Encyclopedia article
Evolution
Change in the heritable characteristics of biological populations over successive generations, and the scientific theory that explains it.
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See every claimEvolution is the change in the heritable characteristics of biological populations over successive generations 1. The University of California Museum of Paleontology's Understanding Evolution resource defines it as descent with inherited modification, which covers both small-scale changes in gene-variant frequencies and the descent of species from shared ancestors 10. The theory of evolution by natural selection was conceived independently by Charles Darwin and Alfred Russel Wallace in the mid-19th century and was first set out in detail in Darwin's On the Origin of Species, first published in 1859 117. Current research debates concern the relative importance of selection and neutral processes, the predictability of evolution and the scope of evolutionary theory 348. One recent paper describes evolution by natural selection as well established across science and society 4.
Definition and heredity
#Evolution happens when individuals in a population vary in their genetic characteristics and in the numbers of their surviving offspring 1. It is often defined as a change in the frequency of alleles, the variants of a gene found at a specific locus, in a population 115. Because alleles do not necessarily have a visible effect, evolution in the strict biological sense can occur even when an organism's appearance or behaviour is unchanged 1.
Encyclopaedia Britannica describes evolution as a theory postulating that living things have their origin in other preexisting types, with differences due to modifications in successive generations, and calls it one of the fundamental keystones of modern biological theory 11.
In the modern synthesis, the basis for heredity is in DNA molecules that pass information from generation to generation 1.
Mechanisms
#Mutations are changes in the DNA sequence and are the ultimate source of genetic variation in all organisms 1. About half of the mutations in the coding regions of protein-coding genes are deleterious and the other half neutral, with a small percentage conferring a fitness benefit 1.
Natural selection is the process by which traits that enhance survival and reproduction become more common in successive generations 1. It rests on observations that more offspring are often produced than can survive, that traits vary among individuals, that different traits confer different rates of survival and reproduction, and that traits are heritable 1. Fitness is indicated by the proportion of subsequent generations carrying an organism's genes, not simply by its number of offspring 1.
Genetic drift is the random fluctuation of allele frequencies from one generation to the next, and fixation of a neutral allele through drift is more rapid in smaller populations 1. Gene flow is the exchange of genes between populations and species, including horizontal gene transfer, which is most common among bacteria and contributes to the spread of antibiotic resistance 1. Recombination and reassortment do not alter allele frequencies but produce new combinations of alleles 1.
Outcomes: adaptation, speciation and extinction
#Adaptation is the process that makes organisms better suited to their habitat, and adaptations are produced by natural selection 1. One observed example is the evolution, in a long-term laboratory experiment, of Escherichia coli able to use citric acid as a nutrient 1.
Speciation is the divergence of a species into two or more descendant species, and it has been observed both in laboratory conditions and in nature 1. There is no single agreed definition of species; Ernst Mayr defined the Biological Species Concept in 1942 1.
Nearly all species that have lived on Earth are extinct, with an estimated more than 99% lost 1. The Permian–Triassic extinction event drove approximately 96% of marine species to extinction 1. Present-day extinction rates are reported as 100–1000 times the background rate 1.
History of life
#The Earth is about 4.54 billion years old, and the earliest undisputed evidence of life dates from at least 3.5 billion years ago 1. All life on Earth is traced to a last universal common ancestor that lived approximately 3.5–3.8 billion years ago 1. Oxygen concentrations first rose in the Great Oxygenation Event 2.4 billion years ago, which a timetable of evolution published in Science Advances says facilitated the emergence of eukaryotes; a later Neoproterozoic oxygenation is associated with large, complex multicellular organisms 16.
Eukaryotic cells emerged between 1.6 and 2.7 billion years ago 1. Multicellular organisms began to appear around 1.7 billion years ago 1. The Cambrian explosion, beginning approximately 538.8 million years ago, saw most types of modern animals appear in the fossil record over about 10 million years 1.
Morphological and biochemical traits tend to be more similar among species that share a more recent common ancestor 1. DNA comparisons show that humans and chimpanzees share 98% of their genomes 1.
Dates for human evolution vary between sources. Early hominins in Africa date to around 6 million years ago, and human–gorilla divergence may be as early as 8 million years ago 16. The oldest fossil attributed to Homo is dated between ash beds of about 2.82 and 2.67 million years 16, and Britannica gives about 2.8–2.75 million years ago in Ethiopia 18. For modern humans, 16 states that anatomically modern humans emerged after 200,000 years ago, 1 gives around 250,000 years ago, and Britannica reports the oldest known Homo sapiens remains at about 315,000 years ago 18.
