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The theory is supported by evidence from fossils, and by the rapid changes that can be seen to occur in microorganisms such as antibiotic-resistant bacteria.
Many species have become extinct in the past and the extinction of species continues to happen. The basic idea behind the theory of evolution is that different species have developed over time from simple life forms.
These simple life forms first developed more than three billion years ago as the Earth is believed to be about 4. This timeline shows key events in the evolution of life forms on Earth, from the first bacteria to the human species.
You can see a more detailed history of life timeline on BBC Nature. Download 'Tree of Life' poster pdf file, 2. This page is best viewed in an up-to-date web browser with style sheets CSS enabled.
Advancements were also made in phylogenetic systematics , mapping the transition of traits into a comparative and testable framework through the publication and use of evolutionary trees.
Since then, the modern synthesis has been further extended to explain biological phenomena across the full and integrative scale of the biological hierarchy , from genes to species.
One extension, known as evolutionary developmental biology and informally called "evo-devo," emphasises how changes between generations evolution acts on patterns of change within individual organisms development.
Evolution in organisms occurs through changes in heritable traits—the inherited characteristics of an organism.
In humans, for example, eye colour is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of their parents.
The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype.
These traits come from the interaction of its genotype with the environment. For example, suntanned skin comes from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children.
However, some people tan more easily than others, due to differences in genotypic variation; a striking example are people with the inherited trait of albinism , who do not tan at all and are very sensitive to sunburn.
Heritable traits are passed from one generation to the next via DNA, a molecule that encodes genetic information.
The sequence of bases along a particular DNA molecule specify the genetic information, in a manner similar to a sequence of letters spelling out a sentence.
Portions of a DNA molecule that specify a single functional unit are called genes; different genes have different sequences of bases.
Within cells, the long strands of DNA form condensed structures called chromosomes. The specific location of a DNA sequence within a chromosome is known as a locus.
If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles.
DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of the organism.
Recent findings have confirmed important examples of heritable changes that cannot be explained by changes to the sequence of nucleotides in the DNA.
These phenomena are classed as epigenetic inheritance systems. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment.
This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Descendants inherit genes plus environmental characteristics generated by the ecological actions of ancestors.
An individual organism's phenotype results from both its genotype and the influence from the environment it has lived in. A substantial part of the phenotypic variation in a population is caused by genotypic variation.
The frequency of one particular allele will become more or less prevalent relative to other forms of that gene. Variation disappears when a new allele reaches the point of fixation —when it either disappears from the population or replaces the ancestral allele entirely.
Natural selection will only cause evolution if there is enough genetic variation in a population. Before the discovery of Mendelian genetics, one common hypothesis was blending inheritance.
But with blending inheritance, genetic variance would be rapidly lost, making evolution by natural selection implausible.
The Hardy—Weinberg principle provides the solution to how variation is maintained in a population with Mendelian inheritance.
The frequencies of alleles variations in a gene will remain constant in the absence of selection, mutation, migration and genetic drift.
Variation comes from mutations in the genome, reshuffling of genes through sexual reproduction and migration between populations gene flow.
Despite the constant introduction of new variation through mutation and gene flow, most of the genome of a species is identical in all individuals of that species.
Mutations are changes in the DNA sequence of a cell's genome. When mutations occur, they may alter the product of a gene , or prevent the gene from functioning, or have no effect.
Mutations can involve large sections of a chromosome becoming duplicated usually by genetic recombination , which can introduce extra copies of a gene into a genome.
New genes can be generated from an ancestral gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new function while the other copy continues to perform its original function.
The generation of new genes can also involve small parts of several genes being duplicated, with these fragments then recombining to form new combinations with new functions.
In asexual organisms, genes are inherited together, or linked , as they cannot mix with genes of other organisms during reproduction.
In contrast, the offspring of sexual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment.
In a related process called homologous recombination , sexual organisms exchange DNA between two matching chromosomes.
The two-fold cost of sex was first described by John Maynard Smith. This cost does not apply to hermaphroditic species, like most plants and many invertebrates.
The Red Queen hypothesis has been used to explain the significance of sexual reproduction as a means to enable continual evolution and adaptation in response to coevolution with other species in an ever-changing environment.
