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Evolution



For more details on this topic, see Timeline of evolution.
Evolutionary tree showing the divergence of modern species from their common ancestor in the center. The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Evolutionary tree showing the divergence of modern species from their common ancestor in the center.[153] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.

Despite the uncertainty on how life began, it is clear that prokaryotes were the first organisms to inhabit Earth,[154] approximately 3–4 billion years ago.[155] No obvious changes in morphology or cellular organization occurred in these organisms over the next few billion years.[156]

The eukaryotes were the next major innovation in evolution. These came from ancient bacteria being engulfed by the ancestors of eukaryotic cells, in a cooperative association called endosymbiosis.[87][157] The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into either mitochondria or hydrogenosomes.[158] An independent second engulfment of cyanobacterial-like organisms led to the formation of chloroplasts in algae and plants.[159]

The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about a billion years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.[154][160] The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae, cyanobacteria, slime moulds and myxobacteria.[161]

Soon after the emergence of these first multicellular organisms, a remarkable amount of biological diversity appeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the majority of types of modern animals appeared in the fossil record, as well as unique lineages that subsequently became extinct.[162] Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen in the atmosphere from photosynthesis.[163] About 500 million years ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals.[164] Amphibians first appeared around 300 million years ago, followed by early amniotes, then mammals around 200 million years ago and birds around 100 million years ago (both from "reptile"-like lineages). However, despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this process continue to be highly successful and dominate the Earth, with the majority of both biomass and species being prokaryotes.[94]

History of evolutionary thought

For more details on this topic, see History of evolutionary thought.
Charles Darwin at age 51, just after publishing On the Origin of Species.
Charles Darwin at age 51, just after publishing On the Origin of Species.

Evolutionary ideas such as common descent and the transmutation of species have existed since at least the 6th century BC, when they were expounded by the Greek philosopher Anaximander.[165] Others who considered such ideas included the Greek philosopher Empedocles, the Roman philosopher-poet Lucretius, the Arab biologist Al-Jahiz,[166] the Persian philosopher Ibn Miskawayh, the Brethren of Purity,[167] and the Eastern philosopher Zhuangzi.[168] As biological knowledge grew in the 18th century, evolutionary ideas were set out by a few natural philosophers including Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.[169] The ideas of the biologist Jean-Baptiste Lamarck about transmutation of species had wide influence. Charles Darwin formulated his idea of natural selection in 1838 and was still developing his theory in 1858 when Alfred Russel Wallace sent him a similar theory, and both were presented to the Linnean Society of London in separate papers.[170] At the end of 1859 Darwin's publication of On the Origin of Species explained natural selection in detail and presented evidence leading to increasingly wide acceptance of the occurrence of evolution.

Gregor Mendel, who laid the foundation for genetics.
Gregor Mendel, who laid the foundation for genetics.

Debate about the mechanisms of evolution continued, and Darwin could not explain the source of the heritable variations which would be acted on by natural selection. Like Lamarck, he thought that parents passed on adaptations acquired during their lifetimes,[171] a theory which was subsequently dubbed Lamarckism.[172] In the 1880s August Weismann's experiments indicated that changes from use and disuse were not heritable, and Lamarckism gradually fell from favour.[173][174] More significantly, Darwin could not account for how traits were passed down from generation to generation. In 1865 Gregor Mendel found that traits were inherited in a predictable manner.[175] When Mendel's work was rediscovered in 1900, disagreements over the rate of evolution predicted by early geneticists and biometricians led to a rift between the Mendelian and Darwinian models of evolution.

