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Evolutionary thought, the idea that species change over time, has roots in antiquity, in the ideas of the Greeks, Romans, Chinese and Muslims. However, until the 18th century, Western biological thinking was dominated by essentialism, the idea that living forms are unchanging. This started to change when, during the Enlightenment, evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history. Naturalists began to focus on the variability of species; the emergence of paleontology with the concept of extinction further undermined the static view of nature. In the early 19th century, Jean-Baptiste Lamarck proposed his theory of the transmutation of species, which was the first fully-formed scientific theory of evolution.

In 1858, Charles Darwin and Alfred Russel Wallace published a new evolutionary theory, which was explained in detail in Darwin's On the Origin of Species (1859). Unlike Lamarck, Darwin proposed common descent and a branching tree of life. The theory was based on the idea of natural selection, and it synthesized a broad range of evidence from animal husbandry, biogeography, geology, morphology, and embryology.

Darwin's work led to the rapid acceptance of evolution, but the mechanism he proposed, natural selection, was not widely accepted until the 1940s. Most biologists argued that other factors drove evolution, such as inheritance of acquired characteristics (neo-Lamarckism), an innate drive for change (orthogenesis), or sudden large mutations (saltationism). The synthesis of natural selection with Mendelian genetics during the 1920s and 1930s founded the new discipline of population genetics. Throughout the 1930s and 1940s, population genetics became integrated with other branches of biology, finally resulting in the current unified theory of evolution—the modern evolutionary synthesis.

Following the establishment of evolutionary biology, studies of mutation and variation in natural populations, combined with biogeography and systematics, led to sophisticated mathematical and causal models of evolution. Paleontology and comparative anatomy allowed more detailed reconstructions of the history of life. After the rise of molecular genetics in the 1950s, the field of molecular evolution developed, based on DNA, RNA, and protein sequences. The gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral theory of molecular evolution, sparking debates over adaptationism, the units of selection, and the relative importance of genetic drift versus natural selection. In the late 20th century, DNA sequencing led to molecular phylogenetics and the reorganization of the tree of life into the three-domain system. In addition, the newly recognized factors of symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary history.

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Antiquity

Greeks

Greek philosophers discussed ideas that involved forms of organic evolution. Anaximander (c. 610–546 BC) claimed that life had originally developed in the sea and only later moved onto land, and Empedocles (c. 490–430 BC) wrote of a non-supernatural origin for living things. Empedocles even suggested a form of natural selection, which Aristotle summarized as, "Wherever then all the parts came about just what they would have been if they had come be for an end, such things survived, being organized spontaneously in a fitting way; whereas those which grew otherwise perished and continue to perish..."

Plato (c. 428–348 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of evolutionism," as he established the philosophy of essentialism, which he called the Theory of Forms. This theory holds that objects observed in the real world are only reflections of a limited number of essences (eide). Variation is merely the result of an imperfect reflection of these constant essences. In his Timaeus, Plato set forth the idea that God had created the cosmos and everything in it because He is good, and hence, "... free from jealousy, He desired that all things should be as like Himself as they could be." God created all conceivable forms of life, since "... without them the universe will be incomplete, for it will not contain every kind of animal which it ought to contain, if it is to be perfect." This idea, that all potential forms of life are essential to a perfect creation, is called the plenitude principle, and it greatly influenced Christian thought.

Aristotle (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural historian whose work has been preserved in any real detail. His writings on biology were the result of his research into natural history on the isle of Lesbos, and have survived in the form of four books, usually known by their Latin names, De anima (on the essence of life), Historia animalium (inquiries about animals), De generatione animalium (reproduction), and De partibus animalium (anatomy). Aristotle's works contain some remarkably astute observations and interpretations, along with sundry myths and mistakes—reflecting the uneven state of knowledge during his time. However, for Charles Singer, "Nothing is more remarkable than efforts to the relationships of living things as a scala naturæ". This scala naturæ, described in Historia animalium, classified organisms in relation to a hierarchical "Ladder of Life" or "Chain of Being", placing them according to complexity of structure and function, with organisms that showed greater vitality and ability to move described as "higher organisms".

Chinese

Ideas on evolution were expressed by ancient Chinese thinkers such as Zhuangzi (Chang Tzu), a Taoist philosopher who lived around the 4th century BC. According to Joseph Needham, Taoism explicitly denied the fixity of biological species. Instead, humans, nature and the heavens were seen as existing in a state of "constant transformation" known as the Tao.

Romans

Titus Lucretius Carus (d. 50 BC), the Roman Epicurean and atomist, wrote the poem On the Nature of Things (De rerum natura), describing the development of the living Earth in stages: from atoms colliding in the void as swirls of dust to early plants and animals springing from the early Earth's substance, then a succession of animals, including a series of progressively less brutish humans. Lucretius may be seen as the earliest believer in hard inheritance. He said, "For if each organism had not its own genetic bodies, how could we with certainty assign each to its mother?". The essence of Lucretius' ideas was naturalism, and the avoidance of supernatural interventions or explanations.