Gene flow is the exchange of genes between populations and between species. Gene flow can be caused by the movement of individuals between separate populations of organisms, as might be caused by the movement of mice between inland and coastal populations, or the movement of pollen between heavy-metal-tolerant and heavy-metal-sensitive populations of grasses.
Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer.
Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria.
Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and bacteria, during the acquisition of chloroplasts and mitochondria.
It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria and archaea. From a neo-Darwinian perspective, evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms,  for example, the allele for black colour in a population of moths becoming more common.
Mechanisms that can lead to changes in allele frequencies include natural selection, genetic drift, genetic hitchhiking, mutation and gene flow.
Evolution by means of natural selection is the process by which traits that enhance survival and reproduction become more common in successive generations of a population.
It has often been called a "self-evident" mechanism because it necessarily follows from three simple facts: More offspring are produced than can possibly survive, and these conditions produce competition between organisms for survival and reproduction.
Consequently, organisms with traits that give them an advantage over their competitors are more likely to pass on their traits to the next generation than those with traits that do not confer an advantage.
The central concept of natural selection is the evolutionary fitness of an organism. If an allele increases fitness more than the other alleles of that gene, then with each generation this allele will become more common within the population.
These traits are said to be "selected for. Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele becoming rarer—they are "selected against.
Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types.
The first is directional selection , which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. This would be when either short or tall organisms had an advantage, but not those of medium height.
Finally, in stabilising selection there is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity.
A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates.
Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises the survival of individual males.
Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive.
Eugene Odum , a founder of ecology , defined an ecosystem as: These relationships involve the life history of the organism, its position in the food chain and its geographic range.
This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection.
Natural selection can act at different levels of organisation , such as genes, cells, individual organisms, groups of organisms and species. In addition to being a major source of variation, mutation may also function as a mechanism of evolution when there are different probabilities at the molecular level for different mutations to occur, a process known as mutation bias.
Mutation bias effects are superimposed on other processes. If selection would favour either one out of two mutations, but there is no extra advantage to having both, then the mutation that occurs the most frequently is the one that is most likely to become fixed in a population.
Most loss of function mutations are selected against. But when selection is weak, mutation bias towards loss of function can affect evolution.
Loss of sporulation ability in Bacillus subtilis during laboratory evolution appears to have been caused by mutation bias, rather than natural selection against the cost of maintaining sporulation ability.
In parasitic organisms, mutation bias leads to selection pressures as seen in Ehrlichia. Mutations are biased towards antigenic variants in outer-membrane proteins.
Genetic drift is the random fluctuations of allele frequencies within a population from one generation to the next. Genetic drift may therefore eliminate some alleles from a population due to chance alone.
Even in the absence of selective forces, genetic drift can cause two separate populations that began with the same genetic structure to drift apart into two divergent populations with different sets of alleles.
The neutral theory of molecular evolution proposed that most evolutionary changes are the result of the fixation of neutral mutations by genetic drift.
The time for a neutral allele to become fixed by genetic drift depends on population size, with fixation occurring more rapidly in smaller populations.
It is usually difficult to measure the relative importance of selection and neutral processes, including drift. Recombination allows alleles on the same strand of DNA to become separated.
However, the rate of recombination is low approximately two events per chromosome per generation. As a result, genes close together on a chromosome may not always be shuffled away from each other and genes that are close together tend to be inherited together, a phenomenon known as linkage.
A set of alleles that is usually inherited in a group is called a haplotype. This can be important when one allele in a particular haplotype is strongly beneficial: Gene flow involves the exchange of genes between populations and between species.
Due to the complexity of organisms, any two completely isolated populations will eventually evolve genetic incompatibilities through neutral processes, as in the Bateson-Dobzhansky-Muller model , even if both populations remain essentially identical in terms of their adaptation to the environment.
If genetic differentiation between populations develops, gene flow between populations can introduce traits or alleles which are disadvantageous in the local population and this may lead to organisms within these populations evolving mechanisms that prevent mating with genetically distant populations, eventually resulting in the appearance of new species.