Estimates of current species diversity also differ. One estimate gives 10–14 million species, with about 1.9 million named 1; Britannica reports more than 2 million named species and total estimates of 10–30 million 11.
History of evolutionary thought
#Proposals that one type of organism could descend from another go back to early Greek writers including Anaximander and Empedocles 117. According to 1, Ibn Khaldun's Muqaddimah (1377) asserted that humans developed from 'the world of the monkeys'. The sources used here do not describe how this passage relates to modern evolutionary theory 1.
Jean-Baptiste Lamarck's 1809 transmutation theory has been called the first full-fledged evolutionary scheme 1.
Darwin's On the Origin of Species was first published in 1859 17. Darwin did not use the term 'evolution' in the first edition, referring instead to 'descent with modification' 13. Wallace also arrived at the idea of natural selection independently 17. According to the Open University, his view differed from Darwin's in that he did not think natural selection was sufficient to account for the origin of man and held that divine intervention had been required 17.
In the 1880s August Weismann's germ-plasm theory defended natural selection, and after Mendel's work was rediscovered in 1900, Hugo de Vries proposed mutationism, which discounted natural selection 17. In the 1920s and 1930s R. A. Fisher, J. B. S. Haldane and Sewall Wright used mathematical arguments to support natural selection and provide a framework for integrating genetics into Darwin's theory, and Theodosius Dobzhansky's Genetics and the Origin of Species (1937) combined Darwinian selection and Mendelian genetics 17. The resulting modern synthesis has guided evolutionary biology for over 50 years 8.
Current scientific debates
#The relative importance of selection and neutral processes is an area of current research 1.
For human populations, a review in Nature Reviews Genetics states that scientists have debated for over a century how much present-day variation results from natural selection versus neutral processes such as drift or population history 3. It reports that ancestral recombination graphs and ancient DNA have increased the potential power of selection scans, but that there is little consensus on how to detect and interpret selection signatures while avoiding confounders 3. Large genetic biobanks with genome or exome sequencing are also being used to study human evolution 7.
How predictable evolution is remains debated 4. A paper in Nature Reviews Biodiversity argues that prediction can be improved by understanding interactions between sources of selection, which it terms 'complex selection', rather than individual or summed effects 4. In human evolution, a paper in Evolutionary Anthropology states that the field lacks a widely accepted unifying explanation and argues that it need not have one, proposing a framework of 'gradients' for problems of rising magnitude 6.
Kevin Laland and colleagues have argued for an Extended Evolutionary Synthesis, holding that the modern synthesis is not wrong but does not capture the full richness of evolution, including inheritance beyond genes and the role of development 8. Other biologists have pushed back, saying there is little evidence that such a paradigm shift is warranted; both sides presented their cases at a Royal Society meeting 8. A related position discussed by BioLogos holds that organisms, not genes, should be the focal point of evolution, while random genetic mutations and natural selection continue to play a vital role 5.
Applications
#Viruses, bacteria, fungi and cancers evolve resistance to drugs and immune defences 1. According to Quanta Magazine, scientists used the modern synthesis to explain, among other things, why some people are prone to genetic disorders such as sickle-cell anaemia and why pesticides sooner or later fail to keep farm pests in check 8. In computer science, Ingo Rechenberg's work in the 1960s helped establish artificial evolution as a recognised optimisation method 1.
Scripture
Passages quoted from the King James Version. The text is fetched, never written by a model.
And God said, Let the waters bring forth abundantly the moving creature that hath life, and fowl that may fly above the earth in the open firmament of heaven. And God created great whales, and every living creature that moveth, which the waters brought forth abundantly, after their kind, and every winged fowl after his kind: and God saw that it was good. And God blessed them, saying, Be fruitful, and multiply, and fill the waters in the seas, and let fowl multiply in the earth. And the evening and the morning were the fifth day. And God said, Let the earth bring forth the living creature after his kind, cattle, and creeping thing, and beast of the earth after his kind: and it was so. And God made the beast of the earth after his kind, and cattle after their kind, and every thing that creepeth upon the earth after his kind: and God saw that it was good. And God said, Let us make man in our image, after our likeness: and let them have dominion over the fish of the sea, and over the fowl of the air, and over the cattle, and over all the earth, and over every creeping thing that creepeth upon the earth. So God created man in his own image, in the image of God created he him; male and female created he them. And God blessed them, and God said unto them, Be fruitful, and multiply, and replenish the earth, and subdue it: and have dominion over the fish of the sea, and over the fowl of the air, and over every living thing that moveth upon the earth.