This contradiction was reconciled in the 1930s by biologists such as Ronald Fisher. The end result was a combination of evolution by natural selection and Mendelian inheritance, the modern evolutionary synthesis.[176] In the 1940s, the identification of DNA as the genetic material by Oswald Avery and colleagues and the subsequent publication of the structure of DNA by James Watson and Francis Crick in 1953, demonstrated the physical basis for inheritance. Since then, genetics and molecular biology have become core parts of evolutionary biology and have revolutionized the field of phylogenetics.[12]

In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically-oriented disciplines, whose specialist training in particular organisms addressed general questions in evolution. As evolutionary biology expanded as an academic discipline, particularly after the development of the modern evolutionary synthesis, it began to draw more widely from the biological sciences.[12] Currently the study of evolutionary biology involves scientists from fields as diverse as biochemistry, ecology, genetics and physiology, and evolutionary concepts are used in even more distant disciplines such as psychology, medicine, philosophy and computer science.

Social and cultural responses

For more details on this topic, see Social effect of evolutionary theory.
Caricature of Charles Darwin as a quadrupedal ape, reflecting the cultural backlash against evolution.
Caricature of Charles Darwin as a quadrupedal ape, reflecting the cultural backlash against evolution.

Even before the publication of On the Origin of Species, the idea that life had evolved was an active source of debate. Evolution is still a contentious concept in some quarters outside the scientific community. Debate has centered on the philosophical, social and religious implications of evolution, not on the science itself; the proposition that biological evolution occurs through the mechanism of natural selection is standard in the scientific literature.[177]

Although many religions and denominations have reconciled their beliefs with evolution through various concepts of theistic evolution, there are many creationists who believe that evolution is contradicted by the creation myths found in their respective religions.[178] As Darwin recognized early on, the most controversial aspect of evolutionary thought is its implications for human origins. In some countries—notably the United States—these tensions between scientific and religious teachings have fueled the ongoing creation–evolution controversy, a religious conflict focusing on politics and public education.[179] While other scientific fields such as cosmology[180] and earth science[181] also conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more opposition from many religious believers.

Evolution has been used to support philosophical positions that promote discrimination and racism. For example, the eugenic ideas of Francis Galton were developed to argue that the human gene pool should be improved by selective breeding policies, including incentives for those considered "good stock" to reproduce, and the compulsory sterilization, prenatal testing, birth control, and even killing, of those considered "bad stock."[182] Another example of an extension of evolutionary theory that is now widely regarded as unwarranted is "Social Darwinism," a term given to the 19th century Whig Malthusian theory developed by Herbert Spencer into ideas about "survival of the fittest" in commerce and human societies as a whole, and by others into claims that social inequality, racism, and imperialism were justified.[183] However, contemporary scientists and philosophers consider these ideas to have been neither mandated by evolutionary theory nor supported by data.[184][185]

Applications

For more details on this topic, see Artificial selection and Evolutionary computation.

A major technological application of evolution is artificial selection, which is the intentional selection of certain traits in a population of organisms. Humans have used artificial selection for thousands of years in the domestication of plants and animals.[186] More recently, such selection has become a vital part of genetic engineering, with selectable markers such as antibiotic resistance genes being used to manipulate DNA in molecular biology.

As evolution can produce highly optimized processes and networks, it has many applications in computer science. Here, simulations of evolution using evolutionary algorithms and artificial life started with the work of Nils Aall Barricelli in the 1960s, and was extended by Alex Fraser, who published a series of papers on simulation of artificial selection.[187] Artificial evolution became a widely recognized optimization method as a result of the work of Ingo Rechenberg in the 1960s and early 1970s, who used evolution strategies to solve complex engineering problems.[188] Genetic algorithms in particular became popular through the writing of John Holland.[189] As academic interest grew, dramatic increases in the power of computers allowed practical applications, including the automatic evolution of computer programs.[190] Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software produced by human designers, and also to optimize the design of systems.[191]

Understanding macroevolution can have practical applications too. A certain species of coral might be discovered that produces an antibiotic compound with medical potential. Knowing its closest relatives would inform researchers of other species that might produce similar compounds, which could then be investigated.[192]

Further reading

Introductory reading

History of evolutionary thought

Advanced reading

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