaking observer of natural phenomena and collector of specimens. After graduating from Cambridge in 1831, the 22year-old Darwin was taken aboard the English survey ship HMS Beagle, largely on Henslow's recommendation, as an unpaid naturalist on a scientific expedition around the world.

Voyage of the Beagle.

Darwin's job as naturalist aboard the Beagle gave him the opportunity to observe the various geological formations found on different continents and islands along the way, as well as a huge variety of fossils and living organisms. In his geological observations, Darwin was most impressed with the effect that natural forces had on shaping the earth's surface.

At the time, most geologists adhered to the so-called catastrophist theory that the earth had experienced a succession of creations of animal and plant life, and that each creation had been destroyed by a sudden catastrophe, such as an upheaval or convulsion of the earth's surface ( see Geology: History of Geological Thought: 18th and 19th Centuries ). According to this theory, the most recent catastrophe, Noah's flood, wiped away all life except those forms taken into the ark. The rest were visible only in the form of fossils. In the view of the catastrophists, species were individually created and immutable, that is, unchangeable for all time.

The catastrophist viewpoint (but not the immutability of species) was challenged by the English geologist Sir Charles Lyell in his two-volume work Principles of Geology (1830-33). Lyell maintained that the earth's surface is undergoing constant change, the result of natural forces operating uniformly over long periods.

Aboard the Beagle, Darwin found himself fitting many of his observations into Lyell's general uniformitarian view. Beyond that, however, he realized that some of his own observations of fossils and living plants and animals cast doubt on the Lyell-supported view that species were specially created. He noted, for example, that certain fossils of supposedly extinct species closely resembled living species in the same geographical area. In the Galapagos Islands, off the coast of Ecuador, he also observed that each island supported its own form of tortoise, mockingbird, and finch; the various forms were closely related but differed in structure and eating habits from island to island. Both observations raised the question, for Darwin, of possible links between distinct but similar species.

Theory of Natural Selection.

After returning to England in 1836, Darwin began recording his ideas about changeability of species in his Notebooks on the Transmutation of Species. Darwin's explanation for how organisms evolved was brought into sharp focus after he read An Essay on the Principle of Population (1798), by the British economist Thomas Robert Malthus, who explained how human populations remain in balance. Malthus argued that any increase in the availability of food for basic human survival could not match the geometrical rate of population growth. The latter, therefore, had to be checked by natural limitations such as famine and disease, or by social actions such as war.

Darwin immediately applied Malthus's argument to animals and plants, and by 1838 he had arrived at a sketch of a theory of evolution through natural selection ( see Species and Speciation ). For the next two decades he worked on his theory and other natural history projects. (Darwin was independently wealthy and never had to earn an income.) In 1839 he married his first cousin, Emma Wedgwood (1808-96), and soon after, moved to a small estate, Down House, outside London. There he and his wife had ten children, three of whom died in infancy.

Darwin's theory was first announced in 1858 in a paper presented at the same time as one by Alfred Russel Wallace, a young naturalist who had come independently to the theory of natural selection. Darwin's complete theory was published in 1859, in On the Origin of Species. Often referred to as the "book that shook the world," the Origin sold out on the first day of publication and subsequently went through six editions.

Darwin's theory of evolution by natural selection is essentially that, because of the food-supply problem described by Malthus, the young born to any species intensely compete for survival. Those young that survive to produce the next generation tend to embody favorable natural variations (however slight the advantage may be)-the process of natural selection-and these variations are passed on by heredity. Therefore, each generation will improve adaptively over the preceding generations, and this gradual and continuous process is the source of the evolution of species. Natural selection is only part of Darwin's vast conceptual scheme; he also introduced the concept that all related organisms are descended from common ancestors. Moreover, he provided additional support for the older concept that the earth itself is not static but evolving.

Reactions to the Theory.

The reaction to the Origin was immediate. Some biologists argued that Darwin could not prove his hypothesis. Others criticized Darwin's concept of variation, arguing that he could explain neither the origin of variations nor how they were passed to succeeding generations. This particular scientific objection was not answered until the birth of modern genetics in the early 20th century ( see Heredity ; Mendel's Laws ). In fact, many scientists continued to express doubts for the following 50 to 80 years. The most publicized attacks on Darwin's ideas, however, came not from scientists but from religious opponents. The thought that living things had evolved by natural processes denied the special creation of humankind and seemed to place humanity on a plane with the animals; both of these ideas were serious contradictions to orthodox theological opinion.

Later Years.

Darwin spent the rest of his life expanding on different aspects of problems raised in the Origin. His later books-including The Variation of Animals and Plants Under Domestication (1868), The Descent of Man (1871), and The Expression of the Emotions in Animals and Man (1872)-were detailed expositions of topics that had been confined to small sections of the Origin. The importance of his work was well rec ognized by his contemporaries; Darwin was elected to the Royal Society (1839) and the French Academy of Sciences (1878). He was also honored by burial in Westminster Abbey after he died in Down, Kent, on April 19, 1882.

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