Thus, exchange of genetic information between individuals is fundamentally important for the development of the Biological Species Concept BSC.
During the development of the modern synthesis, Sewall Wright developed his shifting balance theory , which regarded gene flow between partially isolated populations as an important aspect of adaptive evolution.
Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptations that are the outcome of natural selection.
These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by cooperating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis.
In the longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed.
These outcomes of evolution are distinguished based on time scale as macroevolution versus microevolution.
Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction; whereas microevolution refers to smaller evolutionary changes within a species or population, in particular shifts in allele frequency and adaptation.
For instance, a large amount of variation among individuals allows a species to rapidly adapt to new habitats , lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated.
In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction.
A common misconception is that evolution has goals, long-term plans, or an innate tendency for "progress", as expressed in beliefs such as orthogenesis and evolutionism; realistically however, evolution has no long-term goal and does not necessarily produce greater complexity.
Adaptation is the process that makes organisms better suited to their habitat. For example, the adaptation of horses ' teeth to the grinding of grass.
By using the term adaptation for the evolutionary process and adaptive trait for the product the bodily part or function , the two senses of the word may be distinguished.
Adaptations are produced by natural selection. Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature.
An example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of the cell.
Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organisation may have different functions in related organisms.
This is the result of a single ancestral structure being adapted to function in different ways. The bones within bat wings, for example, are very similar to those in mice feet and primate hands, due to the descent of all these structures from a common mammalian ancestor.
During evolution, some structures may lose their original function and become vestigial structures. Examples include pseudogenes ,  the non-functional remains of eyes in blind cave-dwelling fish,  wings in flightless birds,  the presence of hip bones in whales and snakes,  and sexual traits in organisms that reproduce via asexual reproduction.
However, many traits that appear to be simple adaptations are in fact exaptations: However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation.
An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations and exaptations.
Interactions between organisms can produce both conflict and cooperation. When the interaction is between pairs of species, such as a pathogen and a host , or a predator and its prey, these species can develop matched sets of adaptations.
Here, the evolution of one species causes adaptations in a second species. These changes in the second species then, in turn, cause new adaptations in the first species.
This cycle of selection and response is called coevolution. In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake.
Not all co-evolved interactions between species involve conflict. For instance, an extreme cooperation exists between plants and the mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil.
Here, the fungi actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals that suppress the plant immune system.
Coalitions between organisms of the same species have also evolved. An extreme case is the eusociality found in social insects, such as bees , termites and ants , where sterile insects feed and guard the small number of organisms in a colony that are able to reproduce.
On an even smaller scale, the somatic cells that make up the body of an animal limit their reproduction so they can maintain a stable organism, which then supports a small number of the animal's germ cells to produce offspring.
Here, somatic cells respond to specific signals that instruct them whether to grow, remain as they are, or die. If cells ignore these signals and multiply inappropriately, their uncontrolled growth causes cancer.
Such cooperation within species may have evolved through the process of kin selection , which is where one organism acts to help raise a relative's offspring.
Speciation is the process where a species diverges into two or more descendant species. There are multiple ways to define the concept of "species.
Despite the diversity of various species concepts, these various concepts can be placed into one of three broad philosophical approaches: Defined by evolutionary biologist Ernst Mayr in , the BSC states that "species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups.
Barriers to reproduction between two diverging sexual populations are required for the populations to become new species. Gene flow may slow this process by spreading the new genetic variants also to the other populations.
Depending on how far two species have diverged since their most recent common ancestor , it may still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.
In this case, closely related species may regularly interbreed, but hybrids will be selected against and the species will remain distinct.
However, viable hybrids are occasionally formed and these new species can either have properties intermediate between their parent species, or possess a totally new phenotype.
Speciation has been observed multiple times under both controlled laboratory conditions see laboratory experiments of speciation and in nature.
There are four primary geographic modes of speciation. The most common in animals is allopatric speciation , which occurs in populations initially isolated geographically, such as by habitat fragmentation or migration.
Selection under these conditions can produce very rapid changes in the appearance and behaviour of organisms. The second mode of speciation is peripatric speciation , which occurs when small populations of organisms become isolated in a new environment.