For thou hast possessed my reins: thou hast covered me in my mother’s womb. I will praise thee; for I am fearfully and wonderfully made: marvellous are thy works; and that my soul knoweth right well. My substance was not hid from thee, when I was made in secret, and curiously wrought in the lowest parts of the earth. Thine eyes did see my substance, yet being unperfect; and in thy book all my members were written, which in continuance were fashioned, when as yet there was none of them.
Because that which may be known of God is manifest in them; for God hath shewed it unto them. For the invisible things of him from the creation of the world are clearly seen, being understood by the things that are made, even his eternal power and Godhead; so that they are without excuse:
And the LORD God formed man of the dust of the ground, and breathed into his nostrils the breath of life; and man became a living soul.
Sources
- 1.Evolution — Wikipedia (opens in a new tab)
en.wikipedia.orgWikipedia (CC BY-SA 4.0)
- 2.The big issues (opens in a new tab)
evolution.berkeley.edu
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- 8.Scientists Seek to Update Evolution (opens in a new tab)
quantamagazine.org
- 9.Evolution Controversy: A Phenomenon Prompted by the ... (opens in a new tab)
pmc.ncbi.nlm.nih.gov
- 10.An introduction to evolution (opens in a new tab)
evolution.berkeley.edu
- 11.
- 12.Science and evolution - PMC - NIH (opens in a new tab)
pmc.ncbi.nlm.nih.gov
- 13.Evolution (Stanford Encyclopedia of Philosophy) (opens in a new tab)
plato.stanford.edu
- 14.Evolution Resources (opens in a new tab)
nationalacademies.org
- 15.Conceptual framework (opens in a new tab)
evolution.berkeley.edu
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- 18.
Truth Ledger
Every checkable claim in the draft, checked by GPT-6.1 Sol and Grok 4.7. A claim is stated as fact only when both checkers confirm it from the cited sources; a split verdict is published with attribution, and a claim neither can confirm is cut.
Showing 45 claims.
- Verified
Evolution is the change in the heritable characteristics of biological populations over successive generations.
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Source [1] gives this exact definition of evolution. / Source [1] uses this exact definition of evolution.
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Understanding Evolution defines biological evolution as descent with inherited modification.
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Source [10] explicitly defines biological evolution as descent with inherited modification. / Source [10] defines biological evolution as descent with inherited modification.
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Darwin and Wallace independently conceived the theory of evolution by natural selection in the mid-19th century.
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Source [1] identifies Darwin and Wallace as independent originators in the mid-19th century. / Source [1] states Darwin and Wallace independently conceived natural selection in the mid-19th century.
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On the Origin of Species was first published in 1859.
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Source [17] dates the first edition to 1859. / Source [17] says the first edition was published in 1859.
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Britannica describes the theory of evolution as one of the fundamental keystones of modern biological theory.
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Source [11] uses this exact characterization. / Source [11] calls evolution a fundamental keystone of modern biological theory.
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Mutations are the ultimate source of genetic variation in all organisms.
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Source [1] explicitly calls mutations the ultimate source of genetic variation in all organisms. / Source [1] states mutations are the ultimate source of genetic variation in all organisms.
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About half of mutations in coding regions of protein-coding genes are deleterious and the other half neutral, with a small percentage beneficial.
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Source [1] states these approximate proportions for mutations in protein-coding regions. / Source [1] says about half are deleterious, half neutral, and a small percentage beneficial.
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Fitness is indicated by the proportion of subsequent generations that carry an organism's genes, not the total number of offspring.
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Source [1] explicitly distinguishes fitness from total offspring number and describes contribution to subsequent generations. / Source [1] distinguishes fitness from offspring count and ties it to gene representation.
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Fixation of a neutral allele by genetic drift is more rapid in smaller populations.
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Source [1] states that neutral-allele fixation by drift is faster in smaller populations, emphasizing effective population size. / Source [1] says neutral-allele fixation by drift is faster in smaller populations.
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Horizontal gene transfer is most common among bacteria and contributes to the spread of antibiotic resistance.
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Source [1] explicitly describes bacterial prevalence and the contribution to antibiotic resistance. / Source [1] says horizontal gene transfer is most common in bacteria and spreads resistance.
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Recombination and reassortment do not alter allele frequencies but produce new combinations of alleles.