Middle Ages

Islamic philosophy and the struggle for existence

Whereas Greek and Roman evolutionary ideas died out in Europe after the fall of the Roman Empire, they were not lost to Islamic scientists and philosophers. In the Islamic Golden Age, early theories on evolution were taught in Islamic schools. John William Draper, the 19th-century scientist, philosopher and historian, discussed the 12th-century writings of al-Khazini as part of what he called the "Mohammedan theory of evolution". He compared these early ideas to later biological theories, arguing that the former were developed "... much farther than we are disposed to do, extending them even to inorganic or mineral things."

The first Muslim biologist and philosopher to speculate in detail about evolution was the Afro-Arab writer al-Jahiz in the 9th century. He considered the effects of the environment on an animal's chances for survival, and described the struggle for existence. Ibn Miskawayh's al-Fawz al-Asghar and the Brethren of Purity's Encyclopedia of the Brethren of Purity (The Epistles of Ikhwan al-Safa) expressed ideas about how species developed: from matter into vapor and thence to water, then minerals into plants and then animals, leading to apes and, finally, humans. The polymath Ibn al-Haytham wrote a book in which he argued for evolutionism (although not natural selection). Numerous other Islamic scholars and scientists, such as Abū Rayhān al-Bīrūnī, Nasir al-Din Tusi, and Ibn Khaldun discussed and developed these ideas. Translated into Latin, these works began to appear in the West after the Renaissance and may have had an impact on Western science.

Christian philosophy and the great chain of being

During the Dark Ages, Greek classical learning was all but lost to the West. However, contact with the Islamic world, where Greek manuscripts were preserved and elaborated on, soon led to a massive spate of Latin translations in the 12th century. Europeans were thus re-introduced to the works of Plato and Aristotle, as well as to Islamic thought. Christian thinkers combined Aristotlean classification with Plato's ideas of the goodness of God, and of all potential life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings into a huge interconnected system: the scala naturæ, or great chain of being.

Within this system, everything that existed could be placed in order, from "lowest" to "highest", with Hell at the bottom and God at the top—below God, an angelic hierarchy marked by the orbits of the planets, mankind in an intermediate position, and worms the lowest of the animals. As the universe was ultimately perfect, the great chain was also perfect. There were no empty links in the chain, and no link was represented by more than one species. Therefore no species could ever move from one position to another. Thus, in this Christianized version of Plato's perfect universe, species could never change, but must remain forever fixed, in accordance with the text of Genesis. For humans to forget their position was seen as sinful, whether they behaved like lower animals or aspired to a higher station than was given them by their Creator.

Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in the saying: Error: {{Lang}}: text has italic markup (help) ("nature does not make leaps"). This basic concept of the great chain of being greatly influenced the thinking of Western civilization for centuries (and still has an influence today). It also formed a part of the argument from design presented by natural theology. As a classification system, it became the major organizing principle and foundation of the emerging science of biology in the 17th and 18th centuries.

Renaissance and Enlightenment

Some evolutionist theories explored between 1650 and 1800 postulated that the universe, including life on Earth, had developed mechanically, entirely without divine guidance. Around this time, the mechanical philosophy of René Descartes began to encourage the machine-like view of the universe which would come to characterise the scientific revolution. However, most contemporary theories of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution was a fundamentally spiritual process. In 1751, Pierre Louis Maupertuis veered toward more materialist ground. He wrote of natural modifications occurring during reproduction and accumulating over the course of many generations, producing races and even new species, and he anticipated in general terms the idea of natural selection.

Later in the 18th century, G. L. L. Buffon suggested that what most people referred to as species were really just well-marked varieties modified from an original form by environmental factors. For example, he believed that lions, tigers, leopards and house cats might all have a common ancestor. He speculated that the 200 or so species of mammals then known might have descended from as few as 38 original forms. Buffon’s evolutionary ideas were strictly limited; he believed each of the original forms had arisen through spontaneous generation and that they were shaped by "internal moulds" that limited the amount of change. Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not only the concept that man had descended from primates, but also that, in response to the environment, creatures had found methods of transforming their characteristics over long time intervals. Charles Darwin’s grandfather, Erasmus Darwin, published Zoönomia in 1796, which suggested that "all warm-blooded animals have arisen from one living filament". In his 1802 poem Temple of Nature, he described the rise of life from minute organisms living in the mud to a state of modern diversity.