This differs from allopatric speciation in that the isolated populations are numerically much smaller than the parental population. Here, the founder effect causes rapid speciation after an increase in inbreeding increases selection on homozygotes, leading to rapid genetic change.
The third mode is parapatric speciation. This is similar to peripatric speciation in that a small population enters a new habitat, but differs in that there is no physical separation between these two populations.
Instead, speciation results from the evolution of mechanisms that reduce gene flow between the two populations. One example is the grass Anthoxanthum odoratum , which can undergo parapatric speciation in response to localised metal pollution from mines.
Selection against interbreeding with the metal-sensitive parental population produced a gradual change in the flowering time of the metal-resistant plants, which eventually produced complete reproductive isolation.
Selection against hybrids between the two populations may cause reinforcement , which is the evolution of traits that promote mating within a species, as well as character displacement , which is when two species become more distinct in appearance.
Finally, in sympatric speciation species diverge without geographic isolation or changes in habitat. This form is rare since even a small amount of gene flow may remove genetic differences between parts of a population.
One type of sympatric speciation involves crossbreeding of two related species to produce a new hybrid species. This is not common in animals as animal hybrids are usually sterile.
This is because during meiosis the homologous chromosomes from each parent are from different species and cannot successfully pair. However, it is more common in plants because plants often double their number of chromosomes, to form polyploids.
Speciation events are important in the theory of punctuated equilibrium , which accounts for the pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis, where species remain relatively unchanged.
As a result, the periods of stasis in the fossil record correspond to the parental population and the organisms undergoing speciation and rapid evolution are found in small populations or geographically restricted habitats and therefore rarely being preserved as fossils.
Extinction is the disappearance of an entire species. Extinction is not an unusual event, as species regularly appear through speciation and disappear through extinction.
The role of extinction in evolution is not very well understood and may depend on which type of extinction is considered. The Earth is about 4.
Microbial mat fossils have been found in 3. More than 99 percent of all species, amounting to over five billion species,  that ever lived on Earth are estimated to be extinct.
Highly energetic chemistry is thought to have produced a self-replicating molecule around 4 billion years ago, and half a billion years later the last common ancestor of all life existed.
All organisms on Earth are descended from a common ancestor or ancestral gene pool. First, they have geographic distributions that cannot be explained by local adaptation.
Second, the diversity of life is not a set of completely unique organisms, but organisms that share morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits and finally, that organisms can be classified using these similarities into a hierarchy of nested groups—similar to a family tree.
Past species have also left records of their evolutionary history. Fossils, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record.
However, this approach is most successful for organisms that had hard body parts, such as shells, bones or teeth.
Further, as prokaryotes such as bacteria and archaea share a limited set of common morphologies, their fossils do not provide information on their ancestry.
More recently, evidence for common descent has come from the study of biochemical similarities between organisms. For example, all living cells use the same basic set of nucleotides and amino acids.
Prokaryotes inhabited the Earth from approximately 3—4 billion years ago. The next major change in cell structure came when bacteria were engulfed by eukaryotic cells, in a cooperative association called endosymbiosis.
The history of life was that of the unicellular eukaryotes, prokaryotes and archaea until about million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.
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.
About million years ago, plants and fungi colonised the land and were soon followed by arthropods and other animals.
Concepts and models used in evolutionary biology, such as natural selection, have many applications. Artificial selection is the intentional selection of traits in a population of organisms.
This has been used for thousands of years in the domestication of plants and animals. Proteins with valuable properties have evolved by repeated rounds of mutation and selection for example modified enzymes and new antibodies in a process called directed evolution.
Understanding the changes that have occurred during an organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human genetic disorders.
Breeding together different populations of this blind fish produced some offspring with functional eyes, since different mutations had occurred in the isolated populations that had evolved in different caves.
Evolutionary theory has many applications in medicine. Many human diseases are not static phenomena, but capable of evolution.
Viruses, bacteria, fungi and cancers evolve to be resistant to host immune defences , as well as pharmaceutical drugs. It is possible that we are facing the end of the effective life of most of available antibiotics  and predicting the evolution and evolvability  of our pathogens and devising strategies to slow or circumvent it is requiring deeper knowledge of the complex forces driving evolution at the molecular level.