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Source [1] states that recombination and reassortment change allele combinations rather than allele frequencies. / Source [1] says recombination and reassortment reshuffle alleles without changing frequencies.
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E. coli evolved the ability to use citric acid as a nutrient in a long-term laboratory experiment.
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Source [1] explicitly gives this long-term experimental example. / Source [1] cites E. coli evolving citrate use in a long-term lab experiment.
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Speciation has been observed under laboratory conditions and in nature.
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Source [1] states that speciation has been observed repeatedly in laboratories and nature. / Source [1] says speciation has been observed in laboratories and in nature.
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Ernst Mayr defined the Biological Species Concept in 1942.
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Source [1] attributes the Biological Species Concept definition to Ernst Mayr in 1942. / Source [1] attributes the 1942 Biological Species Concept definition to Ernst Mayr.
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More than 99% of all species that have lived on Earth are estimated to be extinct.
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Source [1] explicitly reports the estimate of more than 99%. / Source [1] estimates more than 99% of all species that ever lived are extinct.
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The Permian–Triassic extinction event drove approximately 96% of marine species to extinction.
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Source [1] gives approximately 96% marine-species extinction for the Permian–Triassic event. / Source [1] says the Permian–Triassic event drove about 96% of marine species extinct.
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Present-day extinction rates are 100–1000 times greater than the background rate.
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Source [1] explicitly gives the 100–1000-fold estimate. / Source [1] states present extinction rates are 100–1000 times the background rate.
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The Earth is about 4.54 billion years old.
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Source [1] gives Earth's age as approximately 4.54 billion years. / Source [1] states Earth is about 4.54 billion years old.
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The earliest undisputed evidence of life on Earth dates from at least 3.5 billion years ago.
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Source [1] explicitly dates the earliest undisputed evidence to at least 3.5 billion years ago. / Source [1] dates the earliest undisputed evidence of life to at least 3.5 billion years ago.
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The last universal common ancestor lived approximately 3.5–3.8 billion years ago.
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Source [1] explicitly provides this approximate age range for LUCA. / Source [1] places LUCA at approximately 3.5–3.8 billion years ago.
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The Great Oxygenation Event occurred 2.4 billion years ago and facilitated the emergence of eukaryotes.
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Source [16] dates the oxygen increase to 2.4 billion years ago and says it facilitated eukaryote emergence. / Source [16] dates the GOE to 2.4 billion years ago and links it to eukaryotes.
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Eukaryotic cells emerged between 1.6 and 2.7 billion years ago.
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Source [1] explicitly gives the 1.6–2.7-billion-year range. / Source [1] says eukaryotic cells emerged between 1.6 and 2.7 billion years ago.
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Multicellular organisms began to appear around 1.7 billion years ago.
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Source [1] dates the appearance of multicellular organisms to around 1.7 billion years ago. / Source [1] says multicellular organisms began appearing around 1.7 billion years ago.
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The Cambrian explosion began approximately 538.8 million years ago and spanned around 10 million years.
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Source [1] explicitly gives approximately 538.8 million years ago and a roughly 10-million-year span. / Source [1] places the Cambrian explosion at about 538.8 million years ago over roughly 10 million years.
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Humans and chimpanzees share 98% of their genomes.
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Source [1] explicitly reports 98% genomic sharing between humans and chimpanzees. / Source [1] says humans and chimpanzees share 98% of their genomes.
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Early hominins in Africa date to around 6 million years ago, and human–gorilla divergence may be as early as 8 million years ago.
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Source [16] gives both approximate dates and qualifies the human–gorilla divergence estimate. / Source [16] gives early African hominins around 6 Ma and possible human–gorilla divergence at 8 Ma.
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The oldest fossils attributed to Homo date to about 2.8–2.75 million years ago in Ethiopia.
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Source [18] explicitly dates Ethiopian Homo remains to approximately 2.8–2.75 million years ago; [16] provides compatible dating. / Source [18] dates Homo to about 2.8–2.75 million years ago in Ethiopia; [16] is consistent.
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Sources differ on the age of modern humans: after 200,000 years ago, around 250,000 years ago, and oldest known remains about 315,000 years ago.
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The three sources explicitly give these differing dates, with [18] referring specifically to the oldest known remains. / Sources give after 200,000 ([16]), about 250,000 ([1]), and about 315,000 years ([18]).
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Estimates of current species number differ, from 10–14 million to 10–30 million, with roughly 1.9 to over 2 million named.
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Sources [1] and [11] provide these respective species estimates and named-species counts. / Source [1] estimates 10–14 million species and 1.9 million named; [11] gives 10–30 million and over 2 million named.