Early 19th century

Paleontology and geology

In 1796, Georges Cuvier published his findings on the differences between living elephants and those found in the fossil record. His analysis demonstrated that mammoths and mastodons were distinct species different from any living animal, effectively ending a long-running debate over the possibility of the extinction of a species. In 1788 James Hutton described gradual geological processes operating continuously over deep time. William Smith began the process of ordering rock strata by examining fossils in the layers while he worked on his geologic map of England. Independently, in 1811, Georges Cuvier and Alexandre Brongniart published an influential study of the geologic history of the region around Paris, which was based on the stratigraphic succession of layers of rock. These works helped establish the antiquity of the Earth. Cuvier advocated catastrophism to explain the patterns of extinction and faunal succession revealed by the fossil record.

Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th century. By the 1840s, the outlines of the geologic timescale were becoming clear, and in 1841 John Phillips named three major eras, based on the predominant fauna of each: the Paleozoic, dominated by marine invertebrates and fish, the Mesozoic, the age of reptiles, and the current Cenozoic age of mammals. This progressive picture of the history of life was accepted even by conservative English geologists like Adam Sedgwick and William Buckland; however, like Cuvier, they attributed the progression to repeated catastrophic episodes of extinction followed by new episodes of creation. Unlike Cuvier, Buckland and some other advocates of natural theology among British geologists made efforts to explicitly link the last catastrophic episode proposed by Cuvier to the biblical flood.

From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which, building on Hutton's ideas, advocated a uniformitarian alternative to the catastrophic theory of geology. Lyell claimed that, rather than being the products of cataclysmic (and possibly supernatural) events, the geologic features of the Earth are better explained as the result of the same gradual geologic forces observable in the present day—but acting over immensely long periods of time. Although Lyell opposed evolutionary ideas (even questioning the consensus that the fossil record demonstrates a true progression), his concept that the Earth was shaped by forces working gradually over an extended period, and the immense age of the Earth assumed by his theories, would strongly influence future evolutionary thinkers such as Charles Darwin.

Transmutation of species

Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the transmutation of species. Lamarck did not believe that all living things shared a common ancestor but rather that simple forms of life were created continuously by spontaneous generation. He also believed that an innate life force drove species to become more complex over time, advancing up a linear ladder of complexity that was related to the great chain of being. Lamarck recognized that species were adapted to their environment. He explained this by saying that the same innate force driving increasing complexity caused the organs of an animal (or a plant) to change based on the use or disuse of that organ, just as muscles are affected by exercise. He argued that these changes would be inherited by the next generation and produce slow adaptation to the environment. It was this secondary mechanism of adaptation through the inheritance of acquired characteristics that would become known as Lamarckism and would influence discussions of evolution into the 20th century. A radical British school of comparative anatomy that included the anatomist Robert Grant was closely in touch with Lamarck's school of French Transformationism, which contained scientists such as Étienne Geoffroy Saint-Hilaire. Grant developed Lamarck's and Erasmus Darwin's ideas of transmutation and evolutionism, investigating homology to prove common descent. As a young student Charles Darwin joined Grant in investigations of the life cycle of marine animals. In 1826 an anonymous paper, probably written by Robert Jameson, praised Lamarck for explaining how higher animals had “evolved” from the simplest worms; this was the first use of the word “evolved” in a modern sense.

In 1844, the Scottish publisher Robert Chambers anonymously published an influential, and extremely controversial book of popular science entitled Vestiges of the Natural History of Creation. This book proposed an evolutionary scenario for the origins of the Solar System and life on Earth. It claimed that the fossil record showed a progressive ascent of animals with current animals being branches off a main line that leads progressively to humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been woven into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of radicals like Robert Grant, but its implication that humans were only the last step in the ascent of animal life incensed many conservative thinkers. The high profile of the public debate over Vestiges, with its depiction of evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later.

Ideas about the transmutation of species were associated with the radical materialism of the Enlightenment and were attacked by more conservative thinkers. Georges Cuvier attacked the ideas of Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed that the individual parts of an animal were too closely correlated with one another to allow for one part of the anatomy to change in isolation from the others, and argued that the fossil record showed patterns of catastrophic extinctions followed by re-population, rather than gradual change over time. He also noted that drawings of animals and animal mummies from Egypt, which were thousands of years old, showed no signs of change when compared with modern animals. The strength of Cuvier's arguments and his scientific reputation helped keep transmutational ideas out of the mainstream for decades.

In Britain the philosophy of natural theology remained influential. William Paley's 1802 book Natural Theology with its famous watchmaker analogy had been written at least in part as a response to the transmutational ideas of Erasmus Darwin. Geologists influenced by natural theology, such as Buckland and Sedgwick, made a regular practice of attacking the evolutionary ideas of Lamarck, Grant, and The Vestiges of the Natural History of Creation. Although the geologist Charles Lyell opposed scriptural geology, he also believed in the immutability of species, and in his Principles of Geology (1830–1833), he criticized Lamarck's theories of development. Idealists such as Louis Agassiz and Richard Owen believed that each species was fixed and unchangeable because it represented an idea in the mind of the creator. They believed that relationships between species could be discerned from developmental patterns in embryology, as well as in the fossil record, but that these relationships represented an underlying pattern of divine thought, with progressive creation leading to increasing complexity and culminating in humanity. Owen developed the idea of "archetypes" in the Divine mind that would produce a sequence of species related by anatomical homologies, such as vertebrate limbs. Owen led a public campaign that successfully marginalized Robert Grant in the scientific community. Darwin would make good use of the homologies analyzed by Owen in his own theory, but the harsh treatment of Grant, and the controversy surrounding Vestiges, would contribute to his decision to delay publishing his ideas.