In computer science , simulations of evolution using evolutionary algorithms and artificial life started in the s and were extended with simulation of artificial selection.
He used evolution strategies to solve complex engineering problems. In the 19th century, particularly after the publication of On the Origin of Species in , the idea that life had evolved was an active source of academic debate centred on the philosophical, social and religious implications of evolution.
Today, the modern evolutionary synthesis is accepted by a vast majority of scientists. While various religions and denominations have reconciled their beliefs with evolution through concepts such as theistic evolution , there are creationists who believe that evolution is contradicted by the creation myths found in their religions and who raise various objections to evolution.
The teaching of evolution in American secondary school biology classes was uncommon in most of the first half of the 20th century. The Scopes Trial decision of caused the subject to become very rare in American secondary biology textbooks for a generation, but it was gradually re-introduced later and became legally protected with the Epperson v.
Since then, the competing religious belief of creationism was legally disallowed in secondary school curricula in various decisions in the s and s, but it returned in pseudoscientific form as intelligent design ID , to be excluded once again in the Kitzmiller v.
Dover Area School District case. From Wikipedia, the free encyclopedia. This article is about evolution in biology. For related articles, see Outline of evolution.
For other uses, see Evolution disambiguation. Change in the heritable characteristics of biological populations over successive generations.
For a more accessible and less technical introduction to this topic, see Introduction to evolution. Introduction to evolution Evidence of evolution Common descent Evidence of common descent.
History of evolutionary theory. Applications of evolution Biosocial criminology Ecological genetics Evolutionary aesthetics Evolutionary anthropology Evolutionary computation Evolutionary ecology Evolutionary economics Evolutionary epistemology Evolutionary ethics Evolutionary game theory Evolutionary linguistics Evolutionary medicine Evolutionary neuroscience Evolutionary physiology Evolutionary psychology Experimental evolution Phylogenetics Paleontology Selective breeding Speciation experiments Sociobiology Systematics Universal Darwinism.
Evolution as fact and theory Social effects Creation—evolution controversy Objections to evolution Level of support. History of evolutionary thought.
Modern synthesis 20th century. Introduction to genetics , Genetics , Heredity , and Reaction norm. Black morph in peppered moth evolution.
Genetic diversity and Population genetics. Sexual reproduction , Genetic recombination , and Evolution of sexual reproduction. Genetic drift and Effective population size.
Genetic hitchhiking , Hill—Robertson effect , and Selective sweep. Gene flow , Hybrid biology , and Horizontal gene transfer. Assortative mating and Panmixia.
Human timeline and Nature timeline. Evolutionary history of life. Timeline of evolutionary history of life. Common descent and Evidence of common descent.
Evolutionary history of life and Timeline of evolutionary history of life. Applications of evolution , Selective breeding , and Evolutionary computation.
Social effects of evolutionary theory , Oxford evolution debate , Creation—evolution controversy , Objections to evolution , and Evolution in fiction.
Argument from poor design Biocultural evolution Biological classification Evidence of common descent Evolutionary anthropology Evolutionary ecology Evolutionary epistemology Evolutionary neuroscience Evolution of biological complexity Evolution of plants Project Steve Timeline of the evolutionary history of life Universal Darwinism.
National Academies of Sciences, Engineering, and Medicine. Archived from the original on Evolutionary processes are generally thought of as processes by which these changes occur.
Four such processes are widely recognized: The latter two generate variation; the first two sort it. On the Origin of Species 2nd ed. Annual Review of Ecology and Systematics.
Archived PDF from the original on Archived from the original PDF on William ; Kudryavtsev, Anatoliy B. Meet your microbial mom". Archived from the original on June 29, New Brunswick, New Jersey: Leeds International Classical Studies.
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Philosophical Transactions of the Royal Society B: Some of the values in table 1 on p. The errors do not affect the conclusions drawn in the paper.
The corrected table is reproduced below. Journal of Molecular Evolution. Protein Families, Pseudogenes and Proteome Evolution".