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Anaximander and Empedocles proposed that one type of organism could descend from another.
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Source [1] explicitly attributes early proposals of descent between organism types to Anaximander and Empedocles. / Source [1] attributes descent of one organism type from another to Anaximander and Empedocles.
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Ibn Khaldun wrote in the Muqaddimah in 1377 that humans developed from 'the world of the monkeys'.
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Source [1] explicitly gives the author, work, year, and quoted phrase. / Source [1] says Ibn Khaldun’s 1377 Muqaddimah asserted humans developed from “the world of the monkeys.”
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Lamarck's transmutation theory of 1809 was the first full-fledged evolutionary scheme.
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Source [1] calls Lamarck's 1809 transmutation theory the first full-fledged evolutionary scheme. / Source [1] calls Lamarck’s 1809 transmutation theory the first full-fledged evolutionary scheme.
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Darwin did not use the term 'evolution' in the first edition of On the Origin of Species, referring instead to 'descent with modification'.
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Source [13] explicitly states this distinction; Darwin used “evolved,” but not the noun “evolution,” in the first edition. / Source [13] says Darwin used “descent with modification,” not “evolution,” in the first edition.
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Wallace held that natural selection was insufficient to account for the origin of humans.
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Source [17] explicitly states Wallace's view that human origins required more than natural selection. / Source [17] says Wallace thought natural selection insufficient for the origin of humans.
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Fisher, Haldane and Wright used mathematical arguments in the 1920s and 1930s to integrate genetics with natural selection.
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Source [17] identifies these researchers, decades, mathematical arguments, and their role in integrating genetics with natural selection. / Source [17] says Fisher, Haldane, and Wright mathematically supported selection and integrated genetics in the 1920s–30s.
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Dobzhansky published Genetics and the Origin of Species in 1937.
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Source [17] explicitly gives the title, author, and publication year. / Source [17] says Dobzhansky published Genetics and the Origin of Species in 1937.
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The modern synthesis has guided evolutionary biology for over 50 years.
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Source [8] explicitly states that the modern synthesis had guided evolutionary biology for over 50 years. / Source [8] says the Modern Synthesis has guided evolutionary biology for over 50 years.
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The relative importance of selection and neutral processes is an area of current research.
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Source [1] explicitly identifies the comparative importance of adaptive and non-adaptive forces as ongoing research. / Source [1] identifies the relative importance of selection and neutral processes as current research.
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There is little consensus on how to detect and interpret signatures of selection in human genomes while avoiding confounders.
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Source [3] explicitly states that little consensus exists on detecting and interpreting selection signatures while avoiding confounders. / Source [3] says there is little consensus on detecting and interpreting selection while avoiding confounders.
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The extent to which evolution can be predicted remains debated, and 'complex selection' has been proposed to improve prediction.
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Source [4] explicitly describes the predictability debate and proposes understanding complex selection to improve prediction. / Source [4] says evolutionary predictability remains debated and proposes complex selection to improve it.
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Human evolution lacks a widely accepted unifying explanation, according to a paper in Evolutionary Anthropology.
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Source [6] explicitly states that human evolution lacks a widely accepted unifying explanation. / The Evolutionary Anthropology paper in [6] says human evolution lacks a widely accepted unifying explanation.
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Laland and colleagues argue for an Extended Evolutionary Synthesis, while other biologists say there is little evidence a paradigm shift is warranted.
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Source [8] explicitly describes both Laland's proposal and opposing biologists' skepticism about a paradigm shift. / Source [8] reports Laland’s Extended Evolutionary Synthesis proposal and skeptics citing little evidence for a paradigm shift.
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Viruses, bacteria, fungi and cancers evolve resistance to drugs and immune defences.
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Source [1] explicitly lists viruses, bacteria, fungi, and cancers evolving resistance to immune defenses and pharmaceutical drugs. / Source [1] says viruses, bacteria, fungi, and cancers evolve resistance to drugs and immune defences.
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Artificial evolution became a recognised optimisation method through Ingo Rechenberg's work in the 1960s.
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Source [1] explicitly attributes recognition of artificial evolution as an optimization method to Rechenberg's work in the 1960s. / Source [1] credits Rechenberg’s 1960s work with making artificial evolution a recognised optimisation method.
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Text is available under the Creative Commons Attribution-ShareAlike 4.0 licence. Written by Claude Opus 5.5 from the sources listed and checked claim by claim by GPT-6.1 Sol and Grok 4.7.