Anticipations of natural selection

Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of Species published in 1861 Darwin named those he knew about in an introductory appendix, An Historical Sketch of the Recent Progress of Opinion on the Origin of Species, which he expanded in later editions.

In 1813, William Charles Wells read before the Royal Society essays assuming that there had been evolution of humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel Wallace were unaware of this work when they jointly published the theory in 1858, but Darwin later acknowledged that Wells had recognised the principle before them, writing that the paper "An Account of a White Female, part of whose Skin resembles that of a Negro" was published in 1818, and "he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone." When Darwin was developing his theory, he was influenced by Augustin de Candolle's natural system of classification, which laid emphasis on the war between competing species.

Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of "continual balancing of life to circumstance. ... progeny of the same parents, under great differences of circumstance, might, in several generations, even become distinct species, incapable of co-reproduction." Charles Darwin discovered this work after the initial publication of the Origin. In the brief historical sketch that Darwin included in the 3rd addition he says "Unfortunately the view was given by Mr. Matthew very briefly in an Appendix to a work on a different subject ... He clearly saw, however, the full force of the principle of natural selection."

It is possible to look through the history of biology from the Ancient Greeks onwards and discover anticipations of almost all of Darwin's key ideas. However, as the historian of science Peter J. Bowler says "Through a combination of bold theorizing and comprehensive evaluation. Darwin came up with a concept of evolution that was unique for the time." Bowler goes on to say that simple priority alone is not enough to secure a place in the history of science; someone has to develop an idea and convince others of its importance to have a real impact.

T. H. Huxley said in his essay on the reception of the Origin of Species:

The suggestion that new species may result from the selective action of external conditions upon the variations from their specific type which individuals present and which we call spontaneous because we are ignorant of their causation is as wholly unknown to the historian of scientific ideas as it was to biological specialists before 1858. But that suggestion is the central idea of the Origin of Species, and contains the quintessence of Darwinism.

Natural selection

The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands during the voyage of the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first of a series of secret notebooks on transmutation. Darwin's observations lead him to view transmutation as a process of divergence and branching, rather than the ladder-like progression envisioned by Lamarck and others. In 1838 he read the new 6th edition of An Essay on the Principle of Population, written in the late 1700s by Thomas Malthus. Malthus' idea of population growth leading to a struggle for survival combined with Darwin's knowledge on how breeders selected traits, led to the inception of Darwin's theory of natural selection. Darwin did not publish his ideas on evolution for 20 years. However he did share them with certain other naturalists and friends, starting with Joseph Hooker with whom he discussed his unpublished 1844 essay on natural selection. During this period he used the time he could spare from his other scientific work to slowly refine his ideas and, aware of the intense controversy around transmutation, amass evidence to support them.

Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of Creation, already suspected that transmutation of species occurred when he began his career as a naturalist. By 1855 his biogeographical observations during his field work in South America and the Malay Archipelago made him confident enough in a branching pattern of evolution to publish a paper that stated that every species originated in close proximity to an already existing closely allied species. Like Darwin, it was Wallace's consideration of how the ideas of Malthus might apply to animal populations that lead him to conclusions very similar to those reached by Darwin about the role of natural selection. In February 1858 Wallace, unaware of Darwin's unpublished ideas, wrote up his thoughts into an essay and mailed them to Darwin, asking for his opinion. The result was the joint publication in July of an extract from Darwin's 1844 essay along with Wallace's letter. Darwin also began work in earnest on The Origin of Species, which he would publish in 1859.

1859–1930s: Darwin and his legacy

By the 1850s whether or not species evolved was a subject of intense debate, with promininent scientists arguing both sides of the issue. However, it was the publication of Charles Darwin's On the Origin of Species (1859) that fundamentally transformed the discussion over biological origins. Darwin argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy, embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary change could persist: his theory of natural selection.

One of the first and most important naturalists to be convinced by Origin of the reality of evolution was the British anatomist Thomas Henry Huxley. Huxley recognized that unlike the earlier transmutational ideas of Lamarck and Vestiges, Darwin's theory provided a mechanism for evolution without supernatural involvement even if Huxley himself was not completely convinced that natural selection was the key evolutionary mechanism. Huxley would make advocacy of evolution a cornerstone of the program of the X-club to reform and professionalize science by displacing natural theology with naturalism, ending the domination of British natural science by the clergy. By the early 1870s in English-speaking countries, thanks partly to these efforts, evolution had become the mainstream scientific explanation for the origin of species. In his campaign for public and scientific acceptance of Darwin's theory, Huxley made extensive use of new evidence for evolution from paleontology. This included evidence that birds had evolved from reptiles, including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth found in North America. Another important line of evidence was the finding of fossils that helped trace the evolution of the horse from its small five-toed ancestors. However, acceptance of evolution among scientists in non-English speaking nations such as France, and the countries of southern Europe and Latin America was slower. An exception to this was Germany, where both August Weismann and Ernst Haeckel championed this idea: with Haeckel using evolution to challenge the established tradition of metaphysical idealism in German biology, much as Huxley used it to challenge natural theology in Britain.

Darwin's theory succeeded in profoundly shaking scientific opinion regarding the development of life and resulted in a small social revolution. However, this theory could not explain several critical components of the evolutionary process. Namely, Darwin was unable to explain the source of variation in traits within a species, and could not identify a mechanism that could pass traits faithfully from one generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the inheritance of acquired characteristics, proved to be useful for statistical models of evolution that were developed by his cousin Francis Galton and the "biometric" school of evolutionary thought. This idea was, however, of little use to biologists.

Application to humans

Charles Darwin was aware of the severe reaction in some parts of the scientific community against the suggestion made in Vestiges of the Natural History of Creation that humans had arisen from animals by a process of transmutation. Therefore he almost completely ignored the topic of human evolution in The Origin of Species. Despite this precaution, the issue featured prominently in the debate that followed the book's publication. For most of the first half of the 19th century, the scientific community believed that, although geology had shown that the Earth and life were very old, human beings had appeared suddenly just a few thousand years before the present. However, a series of archaeological discoveries in the 1840s and 1850s showed stone tools associated with the remains of extinct animals. By the early 1860s, as summarized in Charles Lyell's 1863 book Geological Evidences of the Antiquity of Man, it had become widely accepted that humans had existed during a prehistoric period – which stretched many thousands of years before the start of written history. This view of human history was more compatible with an evolutionary origin for humanity than was the older view. On the other hand, at that time there was no fossil evidence to demonstrate human evolution. The only human fossils found before the discovery of Java man in the 1890s were either of anatomically modern humans, or of Neanderthals that were too close, especially in the critical characteristic of cranial capacity, to modern humans for them to be convincing intermediates between humans and other primates.

Therefore the debate that immediately followed the publication of The Origin of Species centered on the similarities and differences between humans and modern apes. Richard Owen vigorously defended the traditional classification, suggested by Carolus Linnaeus and Cuvier, that placed humans in a separate order from any of the other mammals. On the other hand, Thomas Henry Huxley sought to demonstrate a close anatomical relationship between humans and apes. In one famous incident, Huxley showed that Owen was mistaken in claiming that the brains of gorillas lacked a structure present in human brains. Huxley summarized his argument in his highly influential 1863 book Evidence as to Man's Place in Nature. Another viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. They agreed that humans shared a common ancestor with apes, but questioned whether any purely materialistic mechanism could account for all the differences between humans and apes, especially some aspects of the human mind.

In 1871, Darwin published The Descent of Man, and Selection in Relation to Sex, which contained his views on human evolution. Darwin argued that the differences between the human mind and the minds of the higher animals were a matter of degree rather than of kind. For example, he viewed morality as a natural outgrowth of instincts that were beneficial to animals living in social groups. He argued that all the differences between humans and apes were explained by a combination of the selective pressures that came from our ancestors moving from the trees to the plains, and sexual selection. The debate over human origins, and over the degree of human uniqueness continued well into the 20th century.

Alternatives to natural selection

Evolution was widely accepted in scientific circles within a few years of the publication of Origin, but the acceptance of natural selection as its driving mechanism was much less widespread. The four major alternatives to natural selection in the late 19th century were theistic evolution, neo-Lamarckism, orthogenesis, and saltationism. Theistic evolution was the idea that God intervened in the process of evolution to guide it in such a way that the living world could still be considered to be designed. However, this idea rapidly fell out of favor among scientists. They became more and more committed to the idea of naturalism and came to believe that direct appeals to supernatural involvement were scientifically unproductive and a form of special pleading. By 1900, theistic evolution had completely disappeared from mainstream scientific discussions.

In the late 19th century, the term neo-Lamarckism came to be associated with the position of naturalists who viewed the inheritance of acquired characteristics as the most important evolutionary mechanism. Advocates of this position included the British writer and Darwin critic Samuel Butler, the German biologist Ernst Haeckel, and the American paleontologist Edward Drinker Cope. They considered Lamarckism to be philosophically superior to Darwin's idea of selection acting on random variation. Cope looked for, and thought he found, patterns of linear progression in the fossil record. Inheritance of acquired characteristics was part of Haeckel's recapitulation theory of evolution, which held that the embryological development of an organism repeats its evolutionary history. Critics of neo-Lamarckism, such as the German biologist August Weismann and Alfred Russel Wallace, pointed out that no one had ever produced solid evidence for the inheritance of acquired characteristics. Despite these criticisms, neo-Lamarckism remained the most popular alternative to natural selection at the end of the 19th century, and would remain the position of some naturalists well into the 20th century.

Orthogenesis was the hypothesis that life has an innate tendency to change, in a unilinear fashion, towards ever-greater perfection. It had a significant following in the 19th century, and its proponents included the Russian biologist Leo Berg and the American paleontologist Henry Fairfield Osborn. Orthogenesis was popular among some paleontologists, who believed that the fossil record showed a gradual and constant unidirectional change. Saltationism was the idea that new species arise as a result of large mutations. It was seen as a much faster alternative to the Darwinian concept of a gradual process of small random variations being acted on by natural selection, and was popular with early geneticists such as Hugo de Vries, William Bateson, and early in his career, T. H. Morgan. It became the basis of the mutation theory of evolution.

Mendelian genetics, biometrics, and mutation

The rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate between two camps of biologists. In one camp were the Mendelians, who were focused on discrete variations and the laws of inheritance. They were led by William Bateson (who coined the word genetics) and Hugo de Vries (who coined the word mutation). Their opponents were the biometricians, who were interested in the continuous variation of characteristics within populations. Their leaders, Karl Pearson and Walter Frank Raphael Weldon, followed in the tradition of Francis Galton, who had focused on measurement and statistical analysis of variation within a population. The biometricians rejected Mendelian genetics on the basis that discrete units of heredity, such as genes, could not explain the continuous range of variation seen in real populations. Weldon's work with crabs and snails provided evidence that selection pressure from the environment could shift the range of variation in wild populations, but the Mendelians maintained that the variations measured by biometricians were too insignificant to account for the evolution of new species.

When T. H. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster, he was a saltationist who hoped to demonstrate that a new species could be created in the lab by mutation alone. Instead, the work at his lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid experimental evidence linking it to chromosomal inheritance. His work also demonstrated that most mutations had relatively small effects, such as a change in eye color, and that rather than creating a new species in a single step, mutations served to increase variation within the existing population.

1920s–1940s

Population genetics

The Mendelian and biometrician models were eventually reconciled with the development of population genetics. A key step was the work of the British biologist and statistician R.A. Fisher. In a series of papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher showed that the continuous variation measured by the biometricians could be produced by the combined action of many discrete genes, and that natural selection could change gene frequencies in a population, resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane, applied statistical analysis to real-world examples of natural selection, such as the evolution of industrial melanism in peppered moths, and showed that natural selection worked at an even faster rate than Fisher assumed.

The American biologist Sewall Wright, who had a background in animal breeding experiments, focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and Wright founded the discipline of population genetics. This integrated natural selection with Mendelian genetics, which was the critical first step in developing a unified theory of how evolution worked.

Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe that Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they observed in the living world. However, as the field of genetics continued to develop, those views became less tenable. Theodosius Dobzhansky, a postdoctoral worker in T. H. Morgan's lab, had been influenced by the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the divide between the theoretical foundations of microevolution developed by the population geneticists and the the field biologists, who were largely concerned with macroevolution, with his 1937 book Genetics and the Origin of Species. Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the assumptions of the population geneticists, these populations had large amounts of genetic diversity, with marked differences between sub-populations. The book also took the highly mathematical work of the population geneticists and put it into a more accessible form.

Ernst Mayr was influenced by the work of the German biologist Bernhard Rensch on how local environmental factors influenced the geographic distribution of sub-species and closely related species. Mayr followed up on Dobzhansky's work with the 1942 book Systematics and the Origin of Species, which emphasized the importance of allopatric speciation in the formation of new species. This form of speciation occurs when the geographical isolation of a sub-population is followed by the development of mechanisms for reproductive isolation. Mayr also formulated the biological species concept that defined a species as a group of interbreeding or potentially interbreeding populations that were reproductively isolated from all other populations.

In the 1944 book Tempo and Mode in Evolution, George Gaylord Simpson showed that the fossil record was consistent with the irregular non-directional pattern predicted by the developing evolutionary synthesis, and that the linear trends that earlier paleontologists had claimed supported orthogenesis and neo-Lamarckism did not hold up to closer examination. In 1950, G. Ledyard Stebbins published Variation and Evolution in Plants, which helped to integrate botany into the synthesis. The emerging cross-disciplinary consensus on the workings of evolution would be known as the modern evolutionary synthesis. It received its name from the book Evolution: The Modern Synthesis by Julian Huxley.

1940s–1960s: Molecular biology

In the 1940s, following up on Griffith's experiment on bacterial transformation, Avery, MacLeod and McCarty definitively identified deoxyribonucleic acid (DNA) as the transforming principle responsible for transmitting genetic information. In 1953, Francis Crick and James D. Watson published their famous paper on the structure of DNA, based on the research of Rosalind Franklin and Maurice Wilkins. These developments ignited the era of molecular biology and led to the understanding of evolution as a molecular process: the mutation of segments of DNA.

During this era of molecular biology, it became clear that a major mechanism causing variation within a population was DNA mutation. In the mid-1970s, Motoo Kimura formulated the neutral theory of molecular evolution, establishing the importance of genetic drift as a major mechanism of evolution. The theory sparked the "neutralist–selectionist" debate, partially solved by the development of the nearly neutral theory of evolution by Tomoko Ohta.

Late 20th century

Gene-centered view

In the mid-1960s, George C. Williams strongly critiqued explanations of adaptations worded in terms of "survival of the species" (group selection arguments). Such explanations were largely replaced by a gene-centered view of evolution, epitomized by the kin selection arguments of W. D. Hamilton, George R. Price and John Maynard Smith. This viewpoint would be summarized and popularized in the influential 1976 book The Selfish Gene by Richard Dawkins. Models of the period showed that group selection was severely limited in its strength; these models have since been shown to be too limited and newer models do admit the possibility of significant multi-level selection.

In 1973, Leigh Van Valen proposed the term "Red Queen", which he took from Through the Looking Glass by Lewis Carroll, to describe a scenario where a species involved in one or more evolutionary arms races would have to constantly change just to keep pace with the species with which it was co-evolving. Hamilton, Williams and others suggested that this idea might explain the evolution of sexual reproduction: the increased genetic diversity caused by sexual reproduction would help maintain resistance against rapidly evolving parasites, thus making sexual reproduction common, despite the tremendous cost from the gene-centric point of view of a system where only half of an organism's genome is passed on during reproduction. The gene-centric view has also led to an increased interest in Darwin's old idea of sexual selection, and more recently in topics such as sexual conflict and intragenomic conflict.

Sociobiology

W. D. Hamilton's work on kin selection contributed to the emergence of the discipline of sociobiology. The existence of altruistic behaviors has been a difficult problem for evolutionary theorists from the beginning. Significant progress was made in 1964 when Hamilton formulated the inequality in kin selection known as Hamilton's rule, which showed how eusociality in insects (the existence of sterile worker classes) and many other examples of altruistic behavior could have evolved through kin selection. Other theories followed, some derived from game theory, such as reciprocal altruism. In 1975, E.O. Wilson published the influential and highly controversial book Sociobiology: The New Synthesis which claimed evolutionary theory could help explain many aspects of animal, including human, behavior. Critics of sociobiology, including Stephen Jay Gould and Richard Lewontin, claimed that sociobiology greatly overstated the degree to which complex human behaviors could be determined by genetic factors. They also claimed that the theories of sociobiologists often reflected their own ideological biases. Despite these criticisms, work has continued in sociobiology and the related discipline of evolutionary psychology, including work on other aspects of the altruism problem.

Evolutionary paths and processes

One of the most prominent debates arising during the 1970s was over the theory of punctuated equilibrium. It was proposed by Niles Eldredge and Stephen Jay Gould who said there was a pattern of fossil species remaining largely unchanged for long periods (what they termed stasis), with relatively brief periods of rapid change during speciation. Improvements in sequencing methods resulted in a large increase of sequenced genomes, allowing the testing and refining of evolutionary theories using this huge amount of genome data. This research is providing insights into the molecular mechanisms of speciation and adaptation. Such genomic analysis has produced fundamental changes, such as the three-domain system of Carl Woese, in our understanding of the evolutionary history of life. Advances in computational hardware and software have allowed for the testing and extrapolation of increasingly advanced evolutionary models and the development of the field of systems biology. One of the results has been an exchange of ideas between theories of biological evolution and the field of computer science known as evolutionary computation, which uses processes that attempt to mimic biological evolution in the development of new computer algorithms. Discoveries in biotechnology are now producing methods for the synthesis and modification of entire genomes, driving evolutionary studies to the level where future experiments may involve the creation of entirely synthetic organisms.

Microbiology and horizontal gene transfer

Microbiology was largely ignored by early evolutionary theory. This was due to the paucity of morphological traits and the lack of a species concept in microbiology, particularly amongst prokaryotes. Now, evolutionary researchers are taking advantage their improved understanding of microbial physiology and ecology, produced by the comparative ease of microbial genomics, to explore the taxonomy and evolution of these organisms. These studies are revealing completely unanticipated levels of diversity amongst microbes, demonstrating that these organisms are the dominant form of life on Earth.

One particularly important outcome from studies on microbial evolution was the discovery in Japan of horizontal gene transfer in 1959. This transfer of genetic material between different species of bacteria has played a major role in the propagation of antibiotic resistance. More recently, as knowledge of genomes has continued to expand, it has been suggested that lateral transfer of genetic material has played an important role in the evolution of all organisms. Indeed, as part of the endosymbiotic theory for the origin of organelles, horizontal gene transfer has been a critical step in the evolution of eukaryotes such as fungi, plants, and animals.

Evolutionary developmental biology

In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under increasing scrutiny. There was a renewal of structuralist themes in evolutionary biology in the work of biologists such as Brian Goodwin and Stuart Kauffman, which incorporated ideas from cybernetics and systems theory, and emphasized the self-organizing processes of development as factors directing the course of evolution. The evolutionary biologist Stephen Jay Gould revived earlier ideas of heterochrony, alterations in the relative rates of developmental processes over the course of evolution, to account for the generation of novel forms, and, with the evolutionary biologist Richard Lewontin, wrote an influential paper in 1979 suggesting that a change in one biological structure could arise incidentally as an accidental result of selection on another structure, rather than through direct selection for that particular adaptation. They called this exaptation.

Molecular data regarding the mechanisms underlying development accumulated rapidly during the 1980s and '90s. It became clear that the diversity of animal morphology was not the result of different sets of proteins regulating the development of different animals, but from changes in the deployment of a small set of proteins that were common to all animals. These proteins became known as the "developmental toolkit". Such perspectives influenced the disciplines of phylogenetics, paleontology and comparative developmental biology, and spawned the new discipline of "evo-devo."

More recent work in this field has emphasized phenotypic and developmental plasticity. It has been suggested, for example, that the rapid emergence of basic animal body plans in the Cambrian explosion was due in part to changes in the environment acting on inherent material properties of cell aggregates, such as differential cell adhesion and biochemical oscillation. The resulting forms were later stabilized by natural selection. Experimental and theoretical research on these and related ideas has been presented in the multi-authored volume Origination of Organismal Form.

Unconventional ideas

Gaia hypothesis

Pierre Teilhard de Chardin formulated theories describing the gradual development of the Universe from subatomic particles to human society, considered by Teilhard as the last stage (see Gaia theory), but his ideas were not accepted by the scientific community. However, this hypothesis was later developed in a more limited and rigorous form by James Lovelock, who proposed that the living and nonliving parts of Earth can be viewed as a complex interacting system with similarities to a single organism. It has also been viewed as an extension of endosymbiosis and exosymbiosis. This modified hypothesis postulates that all living things have a regulatory effect on the Earth's environment that promotes life overall. Although not fully accepted by the scientific community, this hypothesis has been a useful spur to further research and is a topic of current scientific debate.

See also

• The Voyage of the Beagle
• Galápagos Islands
• Faith and rationality

Notes

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References

• Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press. ISBN 0-52023693-9.
• Bowler, Peter J. (2005). Making Modern Science. The University of Chicago Press. ISBN 0-226-06861-7. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
• Darlington, Cyril (1959). Darwin's place in history. Blackwell, Oxford, p85.
• Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (1st ed.). John Murray, London.
• Darwin, Charles (1861). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (3rd ed.). John Murray, London.
• Darwin, Charles (1866). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (3rd ed.). John Murray, London.
• Darwin, Charles (1872). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (6th ed.). John Murray, London.
• Darwin, Erasmus (1825). The Temple of Nature, or The Origin of Society.
• Darwin, Erasmus (1818). Zoonomia.
• Desmond, Adrian (1994). Darwin: The Life of a Tormented Evolutionist. W. W. Norton & Company. ISBN 0393311503. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
• Draper, John William (1878). History of the Conflict Between Religion and Science.
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Belknap Press of Harvard University Press. ISBN 0-674-00613-5.
• Henderson, Jan-Andrew (2000). The Emperor's Kilt: The Two Secret Histories of Scotland. Mainstream Publishing.
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• Lovejoy, Arthur (1936). The Great Chain of Being: A Study of the History of an Idea. Harvard University Press. ISBN 0-674-36153-9.
• Mayr, Ernst (1982). The Growth of Biological Thought: Diversity, Evolution, and Inheritance. The Belknap Press of Harvard University Press. ISBN 0-674-36445-7.
• Mayr, Ernst (1998). The Evolutionary Synthesis: Perspectives on the Unification of Biology. Harvard University Press. ISBN 0-674-27225-0. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
• Needham, Joseph (1995). The Shorter Science and Civilisation in China: An Abridgement of Joseph Needham's Original Text, Vol. 1. Cambridge University Press. ISBN 0521292867. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
• Singer, Charles (1931). A Short History of Biology. Clarendon Press.

External links

• UC Berkeley's History of Evolutionary Thought
• Darwin's precursors and influences by John Wilkins. Part of the Talk.Origins Archive.
• The Alfred Russel Wallace Page

 This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed. (1911). Encyclopædia Britannica (11th ed.). Cambridge University Press. {{cite encyclopedia}}: Missing or empty |title= (help)

• History of evolutionary biology
• History of science
• History of ideas